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Conclusions from Epizoanthus papillosus Benthic surveys have shown that Epizoanthus papillosus is widespread and quite abundant in the seas around the British Isles, apart from the Irish Sea, English Channel and Southern Bight. It most commonly constructs carcinoecia in association with Anapagurus laevis but small, free-living colonies also occur. The latter facilitated cnidom comparisons between equivalent-sized polyps in carcinoecia and free-living colonies. As has been previously demonstrated in other zoanthids (Ryland, et al., 2004), the capsule size in some nematocyst types is correlated with polyp size (and, interestingly, may be correlated with umbrella size in Cyanea (��stman & Hydman, 1997)). This has great relevance to the use of capsule size in taxonomy. Major differences were found between the cnidoms of carcinoecia and free-living colonies, including presence or absence (holotrichs in tentacles of freeliving colonies only), abundance (basitrichs being much more abundant in free-living colonies), and capsule size (both holotrichs and p -mastigophores being larger in free-living colonies). The differences seem probably related to method of feeding, with carcinoecia polyps less dependent on captured live prey and able to benefit from the activities of the hermit crab, but may also be to protect the pagurid itself. This demonstrated paucity of nematocyst types and abundance has a significant, negative impact on the use of cnidom characters in the taxonomy of carcinoecium-forming species of Epizoanthus., Published as part of Ryland, John S. & Ward, Helen, 2016, Carcinoecium-forming Epizoanthus [Hexacorallia: Zoantharia] and the biology of E. papillosus in the eastern Atlantic, with special reference to the cnidom, pp. 489-514 in Zootaxa 4088 (4) on page 510, DOI: 10.11646/zootaxa.4088.4.2, http://zenodo.org/record/263647

Ryland, John S., Ward, Helen (2016): Carcinoecium-forming Epizoanthus [Hexacorallia: Zoantharia] and the biology of E. papillosus in the eastern Atlantic, with special reference to the cnidom. Zootaxa 4088 (4): 489-514, DOI: http://doi.org/10.11646/zootaxa.4088.4.2

Ryland, John S., Holt, Rohan, Loxton, Jennifer, Spencer Jones, Mary E., Porter, Joanne S. (2014): First occurrence of the non-native bryozoan Schizoporella japonica Ortmann (1890) in Western Europe. Zootaxa 3780 (3): 481-502, DOI: http://dx.doi.org/10.11646/zootaxa.3780.3.3

Schizoporella on the Pacific coast of North America Schizoporella unicornis, a Recent bryozoan originally described in a work on the Coralline Crag by Johnston (in Wood 1844; see Tompsett et al. 2009 for details) is a well-known European species (Johnston 1847; Hincks 1880; Ryland 1965; Hayward & Ryland 1995, 1999; but not Marcus 1940 (= S. errata, absent from northern Europe)) that is unknown on the Pacific coast of North America. Because of undue reliance on European literature, generally inappropriate for the Pacific coast (e.g. Ryland & Porter 2012), the characteristics of this species (Hayward & Ryland 1995, 1999; Tompsett et al. 2009), especially the marginally fluted but virtually non-porous ovicell, were missed by authors or deliberately ignored (e.g. Ross & McCain 1976) and the name unicornis has been incorrectly applied to at least two quite different species. Osburn (1952) —before S. japonica had been recognized in the northwest—included only one nominate species that would now be included in the genus Schizoporella, using the name S. unicornis, from various localities in California. Osburn’s account is now known to have been based on a mixture of S. japonica and S. errata (Powell 1970) and S. pseudoerrata (described by Soule et al. 1995). The current distribution of S. errata is unclear since the species is not discussed by Soule et al. (1995, 2007), though it is certainly common in San Francisco Bay (Zabin et al. 2010). The differences between S. unicornis and S. errata are in fact numerous and considerable (this paper and Ryland 1965; Hastings 1968; Hayward & Ryland 1999; Hayward & McKinney 2002; Tompsett et al. 2009). Whether S. errata should be regarded as a single species, a complex, or several species is another issue (Winston & Hayward 2012), which cannot be resolved here. While Schizoporella errata, being a well-known fouling species, seems likely to have been introduced to the Pacific coast well before its first recorded occurrences (as S. unicornis, by Osburn 1952), S. japonica is most certainly a recent alien. However, Powell (1970) established that it (as S. unicornis), as opposed to S. errata, was present in Newport Bay, Los Angeles, as long ago as 1938 (material collected by G. E. MacGinitie); he assumed that it had arrived with Pacific oysters, Crassostrea gigas, which had been imported from Japan (first to Morro Bay) since 1932. This is the earliest record for this species on the west coast of North America. Powell (1970) referred to additional material in USNM from Newport, collected 1943, and Morro Bay, collected 1968. Powell (1970) himself found it (still using the name S. unicornis) from the Strait of Georgia, Canada. It had not been found earlier by O’Donoghue & O’Donoghue (1923, 1925, 1926) but was found by Powell during 1966–69 from several stations in the San Juan Islands, on Vancouver Island, and from as far north as Pendrell Sound (50° N). As for California, he attributed its arrival to the extensive importation of Pacific oysters from Japan in the period 1926– 1935. Its recorded range was extended to further localities in Washington State by Ross & McCain (1976), who conducted a thorough study of zooidal shape [the variability noted earlier that arises from the growth pattern of circular colonies, a topic also investigated in this species by Thorpe & Ryland (1987)]. It was also collected in San Francisco Bay during 1977 (NHMUK 1978.1.4.2). Whereas S. errata, as a warm-water species, is likely to be commonest south of San Francisco, the converse is true for S. japonica. It has spread northwards through Canada to southern Alaska (Dick et al. 2005) but, as already noted, it extends southwards beyond San Francisco to Morro Bay (#14) and (historically at least) to the Los Angeles area. Thus it now appears that three distinct species of Schizoporella occur in central California (i.e. the Monterey Bay area), S. errata, S. japonica, and S. pseudoerrata, the first two, at least, being introductions. Sorte et al. (2010) listed an unidentified Schizoporella from Bodega Harbor, most likely S. japonica (see #21) but possibly S. errata, and Zabin et al. (2012) recorded unidentified Schizoporella from two sites at Santa Cruz. Schizoporella pseudoerrata at present has a very localized confirmed distribution (Soule et al. 1995) although it has been listed as present at two sites in the northern part of San Francisco Bay—Richmond Marina (Blum et al. 2007) and Tiburon (Crooks et al. 2011). It is possible that these records are based on misidentifications and might be either of the other two species. With all three species now recognized and described, it should be possible to correctly identify Schizoporella specimens from the Pacific coast. The characteristics of the three Californian species are summarised in Table 7 (particularly note the distinctive condyles, observation of which requires the preparation of specimens with a hypochlorite bleach such as Clorox: see Material and Methods above), which should be used in conjunction with Figures 2–4 and 10. In addition to morphometric methods, genetic techniques have recently been applied in two non-native fouling bryozoan species-groups. In Bugula neritina (Linnaeus), three biological species were identified. One of these, haplotype S, was globally distributed (Fehlauer-Ale et al. 2013). In Watersipora subtorquata, three clades were identified by Mackie et al. (2012). These studies provide evidence for cryptic speciation in the fouling community. Genetic studies on Schizoporella japonica are currently underway to investigate the potential for cryptic species in this taxon., Published as part of Ryland, John S., Holt, Rohan, Loxton, Jennifer, Spencer Jones, Mary E. & Porter, Joanne S., 2014, First occurrence of the non-native bryozoan Schizoporella japonica Ortmann (1890) in Western Europe, pp. 481-502 in Zootaxa 3780 (3) on pages 495-496, DOI: 10.11646/zootaxa.3780.3.3, http://zenodo.org/record/4910530, {"references":["Wood, S. V. (1844) Descriptive catalogue of the zoophytes of the Crag. Annals and Magazine of Natural History, 13, 10 - 21. http: // dx. doi. org / 10.1080 / 03745484409442561","Tompsett, S., Porter, J. S. & Taylor, P. D. (2009) Taxonomy of the fouling cheilostome bryozoans Schizoporella unicornis (Johnston) and S. errata (Waters). Journal of Natural History, 43, 2227 - 2243. http: // dx. doi. org / 10.1080 / 00222930903090140","Johnston, G. (1847) A History of the British Zoophytes. J. van Voorst, London, 488 pp., 74 pls.","Hincks, T. (1880) A History of the British Marine Polyzoa. van Voorst, London, 772 pp.","Ryland, J. S. (1965) Polyzoa. Catalogue of Main Marine Fouling Organisms, 2, 1 - 83.","Hayward, P. J. & Ryland, J. S. (1995) The British species of Schizoporella (Bryozoa: Cheilostomatida). Journal of Zoology (London), 237, 37 - 47. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1995. tb 02744. x","Hayward, P. J. & Ryland, J. S. (1999) Cheilostomatous Bryozoa. Part 2, Hippothooidea - Celleporoidea. Synopses of the British Fauna, n. s., 14, 1 - 416.","Marcus, E. (1940) Mosdyr (Bryozoer eller Polyzoer). Danmarks Fauna, 46, 1 - 102.","Ryland, J. S. & Porter, J. S. (2012) Species of Alcyonidium (Ctenostomatida) from the Pacific coast of North America: a preliminary account. In: Ernst, A., Schaefer, P. & Scholz, J. (Eds.), Bryozoan studies 2010. Springer, Heidelberg, pp. 289 - 302.","Ross, J. & McCain, K. (1976) Schizoporella unicornis (Ectoprocta) in coastal waters of northwestern United States and Canada. Northwest Science, 50, 160 - 171.","Osburn, R. C. (1952) Bryozoa of the Pacific coast of America. Part 2, Cheilostomata-Ascophora. Allan Hancock Pacific Expeditions, 14 (2), 1 - 611.","Soule, D. F., Soule, J. D. & Chaney, H. W. (1995) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and western Santa Barbara Channel: The Bryozoa. Irene McCulloch Foundation Monograph Series, 2, 1 - 344.","Soule, D. F., Soule, J. D., Morris, P. A. & Chaney, H. W. (2007) Bryozoa. In: Carlton, J. T. (Ed.) The Light and Smith Manual: intertidal invertebrates from central California to Oregon. University of California Press, Berkeley and Los Angeles, pp. 866 - 904.","Zabin, C., Obernolte, R., Mackie, J., Gentry, J., Harris, L. & Geller, J. (2010) A non-native bryozoan creates novel substrate on the mudflats in San Francisco Bay. Marine Ecology Progress Series, 412, 129 - 139. http: // dx. doi. org / 10.3354 / meps 08664","Hastings, A. B. (1968) Some type and other specimens of species involved in the problem of Stylopoma Levinsen (Polyzoa). Bulletin of the British Museum (Natural History), Zoology, 16, 355 - 364.","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. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2002) 270 2.0. co; 2","Winston, J. E. & Hayward, P. J. (2012) The marine bryozoans of the northeast coast of the United States: Maine to Virginia. Memoirs of the Virginia Museum of Natural History, 11, 1 - 180.","O'Donoghue, C. & O'Donoghue, E. (1923) A preliminary list of Bryozoa (Polyzoa) from the Vancouver Island region. Contributions to Canadian Biology and Fisheries, n. s., 1, 143 - 201. http: // dx. doi. org / 10.1139 / f 22 - 010","O'Donoghue, C. H. & O'Donoghue, E. (1925) Notes on certain Bryozoa in the collection of the University of Washington. Washington University Puget Sound Biological Station Publications, 5, 15 - 23.","O'Donoghue, C. H. & O'Donoghue, E. (1926) A second list of the Bryozoa (Polyzoa) from the Vancouver Island region. Contributions to Canadian Biology and Fisheries, N. S., 3, 47 - 131. http: // dx. doi. org / 10.1139 / f 26 - 003","Thorpe, J. P. & Ryland, J. S. (1987) Some theoretical limitations on the arrangement of zooids in encrusting Bryozoa. In: Ross, J. R. P. (Ed.), Bryozoa: present and past. Western Washington University, Bellingham, pp. 277 - 283.","Dick, M. H., Grischenko, A. V. & Mawatari, F. S. (2005) Intertidal Bryozoa (Cheilostomata) of Ketchikan, Alaska. Journal of Natural History, 39, 3687 - 3784.","Sorte, C., Fuller, A. & Bracken, E. (2010) Impacts of a simulated heatwave on composition of a marine community. Oikos, 119, 1901 - 1918.","Zabin, C., Danner, E., Baumgartner, E., Spafford, D., Miller, K. & Pearse, J. (2012) A comparison of intertidal species richness and composition between Central California and Oahu, Hawaii. Marine Ecology, 1 - 26. http: // dx. doi. org / 10.1111 / maec. 12007","Blum, J. C., Chang, A. L., Liljesthrom, M., Schenk, M. E., Steinberg, M. K. & Ruiz, G. M. (2007) The non-native solitary ascidian Ciona intestinalis (L.) depresses species richness. Journal of Experimental Marine Biology and Ecology, 342, 5 - 14. http: // dx. doi. org / 10.1016 / j. jembe. 2006.10.010","Crooks, J. A., Chang, A. L. & Ruiz, G. M. (2011) Aquatic pollution increases the relative success of invasive species. Biological Invasions, 13, 165 - 176. http: // dx. doi. org / 10.1007 / s 10530 - 010 - 9799 - 3","Fehlauer-Ale, K. H., Mackie, J. A., Lim-Fing, G. E., Ale, E., Pie, M. R. & Waeschenbach, A (2013) Cryptic species in the cosmopolitan Bugula neritina complex (Bryozoa, Cheilostomata). Zoologica Scripta, 43 (2), 193 - 205. http: // dx. doi. org / 10.1111 / zsc. 12042","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"]}

Schizoporella japonica Ortmann, 1890 (Figures 2–5) Schizoporella unicornis var. japonica Ortmann, 1890: 49, pl. 3, fig. 35. Schizoporella unicornis: Okada 1929: 20, fig. 7; Powell 1970: 1849, figs 2–3; Ross & McCain 1976: 164, figs 1–6; Kubota & Mawatari 1985: 201, fig. 3A–E; Thorpe & Ryland 1987: 281 (Friday Harbor, in part); Osburn 1952: 317, pl. 37, figs 1–2 (part; some = S. pseudoerrata); Soule et al. 1995: 204, pl. 75A–F. Schizoporella japonica: Dick et al. 2005: 3742, figs 15–16; Grischenko et al. 2007: 1115. Material examined. See Appendix 1. Holotype. Strasbourg Museum, MZS Bry001 as S. unicornis var. japonica; type Locality Sagami Bay, Japan, collected Dr L. Döderlein, 1880–1881. A photograph of the supporting stone and holotype colony is shown in Fig. 4E. Description of British material. Colonies at first more or less circular (Fig. 4F), rapidly becoming extensive, pale whitish-pink to vivid orange-red (Figs 2, 4; British material between Munsell 2.5YR6/14 and 10.0R6/10 (see Kelly & Judd 1955)); mainly unilaminar but frequently with raised edges or displaying slightly elevated lobes (Fig. 2B, C). Zooids generally in obviously linear series, quincuncial away from bifurcations; rectangular; conspicuously longer (often twice as long) than broad (0.5–0.7 × 0.25–0.35 mm; Table 1); length:width proportions (1.7–2.5:1) varying according to distance from a bifurcation (Figs 2D–E, 3A, E); the dividing line between series distinct, slightly depressed; frontal shield with marginal areolae and regularly distributed pseudopores, except sometimes (in Holyhead material) for an incipient suboral umbo; the distolateral pair of areolae somewhat larger. Frontal pseudopores very numerous (c. 600 mm -2, range ~ 400−800 mm -2). Orifice shallower than wide (0.8–0.9:1), though variable within a colony (110–140 × 150–175 µm); the sinus shallow and broad (0.3–0.4:1; 20–35 × 80–120 µm), with sinuous margins, delimited by horizontal, obtusely pointed condyles (Figs 2F, 3D); distal margin of orifice and lip of sinus with minute tubercles. Operculum matching the orifice, with no additional sclerites (Fig. 2F). Some Scottish specimens with occasional orifices closed by perforated calcification (Fig. 5H). Holyhead specimens most commonly with a single avicularium lateral to the orifice but frequently none; distolaterally directed, inner end of hinge-line level with the condyles; mandibles triangular, their height scarcely exceeding the hinge width (Fig. 3A, B, D, E). The Scottish material has 0–5 avicularia per autozooid, with most colonies typically featuring 1–3; there may also be frontal avicularia, of the same basic form but slightly larger and with an elevated chamber (Fig. 5B, C). Ovicells prominent, subglobular, with numerous pores except near the mid-proximal margin; with slender sinuous ridges ascending from the distal zooid, between the pores, and converging in a mid-proximal direction (Fig. 3B, C); sometimes>1 (up to 5 in Scottish material) per autozooid (Fig. 5F–G, and discussion below). Polypide with c. 19 tentacles (Friday Harbor). Embryos reddish, apparently increasing in size during development; half-sized embryos present at Holyhead even in midwinter (February). Ancestrula with D-shaped orifice and 8 marginal spines; a central, patterned circular area on the frontal calcification (Fig. 4A–C); approximately 350–400 × ~ 300 µm overall (settlement panel, Stromness). Additional descriptions. Colonies collected from different parts of the world may vary, may offer reproductive stages not seen elsewhere, or be described in a slightly different manner. Full descriptions accompanied by SEM illustrations have recently been provided for each of the main geographic areas from which S. japonica is known—Alaska and the Pacific coast of North America (Dick et al. 2005) and Japan (Grischenko et al. 2007). Salient features have been selected. Alaska. Colony encrusting, unilaminar but sometimes bilayered as a result of overgrowth; colour ranging from whitish to red. Zooids distinct, separated by suture lines and shallow grooves. Frontal surface slightly to moderately convex, with marginal areolae and frontal pseudopores; pseudopores becoming infundibular with age and thickening calcification, the frontal shield becoming reticulate. Orifice medial or offset, with an avicularium beside it; usually broader than long, anter semi-circular, separated by paired blunt stout condyles, directed medially; operculum light golden brown, transparent. Avicularia paired, single or absent on any given zooid; additionally, occasional zooids bearing a somewhat larger frontal avicularium with raised rostrum and chamber. With increasing secondary calcification, the ovicells—similar to those from Holyhead—become increasingly rugose (Dick et al. 2005). Japan. Colonies as from Alaska but red to bright orange. Zooids with frontal shield uniformly porous except suborally, with 7–9 larger areolae along each margin; usually with a small suboral umbo. Oral avicularia most commonly single but frequently absent or paired; situated lateral or proximolateral to orifice; mandible elongatetriangular, its tip acute, with distal to distolateral orientation; crossbar complete; chamber comparatively small, with 1–3 minute pores laterally around the base; sometimes, in older parts of the colony and associated with complete ovicells, one avicularium is larger, with raised chamber. Zooidal communication via 3–5 distal and 6 lateral basal pore-chambers. Ovicells prominent, hemispherical, partially overhanging the orifice, evenly porous, with larger slit-like pores around the base; sparsely distributed or in a reproductive band within the colony. Ancestrula oval, imperforate, 0.33 × 0.28 mm; orifice D- shaped, 0.13 × 0.15 mm, with 8 marginal spines; 3 zooids budded distally (Grischenko et al. 2007). Remarks. Variations and discussion. Some striking variations that appear characteristic of S. japonica have been reported earlier and observed by us. A remarkable feature is the occurrence in Scottish material of multiple ovicells, arranged serially one behind another, and occasionally stacked (Fig. 5F–G). Powell (1970) and Powell et al. (1970) earlier described and illustrated a similar aberration in specimens from British Columbia and Washington State, and in a colony found on a scallop shell transplanted with oysters (Crassostrea gigas) from Onagawa Bay, on the Pacific coast of Honshu, Japan. Powell et al. (1970), using transplant experiments in Willapa Bay, Washington, attributed this occurrence of multiple ovicells to creosote-treated wood and the presence of petroleum derivatives in the water of harbours and marinas. It is not clear whether the occurrence of this phenomenon in Scotland is attributable to pollution; however, multiple ovicells were observed at sites all around the Scottish coastline. Powell et al. (1970) also noted that S. japonica (as S. unicornis) occurred in hyposaline water, down to salinities of 15. All of the British occurrences have been in marinas, suggesting that—unlike the Pacific coast of North America (see later)—small, ocean-going vessels must have been the vectors (Fig. 4F shows colonies on a boat hull). We have no evidence to suggest whether Japan or North America was the source. Distinction from Schizoporella unicornis. On the Atlantic coasts of Western Europe, including the British Isles, confusion of S. japonica is likely only with S. unicornis, although the usual habitats of the two species are quite different. Schizoporella japonica is so far known only from harbours and marinas, and is a typical fouling species; S. unicornis occurs in non-fouling situations, on stones, rocks, shells and kelp holdfasts on the lower shore and sublittorally. Comprehensive descriptions, variously illustrated, are available (Hayward & Ryland 1979; 1999; Ryland 1990; 1995; Tompsett et al. 2009) but, to facilitate ready comparison the morphological differences are summarized in Table 2., Published as part of Ryland, John S., Holt, Rohan, Loxton, Jennifer, Spencer Jones, Mary E. & Porter, Joanne S., 2014, First occurrence of the non-native bryozoan Schizoporella japonica Ortmann (1890) in Western Europe, pp. 481-502 in Zootaxa 3780 (3) on pages 485-490, DOI: 10.11646/zootaxa.3780.3.3, http://zenodo.org/record/4910530, {"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.","Okada, Y. (1929) Report of the biological survey of Mutsu Bay. 12. Cheilostomatous Bryozoa of Mutsu Bay. Scientific Reports of the Tohoku Imperial University, 4 (4), 11 - 35, pls 1 - 5.","Ross, J. & McCain, K. (1976) Schizoporella unicornis (Ectoprocta) in coastal waters of northwestern United States and Canada. Northwest Science, 50, 160 - 171.","Kubota, K. & Mawatari, S. (1985) A systematic study of bryozoans from Oshoro Bay, Hokkaido. 2. Ascophora. Environmental Science, Hokkaido, 8, 195 - 208.","Thorpe, J. P. & Ryland, J. S. (1987) Some theoretical limitations on the arrangement of zooids in encrusting Bryozoa. In: Ross, J. R. P. (Ed.), Bryozoa: present and past. Western Washington University, Bellingham, pp. 277 - 283.","Osburn, R. C. (1952) Bryozoa of the Pacific coast of America. Part 2, Cheilostomata-Ascophora. Allan Hancock Pacific Expeditions, 14 (2), 1 - 611.","Soule, D. F., Soule, J. D. & Chaney, H. W. (1995) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and western Santa Barbara Channel: The Bryozoa. Irene McCulloch Foundation Monograph Series, 2, 1 - 344.","Dick, M. H., Grischenko, A. V. & Mawatari, F. S. (2005) Intertidal Bryozoa (Cheilostomata) of Ketchikan, Alaska. Journal of Natural History, 39, 3687 - 3784.","Grischenko, A. V., Dick, M. H. & Mawatari, S. F. (2007) Diversity and taxonomy of intertidal Bryozoa (Cheilostomata) at Akkeshi Bay, Hokkaido, Japan. Journal of Natural History, 41, 1047 - 1161. http: // dx. doi. org / 10.1080 / 00222930701391773","Kelly, K. L. & Judd, D. B. (1955) The ISCC-NBS Method of Designating Colors and a Dictionary of Color names. National Bureau of Standards, Washington, D. C., 158 pp.","Powell, N., Sayce, C. S. & Tufts, D. F. (1970) Hyperplasia in an estuarine bryozoan attributable to coal tar derivatives. Journal of the Fisheries Research Board of Canada, 27, 2095 - 2096. http: // dx. doi. org / 10.1139 / f 70 - 234","Hayward, P. J. & Ryland, J. S. (1979) British ascophoran bryozoans. Synopses of the British Fauna, n. s., 14, 1 - 312.","Hayward, P. J. & Ryland, J. S. (1999) Cheilostomatous Bryozoa. Part 2, Hippothooidea - Celleporoidea. Synopses of the British Fauna, n. s., 14, 1 - 416.","Ryland, J. S. (1990) The lophophorate phyla: Phoronida, Bryozoa, and Brachiopoda. In: Hayward, P. J. & Ryland, J. S. (Eds.), Marine Fauna of the British Isles and North-west Europe. The Clarendon Press, Oxford, pp. 794 - 838.","Tompsett, S., Porter, J. S. & Taylor, P. D. (2009) Taxonomy of the fouling cheilostome bryozoans Schizoporella unicornis (Johnston) and S. errata (Waters). Journal of Natural History, 43, 2227 - 2243. http: // dx. doi. org / 10.1080 / 00222930903090140"]}

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."]}

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."]}

Forty-nine species of erect Bryozoa from a broad range of Cyclostome, Ctenostome, and Cheilostome families are described and illustrated, and an artificial dichotomous key is provided for their identification. In general, the marine bryozoan faunas of the northeastern coasts of theUnited States are poorly known; species records are sparse and voucher collections few, and it is certain that many more species occur in this region than are presently known. The species described here occur in intertidal, coastal or offshore habitats; some are well known and have been recorded on numerous previous occasions, others have been only rarely reported, while a few are known to occur commonly in the north of the region but have yet to be recorded south of Cape Cod. Some of the species described have not been recorded at all on northeastern coastsof the United States, but are widely distributed in North Atlantic continental shelf habitats and perhaps occur in similar parts of the outer shelf of this region. This fauna is thus provisional, but is intended to stimulate further work on the Bryozoa. (PDF file contains 52 pages.)