Mycale (Mycale) grandis Gray 1867
Creators: name: Van, Rob W. M.
name: Aryasari, Ratih
name: De, Nicole J.
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Porifera
Demospongiae
Poecilosclerida
Mycalidae
Mycale
Mycale grandis

description: Mycale (Mycale) grandis Gray, 1867 ‘red’
Figs 78 a–g, 79a–k, 80a–j, 81a–k, Tables 6, 7
Esperia sp. Schmidt 1864: 34, pl. III fig. 11.
Mycale grandis Gray, 1867: 533; Hentschel 1912: 337, pl. XVIII fig. 13 (Indonesia); Burton 1934: 547 (no description, North Australia); Burton 1937: 23, pl. II fig. 13 (India); Lévi 1958: 29, fig. 24 (Red Sea); Vacelet & Vasseur 1965: 104 (Madagascar); Vacelet & Vasseur 1971: 86; Thomas 1986: 6, fig. 1n (Minicoy Island, Lakshadweep);? Tanita & Hoshino 1989: 116, fig. 70 (Japan); Pulitzer-Finali 1993: 291 (Kenya); Madrigal 1999: 4 (American Samoa); Coles & Bolick 2007: 911, fig.1 (Oahu, Hawaii, assumed invasive since mid 1990s); Vicente et al. 2016: 1 (Hawaii).
Esperia pellucida Ridley, 1884: 437, pl. XL fig. K, pl. XLII fig. h (from Torres Straits, North Australia); Chervyakova 2007: 242 (Southern Vietnam, listed only).
Esperia obscura; Ridley 1884: 438 (North Australia) (not: Carter 1882 = M. (Naviculina) obscura).
Esperia indica Carter, 1887: 72, pl. VI figs 3–6 (Andaman Sea).
Esperella pellucida; Topsent 1897: 462 (no description, Ambon);? Dawydoff 1952: 50 (Vietnam, listed only).
Mycale armata Thiele, 1903: 950, fig. 16 (Ternate); De Laubenfels 1954: 151, text-fig. 99 (mid Pacific); De Laubenfels 1955: 139 (mid Pacific); Eldredge & Smith 2001: B9–B10 (Hawaii); Coles et al. 2007: 1–30 (Hawaii).
Mycale indica; Burton & Rao 1932: 327 (India); Rao 1941: 445 (India).
Oxymycale strongylophora De Laubenfels, 1954: 94, text-fig. 57 (mid Pacific).
Mycale (Aegogropila) grandis; Coles et al. 1999: table 1 (Oahu, Hawaii, called non-indigenous); Putchakarn 2007: 1637, fig. 12 (Thailand).
Mycale (Mycale) indica; Pattanayak 2006: 64, fig. 43a–i, pl. VII fig. F (Andaman Islands).
? Mycale (Aegogropila) pellucida; Minh-Quang Thai 2013: 114 (Vietnam, listed only).
Mycale (Mycale) grandis; Calcinai et al. 2013: 33, figs 21A–L (Hawaii); Nuñez-Pons et al. 2017: 13 (Hawaii).
Material examined. Lectotype slide BMNH 1867.3.11.60 (‘Esp. ind’) from Schmidt (upon which Gray based his Mycale grandis). No further locality data.
Holotype slides BMNH 1882.2.23.224 (2 slides, ‘from type’), Esperia pellucida Ridley, 1884, Australia, Torres Strait, Alert Island, depth 13 m, bottom sand.
Holotype slides BMNH 1887.6.1.2 (3 slides, 2 of Carter, one labeled Bk (=Bowerbank) 1536), Esperia indica Carter, 1887, Myanmar, Mergui Archipelago, King Island.
Holotype fragment ZMB 3157, Esperella armata Thiele, 1903, Indonesia, Halmahera, Ternate, coll. W. K̹kenthal.
Holotype fragment, USNM 22937, Oxymycale strongylophora De Laubenfels, 1954, Ailing-lap-lap Atoll, Bikajela Island, depth 10 m, on dead coral, coll. ‘diver’, 20 June 1949.
ZMA Por. 01581, Indonesia, Lesser Sunda Islands, Sumba, Bay of Nangamessi, bottom coral sand, 9.6456°S 120.2642°E, depth 0–36 m, trawl, coll. Siboga Expedition stat. 053, field nr. SE 1487 IV, 21 April 1899; ZMA. Por. 01582, Indonesia, Maluku, Banda anchorage, bottom black sand, coral, 4.5398°S 129.9084°E, depth 9–45 m, trawl and dredge, coll. Siboga Expedition stat. 240, field nr. SE814, 22 November 1899; ZMA Por. 01583, Indonesia, Papua, NW coast Waigeu, Wunoh Bay, bottom coralline algae, 0.0901°S 130.2827°E, depth 32 m, dredge, coll. Siboga Expedition stat. 152, field nr. SE 1542 II, 12 August 1899; ZMA Por. 01584, Indonesia, Lesser Sunda Islands, E coast of Roti Island, Pepla Bay, bottom coral, mud, coralline algae, 10.6333°S 123.42°E, depth 22 m, dredge, coll. Siboga Expedition stat. 301, field nr. SE207 CXII, 30 January 1900; ZMA Por. 01585, Indonesia, Maluku, Banda anchorage, bottom black sand, coral, 4.5398°S 129.9084°E, depth 9–45 m, trawl and dredge, coll. Siboga Expedition stat. 240, field nr. SEDEIII, 22 November 1899; ZMA Por. 01586, Indonesia, Sulawesi, Salayar anchorage and surroundings, 6.0963°S 120.4481°E, depth 0–36 m, trawl, coll. Siboga Expedition stat. 213, field nr. SE3019, 26 September 1899; ZMA Por. 01587, Indonesia, Lesser Sunda Islands, E coast of Roti Island, Pepla Bay, bottom coral, mud, coralline algae, 10.6333°S 123.42°E, depth 22 m, dredge, coll. Siboga Expedition stat. 301, field nr. SE4XVI, 30 January 1900; ZMA Por. 01588, Indonesia, Papua, Aru Islands, Pearl Banks, anchorage off Pulau Jedan, bottom sand, shells, 5.4134°S 134.6677E, depth 13 m, trawl and dredge, coll. Siboga Expedition stat. 273, field nr. SE138, 23 December 1899; ZMA Por. 01589, Indonesia, Timor, S coast, anchorage off Noimini, 10.2333°S 124.0916°E, depth 8–36 m, trawl and dredge, coll. Siboga Expedition stat. 296, field nr. SE207 CXII, 24 January 1900; ZMA Por. 01590, Indonesia, Kalimantan, Borneo Bank, Pulau Sebangkatan, bottom coralline algae, 2.2183°S 117.4039°E, depth 34 m, dredge, coll. Siboga Expedition stat. 081, field nr. SE 1432 II, 14 June 1899; ZMA Por. 01591, Indonesia, Papua, Aru Islands, Pearl Banks, anchorage off Pulau Jedan, bottom sand, shells, 5.4134°S 134.6677°E, depth 13 m, trawl and dredge, coll. Siboga Expedition stat. 273, 23 December 1899; ZMA Por. 01592, Indonesia, Kalimantan, Borneo Bank, Pulau Sebangkatan, bottom coralline algae, 2.2183°S 117.4039°E, depth 34 m, dredge, coll. Siboga Expedition stat. 081, field nr. SE16, 14 June 1899; ZMA Por.P. 11923 (slide only), Indonesia, Kalimantan, Borneo Bank, Pulau Sebangkatan, bottom coral, coral sand, 2.2183°S 117.4039°E, depth 34 m, dredge, coll. Siboga Expedition stat. 081, field nr. SE889, 14 June 1899; ZMA Por. 02887, Indonesia, Sulawesi, anchorage off S point Kabaëna Island, 5.5165°S 121.9502°E, bottom coarse sand, depth 22 m, dredge, coll. Siboga Expedition stat. 209, field nr. SE397 II, 23 September 1899; ZMA Por. 07979, Indonesia, Sumba, NE coast, E of Melolo, reef flat, 9.9167°S 120.75°E, depth 1–4 m, snorkeling, coll. R. W.M. van Soest, Indonesian-Dutch Snellius II Expedition stat. 052, field nr. 052 / II/25, 14 September 1984 (brown); ZMA Por. 08997, Indonesia, SE Sulawesi, Tukang Besi Islands, Binongko, SW of Taipabu, 5.9333°S 123.975°E, depth 15–30 m, SCUBA, coll. H.A. Ten Hove, Indonesian-Dutch Snellius II Expedition stat. 044, field nr. 44 / V/05, 11 September 1984 (orange); ZMA Por. 09556, Singapore, Pulau Salu, 1.2164°N 103.7069°E, depth 0–2 m, snorkeling, coll. H. Moll, 22 December, 1977 (blood-red); ZMA Por. 09809, Australia, Queensland, Heron Island, SE, depth 0–8 m, snorkeling, coll. D. Vethaak, 1 April 1983 (red); ZMA Por. 10849, Indonesia, Nusa Tenggara, Lombok, Gili Air, 8.35°S 116.0333°E, littoral, snorkeling, coll. C.C. Hofman, field nr. 25, 7 August 1991 (orange); ZMA Por. 11157, Seychelles, Mahé, S of Aride Island, 4.2167°S 55.6667°E, depth 35 m, rectangular dredge, coll. R. W.M. van Soest, Netherlands Indian Ocean Expedition stat. 713, field nr. IOP-E 713/09, 19 December 1992 (red); ZMA Por. 11293, Seychelles, Amirantes, northern slope of Île Desnoeufs platform, 6.2167°S 53.0167°E, reef slope, depth 12-15 m, SCUBA, coll. R. W.M. van Soest, Netherlands Indian Ocean Expedition stat. 783, field nr. IOP-E 783/18, 2 January 1993 (orange); ZMA Por. 11679, Seychelles, Mahé, E of Mahé, N of Moyenne Island, 4.6167°S 55.5167°E, reef slope, depth 1–7 m, SCUBA, coll. R. W.M. van Soest, Netherlands Indian Ocean Expedition stat. 749, field nr. IOP-E 749/25, 25 December 1992 (red); ZMA Por. 11706, Seychelles, Amirantes, St. Joseph Atoll, S rim, 5.45°S 53.35°E, reef slope, depth 10 m, SCUBA, coll. R. W.M. van Soest, Netherlands Indian Ocean Expedition stat. 759, field nr. IOP-E 759/18, 28 December 1992 (orange); ZMA Por. 12145, Seychelles, Amirantes, N of Platte Island Atoll, 5.6167°S 55.3667°E, reef, depth 6 m, SCUBA, coll. H.A. Ten Hove, Netherlands Indian Ocean Expedition stat. 796, field nr. IOP-E 796/18, 7 January 1993 (red); ZMA Por. 12554, Seychelles, Mahé, SE coast, Anse Royale Bay, 4.7333°S 55.5167°E, reef slope, depth 2–13 m, SCUBA, coll. R. W.M. van Soest, Netherlands Indian Ocean Expedition stat. 740, field nr. IOP-E 740/13, 24 December 1992 (red); ZMA Por. 13017, American Samoa, Tutuila Island, Nu’uuli Lagoon, 14.2997°S 170.7225°W, encrusting underside of rubble, snorkeling, coll. L.G. Madrigal, field nr. S2, 24 May 1998 (red-orange); ZMA Por. 13026, American Samoa, Tutuila Island, Tafanal Reef, 14.2879°S 170.6631°W, encrusting underside of rubble, snorkeling, coll. L. Madrigal, field nr. S11, 24 May 1998 (red-orange); ZMA. POR. 18679, Thailand, Rayong, Samet islands, W side of Ko Samet, Hin Khan Na, 12.5539°N 101.4696°E, depth 3 m, under rock, SCUBA, coll. Sumaitt Putchakarn, field nr. SAMA–02, 26 January 2001 (red); ZMA Por. 18695, Thailand, Samet Islands, Ao Pgarung, S of Ko Samet, 12.5539°N 101.4696°E, on dead coral, depth 5 m, SCUBA, coll. Sumaitt Putchakarn, field nr. SAMB–06, 27 October 2001 (red); ZMA Por. 18727, Thailand, Rayong, Klang, Mon Islands, N of Ko Nai, 12.6073°N 101.6905°E, fringing reef, depth 4 m, SCUBA, coll. Sumaitt Putchakarn, field nr. MONB–04, 29 October 2001 (red); ZMA Por. 18753, Thailand, Trad, Chang Islands, S of Ko Mark, 11.786°N 102.4873°E, fringing reef, depth 3 m, SCUBA, coll. Sumaitt Putchakarn, field nr. CHAD–04, 19 November 2001 (red); ZMA Por. 18837, Thailand, Suratthani, Ko Tae-nai, W side of Ko Phangan, 9.7047°N 99.976°E, coral reef, depth 3 m, SCUBA, coll. Sumaitt Putchakarn, field nr SAID–01, 21 December 2001 (red, dried specimen); ZMA Por. 18908, Thailand, Rayong, Ban-Pae fishing pier, depth 15 m, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–41, 26 January 2003; ZMA Por. 18915, Thailand, Trad, Chonburi, Pataya fishing pier, depth 15 m, SCUBA, coll. Sumaitt Putchakarn, field nr. RU–POR–48, 15 December 1999; RMNH Por. 1545, Java Sea, 6.1417°S 107.7917°E, ‘ Gier’ Expedition stat. 3, 16 October 1907; RMNH Por. 2596, Singapore, Pulau Hantu Kecil, NE coast, 1.2281°N 103.7493°E, depth 9 m, SCUBA, coll. N.J. de Voogd, field nr. SIN.21/030406/158, 3 April 2006 (red); RMNH Por. 4400, Indonesia, Sulawesi, Spermonde Archipelago, no further data, depth 17 m, SCUBA, coll. N.J. de Voogd, 11 May 1997; RMNH Por. 8679, Thailand, coll. Jeep, field nr. 57/1, 2014 (red); RMNH Por. 11769, Indonesia, Sulawesi, SW Sulawesi, Spermonde Archipelago, Polewali, 4.8456°S 119.4011°E, depth 10 m, SCUBA, coll. N.J. de Voogd, field nr. CEL058, 24 April 2018 (red); RMNH Por. 11770, Indonesia, Sulawesi, SW Sulawesi, Spermonde Archipelago, Karanrang, 4.8556°S 119.3769°E, depth 10 m, SCUBA, coll. N.J. de Voogd, field nr. CEL113, 27 April 2018 (red).
Description (Figs 78 a–f, 80a). Red or red-orange lobate sponges growing inbetween live and dead corals, often cryptic, but more thinly encrusting specimens are also common (Fig. 78a); occasionally, parts of specimens may bear thin outgrowths. In living condition lobes carry large oscules, up to 1 cm diameter or more (Figs 78 b–d). Size of specimens usually modest, but may grow to 8 x7 x 5 cm. Surface often covered by sediment (Figs 78 a–b), but optically smooth when free. On deck (Fig. 78e), out of the water, the red color is seen to be the same inside as outside. In preserved condition (Figs 78f, 80a), the color becomes dirty white throughout, with shiny, slightly translucent surface, in older preserved specimens the color is often beige. Consistency firm, but compressible.
Skeleton (Figs 79 a–c). Choanosomal skeleton (Fig. 79b) plumoreticulate, with thick spicule tracts, in the interior up to 400 µm in diameter (up to 25 spicules in cross section), about 2 mm or more apart, irregularly interconnected by thinner tracts. Towards the surface the thick tracts subdivide into thinner tracts and fan out to carry the surface skeleton. This tangential ectosomal skeleton (Fig. 79a) is an irregular layer of individually intercrossing megascleres, occasionally forming short tracts of two or three spicules thick. Anisochelae I are not found in the ectosomal skeleton and do not form rosettes. Instead they ‘echinate’ the peripheral choanosomal tracts (Fig. 79c), hooked individually in low density into these with their lower alae.
Spicules (Figs 79 d–k, 80b–j, 81a–k). Mycalostyles, three-four size classes of anisochelae, two size categories of sigmas, trichodragmas.
Mycalostyles (Figs 79d,d 1, 80b,b 1, 81e), often curved but also straight, with elongated heads, mostly with pointed ends mucronate or more gradually tapering, occasionally rounded, variable in length and thickness, possibly regionally restricted, 351– 515.5 –672 x 6– 14.3 – 24 µm.
Anisochelae I, in most specimens divisible in two overlapping size categories, which also have subtle different shapes, here dubbed anisochela Ia and Ib.
Anisochela Ia (Figs 79e, 80c, 81 a–d,f), characteristically with a free part of the shaft making up 50% or more of the spicule length, with upper alae predominantly pointed although not truly unguiferous, occasionally oval, considerably standing off from the shaft, lower alae rounded, lower lateral alae positioned slightly above the median alae, size variable, possibly regionally determined, occasionally rare in individual specimens, total length 90– 117.8 – 156 µm.
Anisochelae Ib (Figs 79f, 80d), similar in overall shape to anisochelae Ia, but with shorter shaft, lower lateral and median alae more or less in the same position towards the shaft, alae rounded, not sharply pointed like anisochela Ia; these spicules are often rare (e.g. not observed in slide of Schmidt’s type material, cf. Fig. 81), but occasionally they are the dominant anisochelae I type in specimens, overall size slightly overlapping with anisochelae Ia, but the two types are clearly separate in size in individual specimens, 51– 73.2 – 96 µm.
Anisochelae II (Figs 79g, 80e,e 1, 81g), often rare or sometimes not found, but occasionally rather abundant. They are robust, with free part of the shaft short, 15–20% of spicule length, with well developed alae at both ends, the upper alae elongated with sharp-angled but rounded edges, the lower alae rounded, no spur, 22– 27.1 – 32 µm.
Anisochelae III (Figs 79h, 80f,f 1, 81h), usually abundant, with upper alae dominating the spicule, with lower alae absent or only visible vaguely as little knobs on the shaft, which is provided at the distal end with a prominent spur, and then curves upwards to form a stick-like reduction of the median alae, 12– 17.1 – 24 µm.
Sigma I (Figs 79i, 80g, 81i), usually abundant, but may be rare in some specimens, thin to moderately thick (1–2 µm), usually asymmetrical, variable in size among specimens, possibly regionally determined, 32– 53.8 – 81 µm.
Sigma II (Figs 79j, 80h, 81j), usually abundant, thin, likewise asymmetrical, 9– 17.3 – 23 µm.
Trichodragmas (Figs 79k, 80 i–j, 81k), usually common, usually fusiform in shape, but squarish forms are also found, size range overall considerable, but usually limited within an individual, occasionally both smaller and larger occur within the same specimen, 18– 57.1 –146 x 9– 11.2 – 18 µm.
Distribution and ecology (Fig. 86). This is one of the most widespread and common Mycale species in the study region, reported apparently correctly from the whole Indo-West Pacific region, from Hawaii to East Africa and from South China to North Australia. Specimens recorded here were all collected in shallow reef habitats, down to 45 m. In Hawaii the species is considered a recent invader (Coles et al. 1999; Eldredge & Smith 2001; Coles et al. 2007, as M. armata; Coles & Bolick 2007, as M. grandis). Distributions of sponges in the mid 20 th century were poorly known, but M. (M.) grandis was already known from Mid-Pacific localities previously (De Laubenfels 1954: as M. grandis and Oxymycale strongylophora; De Laubenfels 1955: as M. armata), and our present specimens from American Samoa add to bridging the gap. De Laubenfels’ Hawaii sponge investigations were not at all exhaustive, some sponges were studied from aquaria and where they were collected in the field it was done using local divers. Despite the great efforts of Coles et al. to assess the ecological impact of the species, the extensive Hawaii occurrence is likely to be at most a range extension or population increase rather than a genuine alien invasion.
The record from off Oshima Island near Tokyo, Japan by Tanita & Hoshino (1989) is perhaps doubtful, because no mention is made of trichodragmas. There are no further records of the species so far north.
Remarks. The only available type material of this species, Schmidt’s slide from the Natural History Museum, London, did not contain anisochelae Ib and anisochelae II, so formally our specimens do not conform to Gray’s Mycale grandis. However, the shape of anisochelae Ia (Fig. 81a) is obviously the same spicule as that found in all our specimens, and also the spicule measurements fall within the range observed in our specimens. No precise locality from which the slide was made is known for Schmidt’s specimen (‘von dem indischen Meere’), but it is likely from our study area. No further specimen details are known, as Schmidt obtained his material by dissolving it from a coral specimen. As the live color of the original specimen is unknown, it cannot be excluded that Schmidt’s material was from a white specimen. This is one of the reasons we here assign both the red and the white specimens (cf. below) to Mycale (Mycale) grandis sensu lato. Abundance of anisochelae II was found to be low in many specimens of both ‘species’, so it is proposed here to consider the deficient presence of anisochelae II in the lectotype of Mycale grandis as an issue of minor importance.
We re-examined type slides of Esperia pellucida Ridley, 1884 (Fig 81b) and E. indica Carter, 1887 (Fig. 81c) and found these two species without doubt to be identical to M.(M.) grandis sensu lato. The measurements provided by Carter (1887: 73) do not all appear to be accurate, as e.g. the size of the mycalostyles cited by Carter (1218 x 37.8) far exceeds those we measured in Carter’s slide BMNH 1887.6.1.2 (504–564 x 14–24 µm, roughly about half the size given by Carter). Later records of Mycale indica by Burton & Rao (1932: 327) give measurements similar to ours and do not at all match those provided by Carter, so it is likely Carter made a mistake. Unfortunately, he did not give magnifications with the drawings of the spicules in his pl. VI figs 3–6, just stating that the spicules were all magnified to the same scale. This is obviously incorrect.
We also re-examined a slide of Ridley’s (1884) specimen described under the name Esperia obscura and this was found to be misidentified, conforming to the present species and not to Carter’s species, which will be treated below as Mycale (Naviculina) obscura. Ridley (p. 439) already suggested that Mycale grandis could be conspecific with his obscura.
Thiele’s (1903) description of Mycale armata failed to mention the presence of anisochelae II and trichodragmas, but re-examination of a slide of the type material ZMB 3157 revealed the rare presence of anisochelae II and abundant presence of trichodragmas. We also found anisochelae II in a fragment of the holotype (cf. Fig. 81g).
Lévi’s (1958) Red Sea record is undoubtedly M. (M.) grandis. He reports the presence of four anisochelae types, of which the second largest (’ type normal’) clearly represents what we call here anisochelae Ib, whereas the third largest were characterized as ‘identique aux précédents’. We have clearly shown with SEM that the third largest anisochelae are distinct in shape from anisochelae Ib.
Pulitzer-Finali’s (1993) record of the species from Kenya also contained two sizes of anisochelae I, viz. 110– 130 µm and 55–70 µm, next to anisochelae II (23–29 m) and anisochelae III (14.5–17 µm). The specimens were orange in color.
Mycale (Mycale) anisochela Lévi, 1963 is a greyish subglobular species from the west coast of South Africa, falling outside the region we focus on in this study. It has a spicule complement virtually identical to Mycale (Mycale) grandis, including the characteristic shape of the anisochelae I, but the measurements of all spicules (except the sigmas) are clearly in excess of those of the many specimens described here from the tropical Indo-West Pacific: mycalostyles 950– 1050 x 26–28 µm, anisochelae I 220 µm, anisochelae II 60– 25 µm, anisochelae III 30 µm, sigmas I 52 µm, sigmas II 32, trichodragmas 50 µm. It is obviously closely related to M. (M.) grandis, as Lévi himself noted, but we consider it a valid species. The large size of the anisochelae is by no means the largest in Mycale, as Caribbean Mycale diaphana (Schmidt, 1870) (p. 57) has anisochelae, quite similar in shape to those of M. (M.) grandis and M. (M.) anisochela, measuring 650 µm. This latter measurement could not be confirmed by Carter (1882: 289), when he studied the type material in the Natural History Museum, London, the sizes he observed were about 120 µm. Maybe Schmidt made an error.
We compared the spicule sizes and categories of red or presumed red M. (M.) grandis specimens from Indo-West Pacific and Indian Ocean (cf. Table 6), but found few if any differences, indicating the likely wide distribution of the species.
description: Published as part of Van, Rob W. M., Aryasari, Ratih & De, Nicole J., 2021, Mycale species of the tropical Indo-West Pacific (Porifera, Demospongiae, Poecilosclerida), pp. 1-212 in Zootaxa 4912 (1) on pages 122-129, DOI: 10.11646/zootaxa.4912.1.1, http://zenodo.org/record/4450930