Mycale (Zygomycale) parishii

Mycale (Zygomycale) parishii (Bowerbank, 1875)

Figs 116 a–f, 117a–d, 118a–i

Raphiodesma parishii Bowerbank, 1875: 283.

Amphilectus parishii; Vosmaer 1880: 119.

? Esperia tunicata; sensu Carter 1880: 49 (Not: Schmidt 1862)

Esperia plumosa Carter, 1882: 299, pl. XI figs 19a–b; Carter 1887: 62.

Esperia parishi; Ridley 1884: 436.

Esperella parishii; Ridley & Dendy 1887: 65; Dendy 1905: 159.

Esperella ridleyi Von Lendenfeld, 1888: 211 (including varieties intermedia and robusta); Hallmann 1914: 402, fig. 13.

Esperella plumosa; Dendy 1905: 159; Dendy 1916: 121, Pl. I figs 4a–g, Pl. III fig. 19; Dendy & Frederick 1924: 503.

Mycale parishii; Burton & Rao 1932: 328.

Zygomycale parishii; De Laubenfels 1950: 25, fig. 16; Lévi 1956: 14; Lévi 1963: 14, pl. IIA, text-fig. 10; Thomas 1973: 38, pl. II fig. 10; Thomas 1979: 58, pl. III fig. 11; Thomas 1986: 6, fig. 1p.

Zygomycale parishi; Bergquist 1968: 58.

Zygomycale plumosa; Pulitzer-Finali 1993: 291.

(Not: Mycale plumosa; Tanita 1958: 133, pl. II figs 10–11, text-fig. 5; Kim et al. 1968: 41; Hoshino 1981: 171; Tanita & Hoshino 1989: 118, text-fig. 72 (identity currently not established).

Mycale parishi; Hooper & Wiedenmayer 1994: 290.

Mycale (Zygomycale) parishi; Lévi et al. 1998: 115; Van Soest & Hajdu 2002: 687, fig. 15; Azzini et al. 2007: Table 1 (listed only).

Mycale (Zygomycale) parishii; Eldredge & Smith 2001: B7–8; Putchakarn 2007: 1637, Fig. 13; Minh-Quang Thai 2013: 114 (listed only); Calcinai et al. 2013: 46, figs 30A–M; Nuñez-Pons et al. 2017: 16 (not: Lakwal et al. 2018: unnumbered fig. p. 212).

Material examined. ZMA Por. 01605, Indonesia, Lesser Sunda Islands, Banda Sea, 7.3233°S 116.825°E, depth 538 m (?), bottom dark brown sandy mud, trawl, coll. Siboga Expedition stat. 316, field nr. SE1261, 19 February 1900; ZMA Por. 01606, Indonesia, Papua, Aru Islands, Pearl Banks, anchorage off Pulau Jedan, 5.4134°S 134.6677°E, bottom sand and shells, depth 13 m, coll. Siboga Expedition stat. 273, field nr SE 1599 III, 23 December 1899; ZMA Por. 01607, Indonesia, Kalimantan, Borneo Bank, 2.7167°S 117.7333°E, bottom coral sand, depth 41–54 m, trawl, coll. Siboga Expedition, stat. 079, field nr SE1554, 12 June 1899; ZMA Por. 01608, Indonesia, Papua, off Raja Ampat, 1.7083°S 130.7916°E, bottom sand, small stones and shells, depth 32 m, coll. Siboga Expedition stat. 164, field nr SE337, 20 August 1899; ZMA Por. 01609, Indonesia, Lesser Sunda Islands, Rotti Island, Cyprus Bay, 10.8733°S 123.0183°E, bottom corals and coralline algae, depth 34 m, coll. Siboga Expedition stat. 299, field nr SE1893, 27 January 1900; ZMA Por. 01610, Indonesia, Papua, Aru Islands, Pearl Banks, anchorage off Pulau Jedan, 5.4134°S 134.6677°E, bottom sand and shells, depth 13 m, coll. Siboga Expedition stat. 273, field nr SE95, 23 December 1899; ZMA Por. 09590, Singapore, Pulau Hantu, 1.2359°N 103.8395°E, littoral, depth 0–2 m, snorkeling, coll. H. Moll, 21 January 1978 (live color grey-brown); ZMA Por. 17073, China, Zhan Jiang, South China Sea, NaoZhou Island, 20.84°N 110.6°E, depth 12 m, SCUBA, coll. Xu Shi Hai, Yang Kai & He Huan Hua, field nr. 2002–05, 8 May 2002 (violet); ZMA Por. 17160, India, Muttom, SW coast, depth 21 m, SCUBA, coll. Anita George, field nr. 35, 2002, no further data (brick red, dried); ZMA Por. 17214, India, Muttom, SW coast, depth 12 m, SCUBA, coll. Anita George, field nr 68, 2002, no further data (red, dried); ZMA Por. 17216, India, Kanuakumari, SE coast, depth 15 m, SCUBA, coll. Anita George, field nr. 70, 2002, no further data (red, dried); ZMA Por. 17524, Thailand, Talokapo, obtained from local fishery, coll. R. G. Moolenbeek & H. Dekker, field nr. Moo 03/06 no. 3, 16 April 2003; ZMA Por. 17537, Thailand, Maruat village, on shells dumped at local fishery, coll. R. G. Moolenbeek & H. Dekker, field nr. Moo 03/06 no. 21, 16 April 2003; ZMA Por. 18604, Thailand, Chon Buri, Ko Khang Khao, S of Ko Sichang, 13.1068°N 100.8148°E, fringing reef, on bivalve shell, depth 5 m, SCUBA, coll. Sumaitt Putchakarn, field nr. SICC–03, 19 September 2001 (greyish purple); ZMA Por. 18665, Thailand, Chon Buri, Pattaya, Lan Islands, W side of Ko Lan, Hat Thain, 12.9162°N 100.7721°E, fringing reef on rocky bottom, depth 5 m, SCUBA, coll. Sumaitt Putchakarn, field nr. LANT–12, 25 September 2001; ZMA Por. 18691, Thailand, Ao Pagarung, Samet Islands, S of Ko Samet, Rayong, 12.5539°N 101.4696°E, fringing reef on sandy bottom, on bivalve shell, depth 3 m, SCUBA, coll. Sumaitt Putchakarn, field nr. LANT–12, 27 October 2001 (pale orange red); ZMA Por. 18694, Thailand, Ao Pagarung, Samet Islands, S of Ko Samet, Rayong, 12.5539°N 101.4696°E, fringing reef on sandy bottom, on rock, depth 5 m, SCUBA, coll. Sumaitt Putchakarn, field nr. SAMB–05, 27 October 2001 (orange); ZMA Por. 18700, Thailand, Ao Pagarung, Samet Islands, S of Ko Samet, Rayong, 12.5539°N 101.4696°E, fringing reef on sandy bottom, on bivalve shell, depth 5 m, SCUBA, coll. Sumaitt Putchakarn, field nr. SAMB–11, 27 October 2001; ZMA Por. 18758, Thailand, Trad, Chang Islands, Ko Ra-yong, 11.7883°N 102.4548°E, coral community on rock, on dead sea whip, depth 9 m, SCUBA, coll. Sumaitt Putchakarn, field nr. CHAE–02–12, 20 November 2001 (red brown); ZMA Por. 18853, Thailand, Trad, Ao Yai, S of Ko Kood, coral reef on sandy bottom, depth 6 m, SCUBA, coll. Sumaitt Putchakarn, field nr. KOGA–01, 4 May 2002 (pale red); ZMA Por. 18879, Thailand, Ban-Pae fishing pier, Rayong, sand, on crag gill net, depth 15 m, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–11, 26 January 2003; ZMA Por. 18885, Thailand, Ban-Pae fishing pier, Rayong, sand, on crag gill net, depth 15 m, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–17, 26 January 2003; ZMA Por. 18904, Thailand, Ban-Pae fishing pier, Rayong, sand, on crag gill net, depth 15 m depth, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–37, 26 January 2003; ZMA Por. 18911, Thailand, Ban-Pae fishing pier, Rayong, sand, on crag gill net, depth 15 m, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–44, 26 January 2003; ZMA Por. 19189, India, SW India, Goa, depth 19 m, SCUBA, coll. Anita George, IERSE, field nr. 25, 18 October 2005; ZMA Por. 19232, India, Kerala, Vizhinjam, SW coast, depth 13 m, SCUBA, coll. Anita George, IERSE, field nr. 72, 1 December 2005; ZMA Por. 19256, India, Lakshadweep Islands, Minicoy, depth 5 m, SCUBA, coll. Anita George, IERSE, field nr. 96, 10 December 2005; ZMA Por. 20681, Madagascar, Nosy Bé, Lokobé, depth 10 m, SCUBA, coll. J.H. Stock, field nr. MD12, 18 December 1963 (orange-red); ZMA Por. 20723, Madagascar, Nosy Bé, Crater Bay, depth 4–10 m, SCUBA, coll. J.H. Stock, 21 December 1963 (red); ZMA. Por. P.12195, specimen not retrieved, slide only, Taiwan, SW coast, depth 40 m, on ‘pectinid’ shell, coll. H.H. Dijkstra, no further data; RMNH Por. 2472, Singapore, Pulau Semakau, NW side, 1.2162°N 103.7556°E, SCUBA, coll. N.J. de Voogd, field nr. SIN.05/2700306/034, 27 March 2006 (orange); RMNH Por. 2473, Singapore, Pulau Semakau, NW side, 1.2162°N 103.7556°E, SCUBA, coll. N.J. de Voogd, field nr. SIN.05/2700306/035, 27 March 2006; RMNH Por. 2533, Singapore, Pulau Tekukor (Monkey Island), SE side, 1.2301°N 103.8402°E, SCUBA, coll. N.J. de Voogd, field nr. SIN.12/300306/095, 30 March 2006 (orange); RMNH Por. 2534, Singapore, Pulau Tekukor (Monkey Island), SE side, 1.2301°N 103.8402°E, SCUBA, coll. N.J. de Voogd, field nr. SIN.12/300306/096, 30 March 2006 (orange); RMNH Por. 5465, Indonesia, Halmahera, Tidore, N of Desa Rum, 0.7433°N 127.3851°E, SCUBA, coll. N.J. de Voogd, Ternate-Halmahera Expedition stat. TER:18, field nr. TER:18/041109/216, 4 November 2009; RMNH Por. 7288b, Taiwan, Penghu Islands, GoPoYu, 23.7127°N 119.5524°E, depth 12 m, SCUBA, coll. Y. Huang, field nr. TW0020, 19 August 2010 (greyish brown); RMNH Por. 7324, Taiwan, Penghu Islands, Suogang, 23.5191°N 119.6116°E, depth 17.2 m, SCUBA, coll. Y. Huang, field nr. TW0059, 4 July 2012; RMNH Por. 7808, Vietnam, Phu Quoc, 9.969°N 104.0158°E, depth 18.9 m, SCUBA, coll. Nguyen Khac Bat, field nr. MC5, 13 May 2013; RMNH Por. 7883, Vietnam, Hon Cau, 12.2311°N 108.8356°E, depth 11 m, SCUBA, coll. Nguyen Khac Bat, field nr. MC15–30, 9 June 2013; RMNH Por. 7891, Vietnam, Hon Cau, 12.2311°N 108.8356°E, depth 11 m, SCUBA, coll. Nguyen Khac Bat, field nr. MC15-90, 9 June 2013; RMNH Por. 8707, Madagascar, 4 Frères, 12.9943°S 48.4875°E, depth 24 m, SCUBA, coll. Anna Bialecki, field nr. MAD12 – IM132, 1 December 2012 (orange).

Description (Figs 116 a–f, 117a–b). Polymorphic habitus, starting out as a thin micronulose crust (Fig. 116c), often growing on other marine invertebrates as molluscs, dead gorgonians, bryozoans, etc. The crusts grow out to thick masses (Fig. 116 d–e) with conulose surface. The conules in more developed specimens extend into shorter or longer spikes. The largest specimens have a massive base with long ramose projections (‘plumose’) covered in spinous processes (Figs 116 a–b, f, 117a–b). Colors in life (Figs 116 a–f) vary between pinkish red, brownish orange, greyish brown or red-brown (e.g. on deck photo of Fig. 117b), in preservation this changes to shades of dirty white or beige (Fig. 117a). Sizes of smaller encrusting specimens are in the 1–3 cm range, with thickness up to 0.5 cm or less. Larger elaborate specimens have bases of 5–8 cm high and wide, with upright branches up to 15 cm high 3–4 cm thick, with spiny projections of 1–2 cm long and 3 mm thick. The surface of thinly encrusting specimens shows faint venal patterns and finely punctate skin in life, and often the surface reticulation is visible in larger magnification. Most specimens have conules, which tend to grow out sharply, but blunt lobate specimens also occur. Oscules are usually moderately large, around 1 cm in diameter, and occur irregularly between the conules and spiny projections. Consistency of the thin specimens firm, compressible, of the larger spiny specimens hard, difficult to cut.

Skeleton (Figs 117 c–d). Choanosomal skeleton plumoreticulate (Fig. 117c), with spicule tracts in more elaborate specimens reaching large dimensions in the lower and central parts of the skeleton, up to 500–600 µm in diameter, dividing and anastomosing in the direction of the surface, with thinner tracts of about 100 µm in diameter interconnecting the thicker upward running tracts. Near the surface, tracts peter out to about 50 µm, fanning out just below the surface membrane to carry the usually strongly developed tangential surface skeleton. This is of the aegogropila-type (Fig. 117d), with strong intercrossing tracts, in various specimens varying 20–110 µm in diameter, enclosing triangular meshes 200–350 µm in widest dimension. Thin encrustations usually have their skeleton less robustly developed and tracts significantly thinner than more elaborate specimens. All types of microscleres are found in the ectosomal meshes, including rosettes of anisochelae of 100–180 µm in diameter. In many specimens the isochelae are crowded around the outlines of the ostia and along the walls of the larger channels.

Spicules (Figs 118 a–i). Mycalostyles, three categories of anisochelae, isochelae, two categories of sigmas, toxas, trichodragmas.

Mycalostyles (Figs 118a,a 1), robust, straight or sometimes slightly wavy, heads elongate, with barely visible ‘neck’, opposite ending sharply pointed, 232– 279.9 –358 x 3– 7.2 – 12 µm.

Anisochelae I (Fig. 118b), well-developed, robust, free part of the shaft 40–45% of spicule length, with comparatively narrow upper median alae, which extends slightly outward, with comparatively less well developed lower median alae, 34– 46.4 – 61 µm.

Anisochelae II (Fig. 118c), well-developed, similar in shape to anisochelae I but with free part of the shaft less than 30% of spicule length, 19– 23.5 – 31 µm.

Anisochelae III (Fig. 118d), well-developed, in profile with upper median alae more narrow, and with free part of the shaft even shorter than in anisochelae II, 13– 17.4 – 21 µm.

Isochelae (Fig. 118e), usually comparatively narrow and lateral alae under-developed, although this varies over the region, 8.5– 9.9 – 12.5 µm.

Sigmas I (Fig. 118f), robust, thickness 3–5 µm, comparatively narrow-shaped, asymmetrical, 58– 78.4 – 126 µm.

Sigmas II (Fig. 118g), thin, often more or less symmetrical, variable in size, 14– 22.1 – 32 µm.

Toxas (Figs 118h), usually thin, often in dragmas of 4 or 5 spicules, curved widely and with small upturned endings, in a wide length range, 18– 57.9 –111 x up to 10 µm.

Trichodragmas (Figs 118i), consisting of moderately thick raphides of about 1 µm in thickness, 18– 28.4 –39 x 5–6 µm

Distribution and ecology (Fig. 119). Indonesia, Singapore, Thailand, Vietnam, South China, Taiwan, India, Madagascar; literature records from Malaysia, Sri Lanka, North and West Australia, Seychelles and Hawaii. Shallow water down to 40 m (except one likely eroneous record from 538 m)

Remarks. The polymorphic habitus has given rise to the distinction of two species names, M. (Z.) parishii and M. (Z.) plumosa, which have persistently been in use for respectively thinly or more massively encrusting specimens and upright ramose specimens. After having studied many specimens of both types and intermediate forms, we cannot confirm that these names represent two species, in stead we recognize only a single polymorphic species.

Hooper & Wiedenmayer (1994) suggested to follow Burton & Rao’s (1932) proposed synonymies of this species. The latter authors decided to ignore reported differences between Indo-West Pacific specimens possessing tiny isochelae because such differences would be subject to variability. They recognized only a single species. We admit that there is considerable variability in frequency of occurrence of the various microsclere categories, but cannot confirm that entire absence is also occurring within specimens assigned to the present species. Studying a large number of samples over a larger region has convinced us that searching intensively for each of the microsclere types and categories almost always is succesfull. Thus we reject Burton & Rao’s conclusions that Mycale (Zygomycale) isochela Hentschel, 1911, and Mycale (Zygomycale) pectinicola Hentschel, 1911, are junior synonyms of the present species. For the latter species, Calcinai et al. (2013) reported the presence of an additional spicule type, micracanthoxeas, which we can confirm (cf. below).

Carter’s (1880) record of the Mediterranean-Atlantic species Esperia tunicata Schmidt, 1862 from the Gulf of Manaar was suggested to belong to the present species by Dendy (1905) (p. 159), followed by Topsent (1924) (p. 99). The descriptive remarks by Carter are so meagre, that no identity can be derived from it, so we must assume Dendy had access to the specimen.

Ridley’s (1884) record of the species from Darwin, North Australia, failed to mention the presence of toxas, but we assume they were overlooked. We follow Hallman (1914: 402) in his conclusion that West Australian Esperella ridleyi Von Lendenfeld, 1888 and its varieties are junior synonyms of the present species. The variety intermedia is a junior secondary homonym of Mycale (Oxymycale) intermedia (Schmidt, 1874). Of the two varieties of M. ridleyi, robusta and intermedia, one must be the typical variety (ICZN Art. 47). Since this is not yet assigned, we assign here Esperella ridleyi var. intermedia Von Lendenfeld, 1888 as the typical variety, to be named Esperella ridleyi var. ridleyi, which removes the secondary homonymy with Mycale (Oxymycale) intermedia (Schmidt, 1874). Both varieties are synonyms of the present species. Hooper & Wiedenmayer (1994) based on Hooper et al. (1992) claimed occurrence of the present species (listed as Mycale ridleyi) along the N, NE and SE coasts of Australia but so far the detailed records remain unpublished.

The junior synonym Mycale plumosa Carter, 1882 was employed by Japanese (e.g. Tanita 1956: 133; Hoshino 1981: 171; Tanita & Hoshino 1989: 118) and Korean (e.g. Kim et al. 1968: 41) authors to describe a Mycale species lacking isochelae, despite Carter’s explicit description and drawing of these spicules. Re-examination of these specimens will be necessary to determine to which species they belong, but they do not conform to the present species. So far, no Mycale (Zygomycale) are known from these Northwest Pacific regions.

One record from India (Lakwal et al. 2018) is a blackish mass of lobed tubes, which is definitely not the present species. Lacking a description this specimen from the Ratnagiri coast of Northwest India remains unidentified.

The Hawaii records are considered invasive by Eldredge & Smith (2001), but the species was already reported as locally common from Kaneohe Bay, Hawaii in 1947 by De Laubenfels (1950 b). Like with M. (M.) grandis (cf. above) the case for an alien invasion is not very strong. The authors suggest the origin of the invasive species is Caribbean because De Laubenfels (1936) used that name for the Florida records of what later became to be recognized as Mycale (Zygomycale) angulosa (Duchassaing & Michelotti, 1864), a closely related but different species. De Laubenfels’ hypothesis of a global distribution of M. (Z.) parishii was followed by Hechtel (1965), but later Caribbean authors, e.g. Van Soest (1984: 16), restricted the name to Indo-West Pacific records. The species is widely distributed over that region, occurring from East Africa to Hawaii. The absence of records from localities between New Caledonia and Hawaii is in our opinion more likely to be caused by lack of research of the many South and Mid Pacific islands, rather than to invasive events. There are similar species in West African (M. (Z.) sierraleonensis Van Soest et al. 2014) and East Pacific waters (M. (Z.) ramulosa Carballo & Cruz-Barrazo, 2010), indicating the likely global tropical distribution of a Tethyan ancestor, differentiated into local species.

We compared spicule sizes of West Pacific and Indian Ocean specimens (cf. Table 8) and concluded that there is hardly any regional difference; perhaps the length range of the toxas in West Pacific specimens (18–111 µm) exceeds that of the Indian Ocean specimens.