Mycale species of the tropical Indo-West Pacific (Porifera, Demospongiae, Poecilosclerida) Author Van, Rob W. M. Author Aryasari, Ratih Author De, Nicole J. 0000-0002-7985-5604 rob.vansoest@naturalis.nl text Zootaxa 2021 2021-01-19 4912 1 1 212 journal article 8641 10.11646/zootaxa.4912.1.1 8a5efe86-cabc-4981-afb4-163791f2530c 1175-5326 4450930 9536C1CF-4AEF-47F8-959B-48CD7A5392D8 Mycale (Carmia) phyllophila Hentschel, 1911 Figs 41 a–e, 42a–c, 43, 44a–e, 45, Table 3 ? Esperella philippensis (sic); sensu Lindgren 1898: 302 , pl. 19 figs 13a–c (not: Dendy 1896 , cf. above). Mycale phyllophila Hentschel, 1911: 294 , fig. 5; Lévi 1963: 10 , text-fig. 3; Van Soest 1982: 88 , fig. 5. ? Mycale phillipensis ; sensu Pulitzer-Finali, 1982b: 101 (not: Dendy 1896 , cf. above). Mycale adhaerens ; Li 1986: 86 (Chinese), 109 (English), pl. I fig. 8, text-fig. 8 (not: Lambe 1894 ). ? Mycale (Carmia) phyllophila ; Minh-Quang Thai 2013: 114 (listed only). Material examined . ZMB Por 4401, syntypes (two specimens, cf. Figs 43 ), Australia , West Australia , Sharksbay, 2.5 miles SW of Denham , coll. Hartmeyer & Michaelsen , Hamburg SW Australia Expedition, 10 June 1905 . ZMA Por. 01614, Indonesia , Sulawesi , Salayar anchorage, 6.0963°S 120.4481°E , depth 0–36 m , reef exploration, coll. Siboga Expedition stat. 213, field nr. SE243, 26 September 1899 ; ZMA Por. 01615, Indonesia , Maluku , Ambon anchorage, 3.6907°S 128.1687°E , depth 36–54 m , dredge, coll. Siboga Expedition stat. 181, field nr. SE1314, 5 September 1899 ; ZMA Por. 01810, Indonesia , Maluku , W coast Kur Island , Killsuin anchorage, 5.3513°S 131.9529°E , depth 20–45 m , dredge, coll. Siboga Expedition stat. 250, field nr. SE272, 6 December 1899 ; ZMA Por. 04481, Hong Kong , Ah Chau , 22.5506°N 114.2662°E , coll. G. Thompson , April 1978 ; ZMA Por. 04482, Hong Kong , Chek Chau , 22.4972°N 114.3486°E , coll. G. Thompson , April 1978 ; ZMA Por. 06523, Indonesia , Nusa Tenggara , Sumbawa, N coast, Bay of Sanggar , 8.3417°S 118.2617°E , coastal reef with sea grass, depth 0–1 m , snorkeling, coll. J. Brouns, Indonesian-Dutch Snellius II Expedition stat. 120, field nr. 120/15, 21 September 1984 (live color yellow) ; ZMA Por. 08035, Indonesia , Nusa Tenggara , Komodo, NE cape, 8.4833°S 119.5683°E , coastal reef, depth 1–4 m , snorkeling, coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 096, field nr. 096 / II/11 , 19 September 1984 (red) ; ZMA Por. 08156, Indonesia , Maluku , Ambon , Ambon Bay near Eri , 3.75°S 128.1333°E , sandy bay with patch reef, depth 4–6 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 006, field nr. 006 / III/1 , 2 September 1984 (orange) ; ZMA Por. 08384, Indonesia , Nusa Tenggara , E of Komodo , Selat Linta , 8.5833°S 119.57°E , reef, depth 4–11 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 079, field nr. 079 / III/08 , 18 September 1984 (red) ; ZMA Por. 08385, Indonesia , Nusa Tenggara , E of Komodo , Selat Linta , 8.5833°S 119.57°E , reef, depth 4–11 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 079, field nr. 079 / III/09 , 18 September 1984 (red) ; ZMA Por. 08820, Indonesia , Nusa Tenggara , Komodo, NE cape, 8.4833°S 119.5683°E , coastal reef, depth 10–17 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 096, field nr. 096 / IV/12 , 19 September 1984 (red) ; ZMA Por. 08904, Indonesia , Sulawesi , SE Sulawesi , NE Take Bone Rate ( Tiger Islands ), 6.5°S 121.1333°E , reef, depth 10–15 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 139, field nr. 139 / IV/23 , 25 September 1984 ; ZMA Por. 08938, Indonesia , Sulawesi , SE Sulawesi , SW Salayar , reef N of Pulau Bahuluang , 6.45°S 120.43°E , depth 10–15 m , SCUBA , coll. R . W.M. van Soest , Indonesian-Dutch Snellius II Expedition stat. 169, field nr. 169 / IV/06 , 30 September 1984 (red-brown) ; ZMA Por. 12148, Seychelles , Amirantes , N of Platte Island Atoll , 5.8167°S 55.3667°E , depth 1 m , snorkeling, coll. E. Coppejans , Netherlands Indian Ocean Expedition stat. 796, fieldnr. IOP-E 796/22, 7 January 1993 (wine-red) ; ZMA Por. 13100, Indonesia , Sulawesi , SW Sulawesi , Samalona , depth 6 m , SCUBA , coll. N.J. de Voogd , field nr. SA /NV/200497/27, 20 April 1997 (red) ; ZMA Por. 13297, Indonesia , Sulawesi , SW Sulawesi , Kudingareng Keke , depth 15 m , SCUBA , coll. N.J. de Voogd , field nr. KK/NV/240497/18, 24 April 1997 (red) ; ZMA Por. 16532, Seychelles , Amirantes , N of Platte Island Atoll , 5.8167°S 55.3667°E , depth 6 m , SCUBA , coll. R . W.M. van Soest , Netherlands Indian Ocean Expedition stat. 796, field nr. IOP-E 796/22, 7 January 1993 (wine-red) ; ZMA Por. 17597, Hong Kong , Wong Shek , near fish farm, 22.4167°N 114.333°E , on rope collected at 150 m depth , coll. H.U. Dahme, 10 May 2003 (red); ZMA Por. 18344, Indonesia , Nusa Tenggara , Bali , 3–20 m depth , coll. Susilo, field nr. TRB72, 15 October 2003 (brownish red); ZMA Por. 21325, China , Hainan , Lingshui , coll. Yu-Wei Guo , field nr. LS–248, 31 August 2004 (red) ; ZMA Por. 21326, China , Hainan , Lingshui , coll. Yu-Wei Guo , field nr. LS–248, 31 August 2004 (yellow) ; ZMA Por. 22182, Mauritius , depth 3–5 m , SCUBA , coll. M. Bhikajee , field nr. VIIsp10, 2010 (reddish brown); RMNH Por. 6583, Indonesia , Sulawesi , N Sulawesi , Lembeh Strait , Teluk Walemetodo , 1.4032°N 125.1723°E , deph 1 m , SCUBA , coll. N.J. de Voogd , field nr. LEM29/150212/224, 15 February 2012 ; RMNH Por. 8202, Vietnam , Ha Long Bay , near Bui Xam island , fish farm, depth 0.5–1 m , coll. N.J. de Voogd, field nr. VIE042, 17 August 2013 . Description ( Figs 41a , 43 ). Thickly to massively encrusting sponges with optically smooth surface. Size up to 15 x 5 cm , thickness up to 0.5–1 cm . Substratum irregularities beneath the crust may occasionally push up the surface into lobes and projections. No visible openings in preservation. Many specimens encrust mangrove roots, algae, dead corals, and rubble. Life colors reported as red, orange, bright orange, red brown, yellow, wine red, (most often times cited as red). In preservation, colors are shades of beige or grey. Consistency crumbly, easily damaged. Skeleton ( Figs 42 a–c). The choanosomal skeleton consist of characteristically wispy, sinuous, closely separated, megasclere tracts, 40–150 m in diameter (up to 20 spicules in cross section), running upwards to the surface, dividing into thinner tracts and ending in often wide spicule brushes of up to 500 µm wide. These brushes are adjacent to each other and make an effective surface cover, but there are no tangential spicules, all lie at various sharp angles to the surface membrane. The tissue between the tracts is charged with dense amounts of individual sigmas and anisochelae ( Fig. 42 b–c). No rosettes. Spicules ( Figs 41 b–e, 44a–e). Mycalostyles, two categories of isochelae, sigmas. Mycalostyles ( Figs 41b,b 1 ), robust, comparatively straight and short, 206– 264.1 –302 x 2.5– 6.1 – 10 µm . FIGURE 41 . Mycale (Carmia) phyllophila Hentschel, 1911 , RMNH Por. 6583 from Indonesia, a, preserved habitus on mangrove roots (scale bar = 1 cm), b–e, SEM images of spicules, b, mycalostyle, b1, details of mycalostyle, c, anisochela I, d, anisochela II (same magnification), e, sigma. Anisochelae I ( Figs 41c , 44 a–e upper row), well developed, narrow-shaped, free part of shaft 20–25% of spicule length, upper alae at the rim slightly curved outwards, 18– 21.7 – 28 µm . Anisochelae II ( Figs 41d , 44 a–e lower row), similar in shape to anisochelae I, with free part of the shaft slightly larger (30–35% of spicule length), 11– 15.3 – 19 µm . Sigmas ( Fig. 41e ), comparatively variable over the region, but no size categories within individual specimens, well developed, thickness 1–2 µm , almost symmetrical in shape with slightly incurved endings, 24– 33.6 – 45 µm . Distribution and ecology ( Fig. 45 ). Indonesia , Seychelles , South Africa , West Australia , Vietnam , China . On mangrove roots, algae, corals, and other hard substratums, from the tidal zone down to 54 m or beyond. Remarks . A common species occurring all over the region. No distinct regional differences in spicule sizes were detected (cf. Table 3 ). FIGURE 42 . Mycale (Carmia) phyllophila Hentschel, 1911 , ZMA Por. 13100 from Indonesia, a–c, light microscopic views of skeleton and spicules, a, cross section of peripheral skeleton, b, detail of the same, showing brushed ending of skeletal tracts and spicule complement, c, microscleres including anisochela I and II (arrows). Lindgren’s misspelled Esperella philippensis Dendy could be this species, although there are other possibilities. Dendy’s species (cf. above) is a member of subgenus Mycale (Aegogropila) , whereas Lindgren described a typical member of Mycale (Carmia) . There is no mention of two size categories of anisochelae, so on paper it could be e.g. M. (C.) lissochela (cf. above), but the abundance of sigmas points more in the direction of the present species. Reexamination of Lindgren’s specimen from 45 m depth off the South Vietnamese coast is necessary. FIGURE 43 . Mycale (Carmia) phyllophila Hentschel, 1911 , type specimens ZMB 4401 from SW Australia, growing on brown algae. FIGURE 44 . Mycale (Carmia) phyllophila Hentschel, 1911 , SEM images of anisochelae I (upper row) and anisochelae II (lower row), to show variation in morphology in five different specimens, a, RMNH Por. 6583, b, ZMA Por. 01615, c, ZMA Por. 01614, d, ZMA Por. 08820, e, ZMA Por. 06523. FIGURE 45 . Mycale (Carmia) phyllophila Hentschel, 1911 , approximate distribution of specimens studied (black squares) and literature records (pink dots). Squares and dots may represent several close collecting localities. Van Soest’s (1982) record from Hong Kong failed to distinguish two size categories of anisochelae, but they were clearly present after re-examination (see ZMA Por. 04481 and 04482). Pulitzer-Finali’s (1982b) report of M. phillipensis (cf. above) from Hong Kong was made in the same general volume on the Hong Kong marine fauna ( Morton & Tseng 1982 ) as Van Soest’s report on M. phyllophila . These two reports were based on small collections sent independently to both spongologists without further coordination. We suggest here that the Hong Kong Mycale ’s described by both concern the same species, M. (C.) phyllophila , but the samples need to be compared to make sure. Li (1986) reported the NE Pacific Mycale adhaerens ( Lambe,1894 ) from the Gulf of Tonkin , S China . His description mentions only a single anisochela size, but his drawing shows two sizes. Since the present species is apparently common in Chinese waters (cf. above), we believe it is likely that Li’s record also belongs here. TABLE 3. Comparison of West Pacific and Indian Ocean Mycale (Carmia) phyllophila Hentschel, 1911 . Specimens West Pacific: ZMA Por. 01614, 01615, 01810, 04481, 04482, 06523, 08156, 08384, 08385, 08405, 13100, 13297, 17597, 18344, 21325, 21326, RMNH Por. 6583. Specimens Indian Ocean: Type ZMB 4401, ZMA Por. 12148/16532/22182 (cf. text for locality data of the specimens). AI= anisochelae I, AII = anisochelae II, SI = sigmas I.

Mycalostyles	AI	AII	SI
West Pacific	234–302 x 2.5–10	16–28	12–19	24–54
Indian Ocean	206–286 x 4–7	19–22	11–18	31–40

East Pacific Mycale (Carmia) cecilia De Laubenfels, 1936 , as extensively redescribed by Carballo & Cruz-Barraza (2010) , apparently does not have two size categories of anisochelae, although the size range is large ( 12.5–27.5 µm ). The authors assume that Hawaii Mycale (Carmia) maunakea De Laubenfels, 1951 is a junior synonym, but in view of the wide geographic separation, we are not convinced (cf. also below). We examined slides from the Galapagos Islands [(ZMA Por. 11246, cf. Desqueyroux-Faúndez & Van Soest 1997: 450 ) and RMNH.Por. 11536 (coll. N.J.DV, field nr. GAP054)] and found the spicules rather variable in length and thickness, subtly different from specimens of Mycale (Carmia) phyllophila . An extensive comparison of specimens, preferably including molecular sequences, from both sides of the Pacific ‘divide’ is necessary to reach a conclusion. This is beyond the ambitions of our present study. Outside our target region, New Zealand Mycale (Carmia) hentscheli Bergquist & Fromont, 1988 (as Carmia hentscheli ) is also close to the present species, as already remarked by the authors themselves. We re-examined a spicule suspension of this species (NNMZ 166), thanks to Eduardo Hajdu’s thesis work in Amsterdam, and found a.o. that the ‘not abundant’ sigmas were so rare that they were virtually absent. The authors described the life color as black-red, dark purple, orange or red, and presence of filamentous algae was noted. Although, it is definitely closely related it is clearly not conspecific with M. (C.) phyllophila . The species is known for its rich source of secondary metabolites (e.g. Rust et al . 2020 ). Mexican-Pacific Mycale (Carmia) contax ( Dickinson, 1945 ) as redescribed by Carballo & Cruz-Barraz (2010) shows spicule similarity in having two closely similar anisochelae sizes as in the present species, but the specimen contains toxas and raphides.