Sponges associated with octocorals in the Indo-Pacific, with the description of four new species Author Calcinai, Barbara Author Bavestrello, Giorgio Author Bertolino, Marco Author Pica, Daniela Author Wagner, Daniel Author Cerrano, Carlo text Zootaxa 2013 3617 1 1 61 journal article 39041 10.11646/zootaxa.3617.1.1 6174b35e-3228-4d50-a6ed-844e244f8b64 1175-5326 248150 4DCCD152-65DA-44A3-AB19-59811384E1E7 Mycale (Zygomycale) parishi ( Bowerbank, 1875 ) ( Fig. 30 A–M) ( Tab. 14 ) Examined material. Samples Port Allen 1, Port Allen 2, Port Allen 3: Hawai’i, Kaua’i Island , Port Allen Harbour, depth between 1–10 m , 0 2 June 2005 ; sample HK 10: Hawai’i, O’ahu Island , Hawai’i Kay, depth between 0.5–3 m , 31 May 2007 ; sample Carrambali 01: Indonesia , Bali Island , Garden eels I, 17 m , 18 October 2007 . Comparative material: holotype of Mycale (Aegogropila) pectinicola Hentschel, 1911 ZMB Por 4404 ( Fig. 31 A-I). Description. Pulpy sponge, about 2–3 mm thick, covering branches of Carijoa riisei up to the anthocodial bases and sometimes joining branches of the octocoral ( Fig. 30 A, B). Consistency elastic and with macroscopically smooth surface ( Fig. 30 B); with stereo-microscope examination, the surface is microhispid due to protruding mycalostyles. Beige alive ( Fig. 30 A), in ethanol pinkish ( Fig. 30 B). The Hawaiian specimens are nine ramified fragments of the octocoral, ~ 10 cm in length ( Fig. 30 A); the Indonesian specimens are two fragments, up to 10 cm , joined by the sponge tissue. Skeleton. Tracts of mycalostyles forming an ectosomal reticulum with triangular meshes ( Fig. 30 C). Choanosome consisting of plumoreticulate, ascending, interconnecting tracts, irregularly arranged ( Fig. 30 D). Anisochelae I arranged in rosettes in the choanosome. Spongin scarce. Spicules. Mycalostyles often flexuous and with a slightly ovoid head ( Fig. 30 E, L), 230 – 360 x 5 – 10 µm. Sigmas in two size classes: sigmas I “C” or “S” shaped ( Fig. 30 F), 45 – 107.5 x 2.5 – 7.5 µm; sigmas II “C” shaped ( Fig. 30 G, L), 14–43 µm. Palmate anisochelae in two size classes: anisochelae I are arranged in rosettes with straight shaft and short alae ( Fig. 30 H), 40–62.5 µm; anisochelae II slightly curved ( Fig. 30 I), 12–27.5 µm. Isochelae with very short lateral alae ( Fig. 30 J), 7–15 µm. Toxas in a large size range ( Fig. 30 K), 27.5–145 µm. Raphides arranged also in trichodragmas ( Fig. 30 M), 20–50 µm. Refer to Tab.14 for complete measurements. Distribution and remarks. Malacca Strait ( Bowerbank 1875 ), Mergui Archipelago ( Carter 1887 ), Australia ( Ridley 1884 ), Philippines ( Ridley & Dendy 1887 ), Indonesia and Palau (Hooper et al. 2000), Vietnam ( Azzini et al. 2007 ) and Hawai’i ( Bergquist 1967 ). TABLE 14. Spicule dimensions in µm of Mycale (Zygomycale) parishi (Bowerbank, 1875) and of the holotype of M. (Z.) pectinicola Hentschel, 1911 .

Mycalostyles	Sigmas I II	Anisochelae I II	Isochelae	Toxas	Raphides/ microxeas
M. (Z.) parishi Port Allen 1	250 – (266.5 ± 7.8) – 275 x 5	92.5 – (95.8 ± 23 – (29.4 3.5) – 100 x 2.5 – ± 7.1) – 43 (4.9 ± 2.1) – 7.5	40 – (45.1 12 – (19.2 ± 2.8) – 50 ± 4.8) – 25	10 – (11.9 ± 1.4) – 15	27.5 – (89.1 ± 34.2) – 145	absent
M. (Z.) parishi Port Allen 2	295 – (306.5 ± 11.3) – 330 x 6.2 – (7.6 ± 1.1) – 10	72.5 – (81.9 ± 14 – (27.2 11.4) – 107.5 x 2.5 ± 7.6) – – (5.1 ± 1.6) – 7.5 37.5	12.5 – 40 – (45 ± (17.3 ± 2.4) – 47.5 3.1) – 22.5	7 – (9.6 ± 1.7) – 13	45.0 – (66.5 ± 17.6) – 87.5	27.5 – (37.8 ± 6.6) – 50
M. (Z.) parishi Port Allen 3	285 – (302.5 ± 9.5) – 320 x 5	72.5 – (76.3 ± 17 – (29.5 3.6) – 82.5 x 5 ± 7.5) – 39	45 – (47.6 18 – (19.1 ± 1.5) – 50 ± 1.2) – 21	10 – (11 ± 0.8) – 12	31 – (65.3 ± 25.7) – 100	absent
M. (Z.) parishi HK10	300 – (330.2 ± 13.8) – 360 x 10	80 – (87.6 ± 12.5 – 3.2) – 95 x 5 – (26.5 ± (6.6 ± 1.2) – 7.5 9.8) – 40	50 – (55.1 20 – (23.6 ± 3.9) – ± 2.4) – 62.5 27.5	10 – (12 ± 1.1) – 12.5	60 – (80.5 ± 11.0) – 100	32.5 to 42.5
M. (Z.) parishi Carrambali 0 1	230 – (258.9 ± 18.4) – 290 x 2.5 – (3.8 ± 1.1) – 5	45 – (68.1 ± 15 – (26.3 15.7) – 80 x 2.5 ± 5.8) – 35 – (4.1 ± 1.2) – 5	17.5 – 40 – (42.9 (18.9 ± ± 2.2) – 45 1.3) – 20	7.5 – (9.5 ± 1.1) – 10	32.5 – (52.8 ± 12.8) – 72	20 – (23.7 ± 2.1) – 25
M. (Z.) pectinicola Holotype ZMB Por 4404	260 – (271 ± 10.9) – 290 x 5 – (5.3 ± 0.9) – 7.5	80 – (84.5 ± 15 – (26 ± 6.7) – 95 x 5 9.4) – 35	10 – (18.3 40 – (43.7 ± 3.5) – ± 1.8) – 45 22.5	10 – (11.5 ± 1.3) – 12.5	15 – (34.6 ± 15) – 60	15 – (19.6 ± 2.2) – 19.6

FIGURE 30. Mycale (Zygomycale) parishi (Bowerbank, 1875) . A, Sponge (HK 10) covering Carijoa riisei ; B, Close up of sponge surface; C, Ectosomal reticulum of triangular meshes of mycalostyles; D, Choanosome of plumoreticulate, ascending, interconnecting tracts; E, Mycalostyle; F, Sigma I; G, Sigma II; H, Anisochela I; I, Anisochela II; J, Isochelae; K, Toxa; L, Magnification of mycalostyle head and sigma II; M, Raphide. According to van Soest et al. (2011) three species belong to this subgenus: M. (Z.) angulosa (Duchassaing & Michelotti, 1864) , M. (Z.) parishi ( Bowerbank, 1875 ) and M. (Z.) ramulosa Carballo & Cruz-Barraza, 2010 . Mycale (Aegogropila) pectinicola Hentschel, 1911 was described with mycalostyles, additional isochelae, two classes of anisochelae, two classes of sigmas and microxeas ( Fig. 31 A–I); the examination of its holotype confirmed the presence of isochelae ( Fig. 31 F), but also of numerous toxas ( Fig. 31 G) and micracanthoxeas ( Fig. 31 I) not originally observed by Hentschel (1911) . Because of the presence of isochelae, this species has to be moved to the subgenus Zygomycale . FIGURE 31. Mycale pectinicola Hentschel, 1911 holotype ZMB Por 4404. A, Mycalostyle and close up of the head; B, Sigma I; C, Sigma II; D, Anisochela I; E, Anisochelae II; F, Isochelae; G, Toxas; H, Microxea; I, Micracanthoxeas. Mycale (Z.) pectinicola differs from our specimens by: i) the shape of the isochelae, which are characterised by a tooth on the central ala, variable in length ( Fig. 31 F, Hentshel 1911 : 300 Fig. 8 d); ii) microxeas instead of raphides; iii) the presence of micracanthoxeas. The dimensions of toxas in M. (Z.) pectinicola are slightly smaller ( Tab. 14 ). The Atlantic species M. (Z.) angulosa (Duchassaing & Michelotti, 1864) is quite similar to M. (Z.) parishi ; the latter shows a large variation in spiculation, which is not reported for M. (Z.) angulosa (van Soest 1984 ) . Mycale (Z.) ramulosa differs from M. (Z.) parishi mainly in the presence of micracanthoxeas. Our specimens are similar to M. (Z.) parishi in skeletal organisation and spicule complement. No differences in general morphology or in spiculation were recorded between the Hawaiian and Indonesian specimens ( Tab. 14 ). Mycale (Z.) parishi is a thinly encrusting species, known to grow over other organisms (molluscs, bryozoans). In Hawai’i it was recorded mainly in the fouling communities of harbours (see Carlton & Eldredge 2009 ). The re-examination of the type of M. ( A .) hentscheli (see M. ( A. ) furcata ) and of M. (Z.) pectinicola revealed the existence of other two species of Mycale with micracanthoxeas, besides the six species known thus far ( Carballo & Cruz-Barraza 2010 ): M. (C.) micracanthoxea , M. (C.) urizae , M. (A.) bamfieldensis , M. (A.) escarlatei , M. (A.) lilianae and M. (Z.) ramulosa . Four of these ( M. ( A .) hentscheli , M. (Z.) ramulosa , M. (Z.) pectinicola and M. (A.) bamfieldensis are described in the Indo-Pacific ocean. It is interesting to note that micracanthoxeas are not a character restricted to genera Carmia and Aegogropila ( Carballo & Hajdu 1998 ) , but are also present in two species of Zygomycale . This subgenus is mainly based on the presence of isochelae, but some authors considered it synonymous with Aegogropila , recognizing the character “isochelae” as plesiomorphic (see comments in van Soest & Hajdu 2002b ). The discovery of micracanthoxeas in two species of Zygomycale adds further controversy to the currently accepted subgeneric classification of Mycale , stressing the need for clarification of these phylogenetic relationships.