taxonID	type	description	language	source
03FBC9096877FFEE50F9FB11FE9DA8C0.taxon	type_taxon	Type species. Lepralia unicornis Johnston in Wood, 1844.	en	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
03FBC9096877FFEE50F9FB11FE9DA8C0.taxon	diagnosis	Diagnosis. Colony encrusting, unilaminar to multilaminar, sheet-like or mammillate, or developing partially erect plates and mounds. Autozooids with an evenly perforated cryptocystidean frontal shield. Orifice with clearly differentiated anter and poster, the latter more or less narrowed to form a sinus; condyles well defined. Avicularia adventitious, occasionally dimorphic, typically adjacent to the autozooid orifice. Oral spines impermanent or absent. Ovicell recumbent on distally succeeding autozooid, prominent, spherical or subglobular, perforate, ribbed or umbonate. Vertical walls with uniporous or multiporous septula (Hayward & Ryland 1999).	en	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
03FBC9096877FFEE50F9FB11FE9DA8C0.taxon	discussion	Remarks. The species of Schizoporella fall into two fairly distinct groups, based on the shape of the sinus. In the first, the sinus is broadly and shallowly U-shaped, wider than deep; in the second, the sinus is narrow, as deep — or deeper — than its greatest width. All the species to be discussed in this paper (S. errata, S. japonica, S. pseudoerrata and S. unicornis) belong in the first group. The main characters traditionally used for species separation in this group are: size, budding pattern, and layering of the colony; the shape of the orifice (including that of the sinus); the appearance of the porous frontal wall; and the morphology of the ovicell, especially ribbing and distribution of pores. Despite being reasonably distinct, the above species have often been confused with each other, especially when found in fouling communities, usually as a result of ignoring well-described characteristics. Known British species (also including S. errata, which has not yet been recorded from Britain) have been described by Hayward & Ryland (1995, 1999), while SEM-illustrated redescriptions of the type specimens of two of these (S. errata and S. unicornis) have been recently published (Tompsett et al. 2009). Details of the specimens we have studied are given in Appendix 1.	en	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
03FBC9096870FFE450F9FF3EFE3EAB82.taxon	description	(Figures 2 – 5)	en	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
03FBC9096870FFE450F9FF3EFE3EAB82.taxon	materials_examined	Material examined. See Appendix 1. Holotype. Strasbourg Museum, MZS Bry 001 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. 4 E.	en	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
03FBC9096870FFE450F9FF3EFE3EAB82.taxon	description	Description of British material. Colonies at first more or less circular (Fig. 4 F), rapidly becoming extensive, pale whitish-pink to vivid orange-red (Figs 2, 4; British material between Munsell 2.5 YR 6 / 14 and 10.0 R 6 / 10 (see Kelly & Judd 1955 )); mainly unilaminar but frequently with raised edges or displaying slightly elevated lobes (Fig. 2 B, 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 2 D – E, 3 A, 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 2 F, 3 D); distal margin of orifice and lip of sinus with minute tubercles. Operculum matching the orifice, with no additional sclerites (Fig. 2 F). Some Scottish specimens with occasional orifices closed by perforated calcification (Fig. 5 H). 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. 3 A, 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. 5 B, 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. 3 B, C); sometimes> 1 (up to 5 in Scottish material) per autozooid (Fig. 5 F – 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. 4 A – 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).	en	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
03FBC9096870FFE450F9FF3EFE3EAB82.taxon	discussion	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. 5 F – 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. 4 F 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.	en	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
03FBC909687AFFE250F9F8F3FEAFA8B1.taxon	description	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.	en	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
