Watersipora subatra ( Ortmann, 1890 ) Vieira & Jones & Taylor, 2014

Watersipora subatra ( Ortmann, 1890) View in CoL n. stat.

( Figs 39–53 View FIGURES 39–42 View FIGURES 43–47 View FIGURES 48–53 , 66 View FIGURES 65–68 , 69 View FIGURES 69–71 ; Table 3 View TABLE 3 )

Schizoporella aterrima var. subatra Ortmann, 1890: 49 [ Japan].

? Watersipora cucullata var. watersi Mawatari, 1952: 12 View in CoL (part), fig. 1G [ Japan].

Dakaria subovoidea: Harmer 1957: p. 1022 (part), pl. 49, figs 11–12, 14 [ Indonesia].

? Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium].

Watersipora subtorquata: Ryland 1974: 345 View in CoL , fig. 3C [Low Island]. Non Cellepora subtorquata d’Orbigny, 1852: 399 [ Brazil]. “ Watersipora subovoidea View in CoL ” fide Harmer: Soule & Soule 1975: 308 (part), pl. 3, fig. 6 [ Australia].

“ Watersipora subovoidea View in CoL ” fide Harmer: Winston & Heimberg, 1986: 15, figs 35–37 [Komodo].

Watersipora subtorquata: Gordon 1989: 40 View in CoL , pl. 20, B–H [ New Zealand].

Watersipora subtorquata: Gordon & Mawatari, 1992: 30 View in CoL [ New Zealand].

Watersipora subtorquata: Ryland et al. 2009: 55 View in CoL View Cited Treatment , figs 3, 4A, C, E, F [ Australia, Bay of Biscay and English Channel]. Watersipora subtorquata: Kelso & Wyse-Jackson 2012: 2010 View in CoL , fig. 1 [ Ireland].

Watersipora subtorquata: Kuhlenkamp & Kind 2013: 3 View in CoL , figs 1B, C, 2A–F [North Sea: Helgoland].

Material examined. Holotype: MZS 002 View Materials , Ortmann Collection , Sagami Bay, Japan, 50–100 fm (15–30 m), 1882 . Other material: NHMUK 1922.10 View Materials .8.3, dry slide, Schizoporella cucullata , sea level, Misaki , Japan, No. 1, Insole coll. NHMUK 1929.12 View Materials .31.1, dry slide, Watersipora atrofusca , Australia, MacGillivray coll. NHMUK 2005.5 View Materials .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 View Materials .14.9, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands , Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12 View Materials .14.10, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands , Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12 View Materials .14.12, balsam slide, Watersipora subtorquata, J.S. Ryland don., on shell, Bay of Arcachon , August 2003, Hans de Blauwe coll. NHMUK 2007.12 View Materials .14.14, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12 View Materials .14.15, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12 View Materials .14.16, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007 , R. Lord coll. NHMUK 2007.12 View Materials .14.17, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007 , R. Lord coll. NHMUK 2007.12 View Materials .14.18, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007 , R. Lord coll. NHMUK 2010.12 View Materials .1.2, dry, Watersipora sp. , Point Loma , Marina, San Diego, California, USA, September 2010. P.D. Taylor & B. Okamura coll. NHMUK 2010.6 View Materials .30.7, dry and dry slide, Long Beach , Los Angeles, USA . NHMUK 2012.6 View Materials .30.6, dry slide [part of NHMUK 2005.5.26.19], Watersipora subtorquata , scraped from hull of Seasprite , hauled out Leigh Wharf / Marina, North Island, New Zealand, 6 October 2004, K. Tilbrook coll.

Description. Colonies encrusting, multiserial, uni- to multilamellar; sometimes erect, foliaceous and bilamellar; colour in life variable, from orange to brownish-purple or greyish to black. Zooids subrectangular to hexagonal, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield slightly convex, with numerous round pseudopores about 25 µm diameter; two large latero-oral intrazooidal septula near lateral zooidal margin, proximolateral to orifice, each with 3–8 (often 5) small pores. Frontal shield obscured by translucent cuticle. Orifice large, subcircular to oval, slightly wider than long, with broad, well-defined proximal, U-shaped sinus; orificial rim thin, sometimes slightly raised; narrow bar-shaped condyles, sometimes inconspicuous. Operculum with broad, biconcave proximal band, without lucidae. Avicularia absent. Ooecia absent.

Remarks. Ortmann (1890) described and figured Schizoporella aterrima from Japan, including the new variety subatra without drawings or description. Ryland et al. (2009) noted that Schizoporella aterrima Ortmann, 1890 is unrecognizable and that the description given by Mawatari (1952) for Watersipora cucullata var. watersi , another species reported from Japan, may include at least two species. One of us (MSJ) has examined Ortmann’s specimens deposited at the Musée zoologique de la Ville de Strasbourg, concluding that they represent two species, Watersipora aterrima (MZS 003; Figs 35–38 View FIGURES 35–38 ; Table 3 View TABLE 3 ) and Watersipora subatra (MZS 002; Fig. 39–42 View FIGURES 39–42 ; Table 3 View 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 View FIGURES 43–47 ), California ( Fig 48–53 View FIGURES 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 View FIGURES 48–53 ) often have a well-demarcated, U-shaped sinus and a smooth suborificial region (without pseudopores), as has been figured in specimens from Brittany ( Ryland et al. 2009, fig. 3A, D, as W. subtorquata ). All specimens assigned to W. subatra have an orifice with a U-shaped sinus and narrow, bar-shaped condyles, opercula with a broad, biconcave band proximally, frontal shields with pseudopores 18–30 µm in diameter, and two latero-oral intrazooidal septula.

Soule and Soule (1975) indicated that Harmer (1957) included at least two species under Dakaria subovoidea , which may be a mixture of W. subtorquata and W. aterrima . According to the list of specimens published by Harmer (1957), however, his description and remarks included at least four species: W. subtorquata , W. cucullata , W. atrofusca and W. subatra (specimens figured from Siboga Stns 181 and 184, see Harmer 1957, pl. 49, figs 11–12, 14). Watersipora subatra is distinguished from W. subtorquata by the shapes of the orifice and condyles, and the presence of latero-oral intrazooidal septula ( Figs 47 View FIGURES 43–47 , 52, 53 View FIGURES 48–53 ). Latero-oral intrazooidal septula are also found in W. cucullata ( Figs 8 View FIGURES 6–11. 6–9 , 29 View FIGURES 25–29 , 34 View FIGURES 30–34 ), W. aterrima ( Fig. 37 View FIGURES 35–38 ) and W. mawatarii n. sp. ( Fig. 58 View FIGURES 54–58 ; see below), but these species have different shapes of orifice and condyles.

Despite the wide distribution of W. subatra in Pacific waters (see Ryland et al. 2009; Mackie et al. 2012; as W. subtorquata ), its occurrence in some localities in the northwest Pacific, as well as the occurrence of W. subtorquata , are unconfirmed and require morphological and molecular investigation.

Distribution. NE Atlantic ( Ireland and British Isles), Indo-West Pacific ( Indonesia) and Pacific ( New Zealand, Australia and California).