Boring sponges (Porifera, Demospongiae) from the Indian Ocean

Italian Journal of Zoology ISSN: 1125-0003 (Print) 1748-5851 (Online) Journal homepage: http://www.tandfonline.com/loi/tizo20 Boring sponges (Porifera, Demospongiae) from the Indian Ocean Barbara Calcinai, Carlo Cerrano, Michele Sarà & Giorgio Bavestrello To cite this article: Barbara Calcinai, Carlo Cerrano, Michele Sarà & Giorgio Bavestrello (2000) Boring sponges (Porifera, Demospongiae) from the Indian Ocean, Italian Journal of Zoology, 67:2, 203-219, DOI: 10.1080/11250000009356314 To link to this article: https://doi.org/10.1080/11250000009356314 Copyright Taylor and Francis Group, LLC Published online: 28 Jan 2009. Submit your article to this journal Article views: 844 View related articles Citing articles: 16 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalinformation?journalcode=tizo20

Ital. J. Zool.; 67. 203-219 (2000) Boring sponges (Porifera, Demospongiae) from the Indian Ocean BARBARA CALCINAI CARLO CERRANO MICHELE SARÀ Dipartimento per lo Studio del Territorio e delle sue Risorse, Università di Genova, C.so Europa 26, I-16132 Genova (Italy) e-mail: zoologia@unige.it GIORGIO BAVESTRELLO Istituto di Scienze del Mare di Ancona, via Brecce Bianche, I-60131 Ancona (Italy) INTRODUCTION The sponge fauna of the Indian Ocean was widely studied in the past; numerous works were produced from South Africa, the Bay of Bengal, the Gulf of Mannar, Sri Lanka etc. (see Thomas, 1976a). Knowledge of the boring sponge fauna of the Indian Ocean comes, in particular, from the works of Carter (1887) for King Island; Dendy (1905, 1921) for the Gulf of Mannar and Seychelles; Annandale (1915a, b) for the Bay of Bengal, the Persian Gulf, and the south-west of the Indian Peninsula; Levi (1956, 1958, 1961) for the Red Sea, Aldabra Island; Vacelet et al. (1976) for Madagascar; Burton (1937, 1959) for the Gulf of Mannar and South Africa; Thomas (1976b, 1979a, 1986, 1989) and Pattanayak (1999) for various parts of the Indian Ocean. In all these papers a total of 43 Indian Ocean species are considered to be boring (Table I). In the present work, a collection of boring sponges from the Maldives and Seychelles archipelagos were studied. This area was previously investigated by Thomas (I973, 1981) who studied demosponges, comprising boring species, from Mahe Island in the Seychelles bank, and the sponge fauna of Laccadives (Thomas, 1989); for the Maldives, the only report is that of Topsent (1905) regarding a specimen of Cliothosa hancocki (Thoosa hancocki) collected by M. J. Stanley Gardiner in Male Atoll (Maldives). Since then, the boring sponges of the Maldives have not been studied until now. With this work three new species and two new records for the area are reported, leading the number of boring species, hitherto known from the Indian Ocean, to 48. MATERIAL AND METHODS ABSTRACT A collection of boring sponges from the Maldives and Seychelles reefs was studied. From the etched organogenous material, 16 boring sponge species (belonging to seven genera and five families), were identified; three of these species are new and two are reported for the Indian Ocean for the first time. A list of the boring species hitherto recorded from the area is given. The boring sponge faunas from the Maldives and Seychelles show certain affinities among them and also with that of the entire Indo-Pacific region. KEY WORDS: Boring sponges - Taxonomy - Indian Ocean. (Received 2 June 1999 - Accepted 20 November 1999) The material for this study was collected in three different periods. In February 1996 and March 1998 in Maldives, at Gangehi and Kuda Rah Islands (Ari Atoll); in January 1997 in the Seychelles, where sampling was conducted at Praslin, La Digue and Les Soeurs. All the examined specimens were studied on dried preserved material coming from dead branches of the stony s of the genera Acropora and Tubastera, from portions of the massive Pontes, from the base of colonies of the hydro Millepora, and from shells of dead molluscs {Hippopus, Pleuroplocá). All samples were collected by SCUBA diving, with hammer and chisel when necessary. Very often the boring activity of these sponges was clearly visible because of the presence of coloured papillae, but sometimes it could be identified only by breaking down dead portions. Often the same substratum was shared by two or more sponges. In the laboratory, substrata infested by sponges were broken down and studied under a stereomicroscope. The sponge tissue filling the chambers was taken for spicular preparations by dissolving the sponge fragments in boiling nitric acid; quick preparations were made to avoid loss of rare and minute spicules and to avoid contamination among sponges sharing the same substratum. Electronmicrographs of the cleaned spicules, coated with gold, were obtained using a PHILIPS 515 scanning electron microscope. The holotypes were deposited at the Museum of Natural History of Genoa (MSNG). Published online 28 Jan 2009

TABLE I - List of the boring species hitherto recorded from the Indian Ocean (new records for the area are reported in bold). to o Species Substrata in the Indian Ocean Principal references Samus anónima Gray, 1867, calcareous algae Jaspis investigatrix Annandale, 1915b shell /. penetrans (Carter, 1880) Dercitus plicatus (Schmidt, 1868) Spirastrella cuspidifera (Lamarck, 1813) S. cunctatrix Schmidt, 1868 not known S. pachispyra Levi, 1958 boring S. vagabunda Ridley, 1884 not known Alectona primitiva Topsent, 1932 Cliona acustella Annandale, 1915a C. annulifera Annandale, 1915a C. celata Grant, 1826 C dichotoma sp. n. C. ensifera Solías, 1878 C. incostans (Dendy, 1887) C. indica Topsent, 1891 C. jullieni Topsent, 1891 C. kempi Annandale, 1915b C. lobata Hancock, 1849 C. margaritiferae Dendy, 1905 shell shell gastropod shell shell, rocks, calcareous algae stony, gastropod, buried in sand not known rock shell C. michelini Topsent, 1888 shell Gulf of Mannar (1); Mahe Is. (2); Seychelles (3); Ceylon Mahe Is. Malay penisula (1); Gulf of Mannar, Palk Bay (2); Mahe Is. (3); Maldives (4) Gulf of Aden, Wide distribution in the Indian Ocean (1); Minicoy (2); Gulf of Mannar, Palk Bay, Red Sea (3) Madagascar (1); Red Sea (2); Seychelles (3) Red Sea (1); Aldabra (2); Madagascar (3); Gulf of Mannar, Palk Bay (4) Ceylon, Mannar, Madagascar (1); Aden (2); Seychelles (3); Zanzibar (4) Madagascar Gulf of Bengal, Ceylon Ceylon Gulf of Bengal, Madras, Gulf of Mannar, Red Sea, Ceylon (1); Mahe Is. (2); West India (3); Mozambique (4); Minicoy (5); Palk Bay (6) Maldives Mergui, Andaman, Gulf of Bengal, Ceylon (1); Madagascar (2); Gulf of Mannar, Palk Bay (3); Maldives (4) Salomon (1); Mannar (2); Aldabra (3); Mahe Is. (4); Madagascar (5); Kenia (6); Mozambique (7); Minicoy (8);Gulf of Mannar, Palk Bay (9); Maldives (10) Ceylon Madagascar Andamans, Port Blair, Bay of Bengala (1); Gulf of Mannar, Palk Bay (2) Madagascar (1); Gulf of Mannar, Palk Bay (2) Ceylon (1), Palk Strait (2); West India (3); Mahe Is, Gulf of Mannar (4); Mozambique (5) Indian Ocean Carter, 1880 (1); Carter, 1887 (2); Thomas, 1972 (3) Thomas, 1979b Thomas, 1973 Solías, 1902 (1); Thomas, 1972 (2); Thomas, 1973 (3); present work (4) Burton, 1959 (D; Thomas, 1980 (2); Thomas, 1986 (3) Levi, 1956 (1); Levi, 1958 (2); present work (3) Levi, 1958 (1); Levi, 1961 (2); Vacelet etal., 1976 (3); Thomas, 1986 (4) Levi, 1956 (1); Topsent, 1893 (2); Dendy, 1921 (3); Lendenfeld, 1897 (4) Vacelet & Vasseur, 1971 Annandale, 1915a Annandale, 1915a Annandale, 1915 (1); Thomas, 1973 (2); Thomas, 1975 (3); Thomas, 1979a (4); Thomas, 1980 (5); Thomas, 1986 (6) present work Annandale, 1915a (1); Vacelet etal., 1976 (2); Thomas, 1972 (3); present work (4) Dendy, 1921 (1); Burton, 1937 (2); Levi, 1961 (3); Thomas, 1973 (4); Vacelet et al, 1976 (5); Thomas, 1976b (6); Thomas, 1979a (7); Thomas, 1980 (8); Thomas, 1986 (9); present work (10) Annandale, 1915a Vacelet et al, 1976 Annandale, 1915b (1); Thomas, 1979b (2) Vacelet et al., 1976 (1); Thomas, 1979b (2) Dendy, 1905 (1); Annandale, 1915a (2); Thomas, 1975 (3); Thomas, 1979b (4); Thomas, 1979a (5) Annandale, 1915a (contidued) n o O CO i

C. mucronata Solías, 1878 C. mussae (Keller, 1891) C nodulosa sp. n. C. Orientalis Thiele, 1900 C. patera (Hardwicke, 1822) C. schmidti (Ridley, 1881) C varians (Duchassaing & Michelotti, 1864) C. viridis (Schmidti, 1862) Pione carpenteri (Hancock, 1865) P. vastifica (Hancock, 1849) Scantittetta evispira (Topsent, 1898) Cliothosa aurivilli (Lindgren, 1897) C. hancocki Topsent, 1888 C. quadrata Hancock, 1849 Delectona higgini Carter, 1880 Dotona pulchella Carter, 1880 Thoosa armata Topsent, 1888 T. investigations Annandale, 1915a T. laeviaster Annandale, 1915a T. radiata Topsent, 1888 T. socialis Carter, 1880 Amorphinopsis excavans Carter, 1887 Zyzzya fuliginosa (Carter, 1879) Aka diagonoxea Thomas, 1968 A. laccadivensis Thomas, 1989 A. maldiviensis sp. n. A. minuta Thomas, 1972 stony Gulf of Mannar (1); Madagascar (2); Gulf of Mannar, Palk Bay (3); Mozambique (4); Maldives (5) Red Sea Maldives dead, Mergui, Malay archipelagos (1), Red Sea, Gulf of Mannar, shell Palk Bay (2); Maldives (3) bivalves and Java gastropods shells rock, Seychelles (1); Amirantes, Madagascar (2); Maldives (3) gastropod Maldives, Seychelles bivalves, s Mergui (1); Mahe Is., Red Sea (2); Gulf of Mannar; Palk Bay (3) bivalves shell, Mergui, Bay of Bengal (1); West India (2); Seychelles (3, 4), Maldives (4) shells,, Madras, Palk Strait, Persian Gulf (1); Red Sea (2); West India (3); calcareous algae Gulf of Mannar, Palk Bay (4); Mozambique (5); Minicoy (6); Seychelles (7) Maldives rock Kenia (1); Palk Bay, Mannar (2), shells Andamans, Java, Maldives (1), Madagascar (2); Gulf of Mannar, Palk Bay (3); Mozambique (4) calcareous algae Andamans, Gulf of Mannar (1, 2); Palk Bay (2) calcareous algae Ceylon (1); Gulf of Mannar, Palk Bay (2) calcareous algae, Mannar (1, 2); Palk Bay (2) shell, stony Andaman (1); Mahe Is., Red Sea (2); Gulf of Mannar s Palk Bay (3) shells Ceylon Mergui (1); Bay of Bengala (2) Mahe Is. calcareous algae Gulf of Mannar (1, 2); Palk Bay, Ceylon (2) rock Mannar, Palk Bay Mahe Is. (1); Maldives (2) not known Mannar, Palk Bay Minicoy Maldives, shell Mahe s. (1); Mozambique (2); Gulf of Mannar, Palk Bay (3) Annandale, 1915a (1); Vacelet et al, 1976 (2); Thomas, 1979b (3); Thomas, 1979a (4); present work (5) Keller, 1891 present work Annandale, 1915a (1); Levi, 1958 (2); Thomas, 1972 (2); present work (3) Annandale, 1915a Ridley, 1884 (1); Vacelet et ai, 1976 (2); present work (3) present work Annandale, 1915a (1); Thomas, 1981 (2); Thomas, 1986 (3) Annandale, 1915a (1); Thomas, 1975 (2); Thomas, 1981 (3); present work (4) Annandale, 1915a (1); Levi, 1958 (2); Thomas, 1975 (3); Thomas, 1979b (4); Thomas, 1979a (5); Thomas, 1980 (6); present work (7) present work Thomas, 1976b (1); Thomas, 1979b (2) Annandale, 1915 (1); Vacelet et al., 1976 (2); Thomas, 1979b (3); Thomas, 1979a (4) Annandale, 1915b (1); Thomas, 1972 (2) Carter, 1880 (1); Thomas, 1972 Carter, 1880 (1); Thomas, 1972 (2) Annandale, 1915a (1); Thomas, 1973 (2); Thomas, 1986 (3) Annandale, 1915a Annandale, 1915a (1); Thomas, 1979b (2) Thomas, 1981 Carter, 1880 (1); Thomas, 1979b (2) Thomas, 1979b, 1986 Thomas, 1981 (1); present work (2) Thomas, 1986 Thomas, 1989 present work Thomas, 1973 (D; Thomas, 1979a (2); Thomas, 1986 (3) «O O en TI g S z o o i The numbers in bracket, reported in the column of the geographic distribution, indicate the relative references.

206 B. CALCINAI, C. CERRANO, M. SARÀ, G. BAVESTREIXO TAXONOMIC ACCOUNTS Rosell & Uriz (1997) published a revision of the excavating Hadromerida. In the following systematic account, some of their results were followed for the genus identification. This collection is formed of 16 species (belonging to five families and seven genera), 11 of which already described for the Indian Ocean; two species: Cliona varians (known only for the Caribbean area) and Scantillel ta levispira (with an Atlanto-Mediterranean distribution) are new records for the Indo-Pacific. Three species described for the Maldives {Cliona nodulosa, Cliona dichotoma and Aka maldiviensis) are new. In total, 14 species have been collected from the Maldives, six from the Seychelles; 10 are present only in the Maldives ÍDercitus plicatus, Cliona inconstans, C. nodulosa, C. dichotoma, C. ensifera, C. schmiditi, C. orientalis, Scantilletta levispira, Zyzzya fuliginosa, Aka maldiviensis), two only in the Seychelles (Pione vastifica and S. cunctatrix), four (C varians, C. cfr. celata, C. mucronata, P. carpenteri) are common to the two zones. Family PACHASTRELLIDAE Carter, 1875 Dercitus plicatus (Schmidt, 1868) Corticium plicatum Schmidt, 1868, p. 2. Samus simplex Carter, 1880, p. 60. Stoeba plicata Annandale, 1915b, p. 458. Halina plicata Thomas, 1972, p. 353. MD6, MD2, in Acropora sp. Kuda Rah, MD12, in Porites sp. Gangehi (Maldives, 1996); 5-15 m depth. The very small sub-spherical specimens (4-10 mm in diameter), which occupied single boring chambers in the internal part of the s, were visible only by breaking the. The dark brown surface of the sponges is smooth, or very hispid with few short digitate processes, sometimes penetrating the foramina of the chambers wall. The skeleton is formed by loose spicules (dichotriaenes, calthrops) and microrhabds; digitate processes are made by numerous microrhabds, densely arranged and rare macroscleres. Spicules - Dichotriaenes widely variable in size (Fig 1A-C); rhabdoma size: 41-102 x 8-15 pm, deuteroclades: 20-82 x 8-20 pm, protoclades: 12-25 x 8-20 pm. Rare chaltrops; rays size: 70-90 pm. Very numerous microrhabds (8-13 x 2-3 pm) in which the central part of Fig. 1 - Dercitus plicatus. A-C, dichotriaenes (scale bars, 35 pm); D-F, microrahbds (scale bar, 2 pm).

BORING SPONGES FROM THE INDIAN OCEAN 207 the spicule is smooth and the spinosity is concentrated at the extremity (Fig. 1D-F). Dercitus plicatus was found insinuating or encrusting by some authors (Topsent, 1895; Pulitzer-Finali, 1970; Boury-Esnault & Lopès, 1985); Thomas (1972, 1979b) recognized the boring habit of the species. The digitate processes of the sponges inserted inside the foramina of the chambers, the presence of numerous chips in the sponge tissue, and the absence of foreign spicules induce us to agree with Thomas' conclusions. Atlantic Ocean, Mediterranean, Indian Ocean, Australian region. Family SPIRASTRELLIDAE Ridley & Dendy, 1886 Spirastrella cunctatrix Schimdt, 1868 Spirastrella cunctatrix Schmidt, 1868, p. 17. Spirastrella decumbens Ridley, 1884, p. 470. Sey6, in fragments of dead Porites sp., Seychelles, 1997; about 10 m depth. The presence of the sponge inside the is shown by circular, brown papillae (6-1 mm) emerging from line algae growing on the. The internal part of the is occupied by boring chambers variable in shape. Dried specimens are brown. Spicules - In general straight tylostyles, (225-324 x 7 pm) with circular or ovoid heads (Fig. 2A). The microscleres are thick spirasters with strong and long spines (40-42 x 30 pm, spines included) and numerous straight or bent spirasters, very variable in shape (8-20 pm, Fig. 2B). This species, characterized by great variability in shape and size of spirasters, is very close to S. decumbens described by Ridley (1884). In fact, many authors (Topsent, 1918; Levi, 1958; 1965; Vacelet et al., 1976) considered the two species synonymous. Mediterranean, Atlantic Ocean, Indo-Pacific. Family CLIONIDAE Gray, 1867 Cliona nodulosa sp. n. Fig. 2 - Spirastrella cunctatrix. A, tylostyle; B, spirasters (scale bar, 25 pm). Holotype MSNG 50029 in Porites sp., Kuda Rah (Maldives, 1998); paratypes MD7, MD4 in the authors' collection. From surface to 5 m depth. The sponge attacks massive s, producing large (10 x 8-17 mm), irregular, deep cavities. These boring chambers are 2 cm in depth in the substratum and are connected with the surface by long papular canals. Both papillae and chambers are dark-yellow, brown. Papillae are circular, small (1-3 mm) and do not fuse. Spicules - Straight tylostyles (310-488 x 9 pm) are irregularly arranged in the choanosome; in the papillae, they are arranged in a palisade; the tylostyles have distinct, sometimes trilobate heads (9.2-16 x 12-20 pm); a second evident swelling is frequently located along the shaft (Figs 3A, 4B, C). Tylostyle swellings are not smooth but are covered by small pronounced outgrowths (Fig. 3A). Rare, nodulose amphiasters are straight or bent and with small spherical or conical knobs (22-30 x 10 pm); they occur only in the papillae (Figs 3B, C, 4A). and etymology This species is named for its characteristic nodulose amphiasters, which are similar to one kind of Cliothosa hancocki amphiasters. Our species differs from this last

208 B. CALCINAI, C. CERRANO, M. SARA, G. BAVESTRELLO Fig. 3 - A, tylostyles with the characteristic swelling of Cliona nodulosa n. sp. (scale bar, 24 vim); B, C, nodulose amphiasters of C. nodulosa n. sp. (scale bar, 6 pm); D, ensiform tylostyle of Cliona dichotoma n. sp. (scale bar, 18 pm); E-H, ramose amphiasters with pointed rays (scale bar, 3 pm). by having only one type of nodulose amphiasters, which are very abundant and more irregular than those of Cliothosa hancocki. Cliona dichotoma sp. n. Holotype MSNG 50030 in Tubastrea sp., paratypes MD12, MD21, in the authors' collection, Kuda Rah (Maldives, 1998), 12 m depth. This species occupied the internal part of the dead ramous and was visible only by breaking the. The chambers made by the sponge are small (1-3 mm), almost circular, and sometimes fused together. They are arranged in a row in the peripheral part of the axis. The colour of the sponge in the dry state is yellowbrown. Spicules - Short, straight or bent, ensiform tylostyles (139-200 x 12 pm), with spherical heads (11-13 x 14-15 pm) (Figs 3D, 5A, B). Ramose amphiasters (11-18 pm), with a short shaft and long, irregular, smooth rays which have pointed tips often branched dichotomically (Figs 3E-H, 5C). and etymology The species is characterized by its short, ensiform tylostyles and smooth, dichotomically branched amphiaster that gave it the name.

BORING SPONGES FROM THE INDIAN OCEAN 209 Cliona ensifera Solías, 1878 Cliona ensifera Solías, 1878, p. 61. (\ Fig. 4 - Cliona nodulosa n. sp. A, amphiasters; B, trilobate heads; C, tylostyles with the frequent, second swelling (scale bar, 25 pm). \ MD1, MD2, MD9, MD11, MD13, in branches of dead Acropora sp., Gangehi (Maldives); MD20 in Serpulorbis colubrinus Kuda Rah (Maldives), 5 m depth. This sponge, which looks pale brown or dark yellow, produces large holes (5-10 mm) and is clearly visible breaking the only, papillae being very small. It presents a confused skeleton arrangement formed by tylostyles and spirasters. Spicules - Ensiform curved tylostyles (234-315 x 8-14 um), thick with a short and sharp tip, rarely blunt, and a spherical, sometimes mucronate head (Fig. 6A). Some thin and fusiform tylostyles are present (width 5 um). Spiny spirasters (33-90 x 3 um) with two or three bends and conical spines (Fig. 6B-C). Indian Ocean. Cliona schmidti (Ridley, I88i) * Vioa schmidtii Ridley, 1881, p. 130. Cliona schmidti Topsent, 1900, p. 77. Cliona schmidti Pang, 1973, P- 8. Fig. 5 - Cliona dichotoma n. sp. A, tylostyle; B, heads of tylostyles; C, ramose amphiasters (scale bar, 50 pm). MD8, MD9, MD11, in Pontes sp., MD20 in Serpulorbis colubrinus, Kuda Rah (Maldives, 1998), 5 m depth. Fig. 6 - Cliona ensifera. A, ensiform tylostyle (scale bar, 85 pm); B, C, spirasters with conical spines (scale bar, 6 pm).

210 B. CALCINAI, C. CERRANO, M. SARA, G. BAVESTRELLO J I Fig. 7 - Cliona schmidti. A, tylostyle; B, spirasters of the two kinds (scale bars, 50 urn). This species is easily recognized by its typically purple colour, which remains in dry preserved specimens. Very small circular papillae (up to 0.5 mm), which do not fuse together, are produced by this sponge in massive s and in the mollusc. In the, C. schmidti, it produces a short papular channel (4-6 mm) connecting groups of small chambers (1-5 mm). Spicules - Straight tylostyles, some very thin and with a spherical head (244-418 X 4.7-8 pm, Fig. 7A). Spirasters of two categories: short and stout spirasters with prominent spines, often located on the extremities (10-40 x 2-6 pm); longer, thin, curved spirasters with more delicate spines, spirally arranged (30-76 x 3-6 pm, Fig. 7B). Cosmopolitan. Cliona orientalis Thiele, 1900 Cliona Orientalis Thiele, 1900, p. 71. Cliona orientalis Annandale, 1915a, p. 13- MD4, MD15, some pieces of dead Acropora sp., Gangehi (Maldives), 5-12 m depth. Fig. 8 - A, boring chamber of Cliona orientalis (scale bar, 4 mm); B-D, spirasters of C. orientalis (scale bars, 4 pm); E, Cliona inconstans in its habitat (scale bar, 1 cm); F, G, curved, spiny spirasters (scale bar, 2 pm).

BORING SPONGES FROM THE INDIAN OCEAN 211 o are on one side of the spicules and not spirally arranged. The presence of frequent c-shaped spirasters, also reported by Thomas (1979b), may be useful element for distinguishing these two species. Indian Ocean, Red Sea, Australian region. Cliona inconstans (Dendy, 1887) Suberites inconstans Dendy, 1887, pp. 154-157. Spirastrella inconstans Thiele, 1899, p. 10. Cliona inconstans Rosell & Uriz, 1997, p. 362. MD 7b, MD 84, on sand, 20 m depth, Gangehi (Maldives, 1996). Fig. 9 - Cliona inconstans. A, tylostyle; B, heads of tylostyles; C, spirasters (scale bars, 50 pm). This sponge infested the internal part of s (Fig. 8A), producing large cavities connected with the surface through papular channels (2-3-5 x 1.5-2 mm). Chambers are irregular in shape and 4 x 20 mm wide. On the surface, a few, yellow, almost circular papillae are visible (1.5-2 mm). The dried sponge is pale brown, yellow. Spicules - Straight, or slightly curved tylostyles (211-282 x 3-6 pm) with rounded (stylote), trilobate, or spherical heads. Numerous spiny spirasters (Fig. 8B-D) with short thorns (5.8-26 x pm); the spirasters are variable in shape, curved with two or more bends, almost straight, or c-shaped with spines arranged on one side of the spicule. As Annandale (1915a) reported this species is close to C. viridis, from which it may be distinguished by the different position of the spines on the spirasters, that This is a dark brown massive sponge which emerges from the sand through cylindrical, digitate conules (20-40 mm high), often connected under the sand (Fig. 8E). The sand incorporated by the sponge shows signs of bioerosion. Oscules are located at the top of the dark brown conules. Dried specimens are brown-dark yellow. The skeleton, constituted by tylostyles, is very confused. Spicules - Slightly curved tylostyles (452-656 pm) with sharp, often stepped, ends (Fig. 9A). Head spherical, often rounded (stylote) or with some modifications (Fig. 9B). Abundant spiny and thin spirasters (4.6-20 x 2-4 pm), straight or with two or three bends and with short but evident spines, some similar to anthosigmas (Figs 8F, G, 9O. Many authors pointed out the great variability of this species, considered in the past as belonging to the genus Spirastrella, and the taxonomic confusion surrounding it (Vacelet & Vasseur, 1971; Vacelet et al., 1976). Our specimens are in agreement with Thomas' description (1973), both on account of the sponge shape, which is tubular, digitate or globular, and grows deeply rooted in the sediment, and of the spicular complement characterized by the spirasters, which are often similar to anthosigmas. Red Sea, Indian Ocean, Australian region, Pacific Ocean. Cliona varians (Duchassaing & Michelotti, 1864) Thalysias varians Duchassaing & Michelotti, 1864, p. 86. Suberites coronarius Carter (not Carter, 1887, p. 74), 1882, p. 352.

212 B. CALCINAI, C. CERRANO, M. SARÀ, G. BAVESTRELLO Fig. 10 - Cliona varians. A, heads of tylostyles (scale bar, 5 pm); C-E, anthosigmas (scale bar, 4.5 pm). Anthosigmella varians de Laubenfels, 1936, p. 143. Cliona varians Rützler, 1990, p. 460. MD3, MD4, MD10, MD11 in dead branches of Acropora sp., Gangehi (Maldives, 1996); Seyll, Seyl4 in Millepora sp., Seychelles, 1997, about 5 m depth. Dried specimens are brownish or dark yellow; circular papillae of the sponge are evident on the surface (1-4 mm). In the the sponge makes irregular hollows or cavities, the size and shape of which are variable. The chambers produced in the Seychelles s seem to be larger (more than 1.5 cm) than those of the specimens from the Maldives (from less than 1 mm up to 5 mm). The skeleton is formed by a confused arrangement of tylostyles. In the papular region, these tend to join together in bundles. Spicules - Straight tylostyles tapering gradually toward the pointed and long end (240-377 X 4.9-6.6 pm). Many thin tylostyles, often bent with an evident spherical head, are present. The head region can be quite spherical, ovoid or elongate (Fig. 10A, B). Abundant c-shaped, regular anthosigmas (16.5-24.7 x 2.5-3 um, Fig. 10D), displaying groups of branched spines regularly arranged along the convex side and in the extremities. A few can be almost straight, and in this case the spines are located on the spicule tops (Fig. IOC, E). Cliona varians is one of the most widespread boring species in our collections. The specimens from the Maldives and the Seychelles are all boring forms with papillae reaching the surface. Pang (1973) described encrusting specimens, but she was not sure of their boring capacity. Vincente (1978) recognized the boring activity of the sponge which 'destabilizes the substratum'. Wiedenmayer (1977) distinguished two different forms.- C. varians forma varians for the most common massive specimens and forma incrustans for the sponges with encrusting habit. Our boring specimens could represent a third form (an alpha stage) which temporally precedes beta (Pang, 1973; Vincente, 1978) and gamma stage. Wiedenmayer (1977) emphasized, (as other authors have done e.g., de Laubenfels, 1953, Hechtel, 1965), the great variability in the presence of real anthosigmas and

BORING SPONGES FROM THE INDIAN OCEAN 213 spirasters: e.g., one Bimini specimen, with an encrusting habit, had rare typical anthosigmas and numerous spined spirasters; two other dry specimens of the encrusting form of Turtle Rock had normal anthosigmas. Our boring samples show a spicular complement with regular anthosigmas and bent anthosigmas becoming spirasters; a similar situation was observed by Hechtel (1965) in Jamaican specimens with anthosigmas, a few spicules as amphiasters and a very few which tending toward the spirastral form. Straight spirasters (not rare in the present specimens) with spines or tubercles located on spicule tips were well illustrated by Topsent (1918). No significant differences between the microscleres of our Indian Ocean samples of Cliona varians and of others collected in the Caribbean were found: in the Caribbean sample, microscleres were not as abundant as in the Indian Ocean samples, but real anthosigmas which tended to be straight or spiral form were present. Until now, C. varians is known only from the West Indies (Pang, 1973). The presence of a Caribbean species in the Indian Ocean is an interesting report; speculations to explain this particular distribution are discussed by Thomas (1983), who reported a list of 13 sponges distributed exclusively in the two areas, and Wiedenmayer (1974). In 1887, Carter reported Suberites coronarius (= C. varians) for the Mergui archipelagos (Indian Ocean), but these samples were considered by Thiele (1900) as Cliona orientalis (see Pang, 1973). Our specimens are therefore the first, sure record of this species for the Indian Ocean. Atlantic, Indian Ocean. Cliona cfr. celata Grant, 1826 Cliona celata Grant, 1826. Cliona warreni Carter, 1881, p. 563; see Topsent, 1891, p. 563. Sey7 in a dead shell of Pleuroploca gigantea, 15 m; Sey8, Sey9 in dead (Porites sp. and Acropora sp.), Seychelles, 1997. This very destructive sponge was observed in numerous samples; it is dark-brown in the dried state, and produces big galleries and chambers (up to 10 mm) which occupy the whole. In the shell of P. gigantea, the sponge has produced holes arranged in rows along the shell; being located in a reduced space, these chambers are smaller (2.5-4 x 3-6 mm). Papillae are circular, numerous and variable in size and shape (from 1 mm up to 7 mm); dark yellow in colour and dark brown in the dried specimens, they often tend to fuse together; in this case, the sponge grows over the substrate, covering it. In the shell, the sponge produces numerous, very regular, circular papillae (0.5-2 mm). Spicules - Straight tylostyles, with sharp tips and oval head sometimes with an apical process (246-393 x 2.3-4.6 pm). Numerous thin, flexuous tylostyles are present. No microsclere were found. The systematic position of boring sponges lacking microscleres and with the presence only of tylostyles is problematic. Several of these specimens have been considered synonymous whit Cliona celata, or erected arbitrarily to new species rank. We follow Thomas (1979b), who reported C. celata in some samples of shells collected in the Indian Seas since his description of the boring pattern and of the spicules shape and size overlaps those of our samples. Cosmopolitan. Cliona mucronata Solías, 1878 Cliona mucronata Solías, 1878, p. 54. Only some small fragments in massive s or in fragments of branched s, mixed to other species, Maldives and Seychelles. Pale yellow sponge occupying small holes; small fragments of skeleton, separating two adjacent chambers, show a characteristic circular framework in which mucronate tylostyles are closely arranged with their tips converging in the centre (Fig. 11 A). n N Fig. 11 - Cliona mucronata. A, circular framework of the mucronate tylostyles (scale bar, 100 um); B, mucronate tylostyles (scale bar, 25 pm).

214 B. CALCINAI, C. CERRANO, M. SARÀ, G. BAVESTRELLO Spicules - Stout mucronate tylostyles (64-82.5 x 11.5 um) often, but not always, with rounded end, trilobate and spherical, evident head (6.6-14 x 8-11.5 urn, Fig. 11B). Not very numerous straight, or slightly curved tylostyles (109-132 x 3 pm). Numerous thin and short tylostyles, probably young spicules are present (59 x 3-5 pm on average). Indian Ocean, Australian region. Pione carpenteri (Hancock, 1867) Cliona carpenteri Hancock, 18Ó7, p. 241. Cliona bacillifera Carter, 1887, p. 76. Pione carpenteri Rosell & Uriz, 1997, p. 362. Seyl, Sey4, in valves of Hippopus sp., Seychelles, 1997; MD5, MD8, in branches of Acropora sp., Millepora sp., Maldives 1996, 5 m depth. Fig. 12 - A-C, spiny oxeas of Pione carpenteri and its magnification (A, B scale bars, 10 pm; C, 2,7 pm); D, E, straight spiny microrhabds of P. carpenteri (scale bar, 2 pm); F, oxea of Pione vastifica (scale bar, 9 pm); G, magnification of the oxea of P. vastifica (scale bar, 3 pm); H, I, curved microrhabds (scale bar, 2 pm).

BORING SPONGES FROM THE INDIAN OCEAN 215 The sponge produced small papillae (< 1 mm) on the surface of the substrata and subspherical chambers (1 mm on average), which occupied all the axis of the in the samples of Acropora sp. or which were located in the upper portion in Hippopus sp. The internal part was yellow, light-orange. Spicules - Straight tylostyles (174-305 x 6 pm) with an evident head; microspiny, fusiform straight, or slightly curved oxeas (66-127 X 3-6 pm, Fig. 12A-C). Straight and microspined microrhabds (11-18 x 3-4.5 pm, Fig. 12D, E). There is a number; of species characterized by tylostyles, spiny oxeas and microspiny rhabds. Following Rosell & Uriz (1997), these species belong to the genus Pione Gray, 1867. In regards to the two species of the genus described here, Topsent (1888, 1891) considered P. carpenteri a good species differing from P. vastifica in the straight and fusiform shape of its microrhabds. Riitzler & Stone (1986) observed in the holotype of P. carpenteri the presence of two classes of oxea that were not found in our samples. Scantilletta levispira (Topsent, 1898) Cliona levispira Topsent, 1898, p. 326. Cliona levispira Rosell & Uriz, 1997, p. 358. MD2, small fragments of this sponge in a branch {Acropora sp.) from Gangehi (Maldives, 1996), 5 m depth. Spicules - Macroscleres are straight or slightly curved oxeas, that sometimes present stepped tips (225-416 X 14 pm. Fig. 13A, B); thin anisoxeas (Fig. 13O, often with a rounded tip (60-132 x 4 pm). In our samples, microscleres consisted predominately by bent, smooth spirasters (Fig. 13D) (up to 4 bends) variable in width (25-51 x 3-6 pm). Rare very long spirasters (> 150 pm). Only in a single small fragment rare spiny spirasters (5-7 x 1.5-2 pm) were found (Fig. 13E). Atlantic Ocean, Indian Ocean, Australian region, Pacific Ocean. Pione vastifica (Hancock, 1849) Cliona vastifica Hancock, 1849. Cliona northumbrica Hancock, 1867, p. 237. Pione vastifica Rosell & Uriz, 1997, p. 362. SeylO, Seyl2, Seyl3, in samples of Acropora sp. and Millepora sp. in Seychelles, 1997, 15 m depth. This sponge which produced small subspherical chambers 1 mm up to 2 mm occupied the whole of the. On the surface, small, circular papillae, light orange in the dried state, were present (< 1 mm). Spicules - Straight tylostyles with evident, rounded heads (151-235 x 5-6 pm); microspiny curved oxeas (117-127 x 6-7.8 pm, Fig. 12F, G). Microrhabds, spiny, curved, S-shaped (9.4-18.8 x 3 pm, Fig. 12H, I). See P. carpenteri. \ I Cosmopolitan. Fig. 13 - Scantilletta levispira. A, oxea (scale bar, 50 pm); B, stepped tip of the oxea; C, thin anisoxeas; D, smooth spirasters; E, rare spiny spirasters (scale bar, 25 pm).

216 B. CALCINAI, C. CERRANO, M. SARÀ, G. BAVESTREUO Small, spiny spirasters described by Topsent (1898) and by Pouliquen (1972) are very rare. Topsent (1898) considered these spicules to be located in the papillae and very difficult to be found. Also Boury-Esnault et al. (1994) pointed out the rarity of this spicule. In our samples, these spicules are rare and only a few spirasters were observed. This species, known for the Mediterranean, Azores, and Caribbean, was in general collected from considerable depths (2000-3000 m), but it had also been recorded from reduced depths in caves. This is the first record for the Indian Ocean. Although Thomas (1973) drew two smooth spirasters that in the plate legend were attributed to «Cliona levispira?» but this species was neither described nor listed in the text. Mediterranean, Atlantic, Indian Ocean. Family IOPHONIDAE Burton, 1929 Zyzzya fuliginosa (Carter, 1879) Suberitesfuliginosus Carter, 1879, p. 347. For a review of Indo-Pacific Zyzzya species see van Soest et al, 1994, p. 166. Fig. 14 - Zyzzya fuliginosa. A, acanthostrongyle with verticillate spination; B, magnification of the tip; C, smooth curved acanthostrongyle, with numerous swellings; D, magnification of the microspined tip; E, curved and flexuous, thin tylotes; F, microspined tip of the tylote (A, C, E scale bars, 40 urn; B, D, F scale bars, 4 um).

BORING SPONGES FROM THE INDIAN OCEAN 217 MD2, in Acropora sp., MD9, MD10, Pontes sp. Gangehi (Maldives, 1996), 5 m depth. This sponge is very destructive, almost entirely occupying the s. Dried specimens are dark brown to black. Spherical boring chambers are very numerous and very small (about 1 mm); the connection of some of them can create elongate and straight hollows, or larger and subcircular chambers (4-6 mm). Numerous small and dark papillae (1-3 mm) are visible on the surface of the, boring also the encrusting line algae; the sponge tends to grow over the surface of its substratum by the merging of a few papillae. The boring habit of our specimen made the study of skeleton arrangement difficult. Spicules - Curved acanthostrongyles (205-270 X 9-28 um spines included) with spines more or less clustered in regular verticils arranged along the shaft (Fig. 14A). Rounded ends are also spined (Fig. 14B). Almost smooth curved acanthostrongyles (205-238 x 8-9 um) with numerous swellings along their surface; they are probably modified or vestigial acanthostrongyles with a very reduced spiny surface and microspined tips (Fig. 14C, D). Straight, curved or flexuous, thin tylotes (Fig. 14E) with microspined heads (Fig. 14F); tylote size: 213-307 x 1.6-4 urn. No microscleres. In a recent work, van Soest et al. (1994) considered Plocamia massalis Dendy, 1921, Damirina verticillata Burton, 1959, Paracornulum atoxa Vacelet et al., 1976 as synonyms of Zyzzya fuliginosa. They pointed out the great variability of the acanthostrongyle size and spinosity and the variable presence of microscleres (toxeas and chelas). Our specimens enlarge the range of variability of the species by having an acanthostrongyles length comparable with that of the tylotes while, generally, the length of tylotes is about twice that of the acanthostrogyles. Moreover, as already shown by Thomas (1981), our specimens are characterized by numerous smooth diactines. Indian Ocean. Family NIPHATIDAE van Soest, 1980 Aka maldiviensis sp. n. Holotype MSNG 50031, paratype MD100, in the authors' collection; Kuda Rah (Maldives, 1998), 12 m depth. Fig. 15 - Aka maldiviensis. A, triangular meshes of the ectosomal skeleton; B, dense oxeas tracts running in the inner surface of the tube (A, B scale bar, 100 pm); C, oxeas (scale bar, 54 um); D, magnification of the opened tip of an oxea (scale bar, 4.8 jam).

218 B. CALCINAI, C. CERRANO, M. SARA, G. BAVESTRELLO The specimens bore within a massive (Pontes sp.) and protrude out of the surface by whitish, pale yellow tubes (5-7 cm high and 8-11 mm in diameter) thinning at the apex. A regular network of triangular meshes builds the ectosomal reticulation (Fig. 15A). Groups of spicules (3-5 spicules wide) form the meshes. The inner surface of the tube is constituted by dense oxeas tracts (Fig. 15B) forming a net that, running to the surface, become more closed and tend to create spicular tufts. Near the base of the tube some spicular tracts, like roots made by oxeas, are visible. Spicules - Short, curved oxeas (127-143 x 4.6-7 urn) with an evident axial canal and opened tips (Fig. 15C, D). Neither styles, nor strongyles are present. From the Indian Ocean, three species of Aka were described: A. mucosa (Bergquist, 1965), known also for the Pacific Ocean, A. diagonoxea, Thomas, 1968, A. minuta Thomas, 1972. A. mucosa consists in blackish tubes 5-10 mm in diameter and 88 mm high; the spiculation consists of two categories of bigger oxeas with modifications to styles or strongyles. Aka diagonoxea is characterized by robust oxeas, often with two bends; the last species 04. minuta) shows an exclusive boring habitat (not protruding from the substrata) and spicules whose dimension and shape do not agree with those of the present sample. Other Aka species, known to produce more or less long chimneys out of the bored substratum, are the Caribbean A. siphona, (de Laubensels, 1949), A. brevitubulata (Pang, 1973), A. cachacrouense, and A. liphagum (Rützler, 1971). Aka siphona consists of tubes protruding from the sand; its spicules are larger and flexuous; A. brevitubulata is lemon yellow and has short tubes; A. cachacrouense forms irregular and dark grey epilithic pillows and its oxeas are larger than our samples; A. liphagum (forma typica, obruta and tubulosa) has two categories of oxeas which can be longer (up to 169.3 pm) than in Maldivian samples, and are frequently modified to styles or strongyles. The colour of these forms is yellow, which may change (in contact with air) to pinkish yellow. Epilithic parts are shorter (up to 50-65 cm) and of different shape (vase-or funnel-shaped). The nearest species to ours is A. amaycaense described by Pulitzer-Finali (1986) and collected in Jamaica at a depth of 40-45 m. The description of this species and the figures presented by the author (hollow, cream-white cylinders) and its spicules complement (shape and size) make it close to our sample collected in the Maldives. 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