Marine Ecology. ISSN 0173-9565 ORIGINAL ARTICLE Sponges boring into precious corals: an overview with description of a new species of Alectona (Demospongiae, Alectonidae) and a worldwide identification key for the genus Barbara Calcinai1, Carlo Cerrano2, Nozomu Iwasaki3 & Giorgio Bavestrello1 1 DiSMar, Università Politecnica delle Marche, Ancona, Italy 2 DipTeRis, Università di Genova, Genova, Italy 3 Usa Marine Biological Institute, Kochi University, Kochi, Japan Keywords Alectona; boring sponge; Japan Sea; new species; Paracorallium japonicum. Correspondence Giorgio Bavestrello, DiSMar, Università Politecnica delle Marche, Via Brecce Bianche, 60131 Ancona, Italy. E-mail: g.bavestrello@univpm.it Accepted: 24 April 2008 doi:10.1111/j.1439-0485.2008.00246.x Abstract Precious corals represent peculiar substrata for several species of boring sponges that exploit their carbonatic scleraxis, strongly decreasing their commercial value. Here we describe a new species of the genus Alectona from Japan recorded in a colony of Paracorallium japonicum (Kishinouye, 1903). The spicular complement of the new species consists of diactinal spicules covered by mushroom-like tubercles, often modified into styloid forms, and fusiform amphiasters with two or sometimes four verticils, each generally made up of six short tubercled rays. A complete survey of the literature on boring sponges recorded in precious corals in the Mediterranean Sea and Pacific Ocean indicates that the species of the family Alectonidae are the most strictly associated to this kind of substratum. Their world distribution, in fact, partially or totally overlap that of their coral hosts. Problem The genera Corallium and Paracorallium (Octocorallia, Corallidae) are known all over the world due to their heavy exploitation by the jewel industry. Considering their generally slow growth rate (Bramanti et al. 2005; Roark et al. 2006) there are several international projects devoted to their protection and management. The main areas where the precious coral fishery is strongly active are the Mediterranean for the red coral Corallium rubrum (Cicogna et al. 1999) and the Pacific Ocean for several other species (Parrish & Baco 2007; UICN 2007). Precious coral colonies are characterised by a scleraxis composed of Mg-calcite organised in an extremely compact structure of acicular to lamellar calcite crystals that grow centripetally, forming fan-like structures (Cortesogno et al. 1999). Often the collected colonies cannot be carved owing to the presence of boring sponges, strongly decreasing the commercial value of the coral. This is one of the most economically damaging defects to be found in commercial stocks (Liverino 1983). When occupied by boring sponges, the scleraxis of the precious coral shows a series of chambers interconnected by channels, giving rise to a ‘‘spongy’’ pattern in the carbonate. Information about which sponge species are involved in this boring activity, their relative destructive power, their distribution and their relationship with the affected coral colonies will help both the coral industry and coral scientists to draw together adequate measures for management and exploitation of this resource. The genus Alectona (Demospongiae) is characterised by the presence of diactinal, sometimes polyactinal, spicules, covered with spines and tubercles and fusiform or branched amphiasters (Rützler 2002). The pitting pattern of the excavated chambers, characterised by concentric Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 273 Calcinai, Cerrano, Iwasaki & Bavestrello Sponges boring into precious corals grooves overlapped by radiating scars, can be used as an additional feature in the diagnosis of this genus (Vacelet 1999; Calcinai et al. 2004a). This group of sponges shows a strong affinity for biogenic substrata. The first two species described for the genus, Alectona millari Carter, 1879 and Alectona wallichii Carter, 1874, were initially recorded as boring into calcareous rocks. However, the former was later observed in Mediterranean red coral (Maldonado 1992) and the latter in Pacific precious corals (Bavestrello et al. 1998). In 1899, Johnson recorded Alectona verticillata from Madeira excavating colonies of Corallium, later redescribed by Calcinai et al. (2004b) boring into Corallium elatius Ridley, 1882 from the Japan Sea. Alectona primitiva Topsent, 1932 was described in the mollusc Haliotis excavata Lamarck. Pang (1973) described Alectona jamaicensis excavating into a colony of Acropora and Bavestrello et al. (1998) recorded Alectona triradiata Lévi & Lévi, 1983 in a colony of C. elatius. The species Alectona sorrentini Bavestrello et al. (1998) and Alectona microspiculata Bavestrello et al. (1998) were recorded respectively in C. elatius and in the hydrocoral Distichopora sp. Vacelet (1999) described the deep Alectona mesatlantica excavating into a calcareous rock that was probably biogenic. The aim of this paper is to describe a new species of the genus Alectona recorded in a colony of Paracorallium japonicum (Kishinouye, 1903) from the Japan Sea. The new species is discussed in the framework of a complete bibliographic survey of all the boring species recorded in precious corals in the Mediterranean and the Pacific Ocean. Methods The material consists of portions of a branch of a colony of P. japonicum collected by Mr Yoshihiko Niiya off Ashizuri Cape (Kochi, Japan) at 120 m depth. Spicule preparations were made by dissolving sponge fragments in boiling nitric acid, and after rinsing in water and ethanol, mounting the spicules on microscope slides. The same preparation was followed for the scanning electron microscope observation. In this case, spicules were put on stubs and sputtered with gold. Ranges and mean spicule size (in brackets) were calculated from 15 measurements for each spicule type. The type material of the new species is deposited at the Museo Civico di Storia Naturale di Genova (MSNG). Results The examined sample of P. japonicum was collected alive, as suggested by the covering of intact coenosarc on the 274 mineral scleraxis. The coral is excavated by discrete, subspherical chambers (0.9–1.8 mm in diameter, 1.3 mm on average) present mainly in the central part of the scleraxis. The chambers communicate by numerous small foramina (0.2–0.4 mm, 0.3 mm on average). These chambers were produced by the boring activity of Holoxea excavans Calcinai et al. 2001, a common borer of Japanese precious corals. This attribution is due both to the large number of the spicules of this species (oxeas of two sizes) filling the boring chambers and to the pitting pattern with a fingerprint-like surface (Calcinai et al. 2004a). Some of the peripheral chambers produced by H. excavans were filled with densely packed groups of diactinal spicules that we attribute to a new species of the genus Alectona described below. Family Alectonidae Rosell, 1996 Genus Alectona Carter, 1879 Alectona sarai n. sp. Material examined Holotype MSNG 54326. Portions of P. japonicum, dry preserved and one spicule slide, 5 ⁄ 05 ⁄ 2007, Ashizuri Cape, Kochi, Japan; other material: two slides in the authors’ collection. Skeleton Proper erosion chambers of Alectona sarai n. sp. were not identified by SEM analysis. In fact, all the examined excavated portions showed scars with a fingerprint-like surface characteristic of Holoxea (Calcinai et al. 2004a), whereas a radial pattern overlapping deep concentric lines was not observed (Vacelet 1999; Calcinai et al. 2004a). The skeleton of the new species of Alectona is made up of: 1 Diactinal spicules (Fig. 1A–C), 175–250 lm long and 12.5–20 lm wide (207.5 ± 27.3 · 17 ± 3 lm); they are generally flexuous, sometimes with a central bend (Fig. 1B). The axial canal is often evident. The tips are sharp and often end with an apical single spine (Fig. 1A– C). These spicules are covered with long mushroomshaped tubercles 5–7.5 lm long, arranged in alternate rows (Fig. 1D). On a few spicules the tubercles are transformed into spines. Several spicules, about half of the total, have a rounded extremity and resemble stout papillose, sometimes bent, styles (Fig. 1E–H). 2 Fusiform, straight amphiasters with a long and generally blunt axis (Fig. 1I–M); 20–40 lm (27 ± 4.9 lm) long and 2.5–3.7 lm thick. In the middle of the axis, two Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd Calcinai, Cerrano, Iwasaki & Bavestrello Sponges boring into precious corals A B E I C F J D G K H L M Fig. 1. Spicules of Alectona sarai n. sp. (A–C) Diactinal tubercled spicules. (D) Magnification of the surface of a diactinal spicule showing the mushroom-shaped tubercles arranged in alternate rows. (E–G) Spicules modified into styles. (H) Magnification of the rounded extremity of a style. (I) Fusiform amphiaster with two verticils of six short tubercled rays. (J–L) Amphiasters with nodules arranged in two accessory distal verticils and scattered along the axis. (M) Amphiaster with a short axis ending with groups of spiny knobs. Scale bars: (A–C) 50 lm; (D,H–M) 10 lm; (E–G) 70 lm. verticils, each generally made up of six (sometimes fewer) short tubercled rays, are present (Fig. 1I). Numerous nodules are often arranged in two other accessory distal verticils or are scattered along the axis (Fig. 1J–L). Several of the latter spicules have one extremity of the axis which is shorter and thinner, ending with groups of spiny knobs (Fig. 1M). Etymology The species was named in memory of our mentor Prof. Michele Sarà. Taxonomic Remarks The species of the genus Alectona are characterised by amphiastrose microscleres which are very variable in shape: they include amphiasters with a small centre and slender long rays, amphiasters with a stout diactine axis (Rützler 2002) and nodular amphiasters with a short axis and microspinations mainly grouped at the extremities. We suggest the term ‘nodular amphiasters’ for the microscleres described as small amphiasters by Rützler (2002). They often have a thick axis and may resemble the nodular amphiasters typical of Thoosa (compare for Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 275 Calcinai, Cerrano, Iwasaki & Bavestrello Sponges boring into precious corals example Fig. 16, b page, 184 and Fig. 1, I page, 282 in Rützler 2002). We suggest the term ‘fusiform’ for the amphiasters with a stout diactine axis and two verticils of spiny knobs, and the term ‘oxyamphiasters’ for the third kind of microsclere with hispid long oxyform axis (Fig. 1, H page 282 in Rützler 2002). Alectona sarai n. sp. is characterised by short diactines, often transformed into styloid forms, with long flattened tubercles and by fusiform amphiasters with two or often four verticils of spined knobs which vary in shape. Six Alectona species have been recorded in the IndoPacific area (Table 1): Alectona microspiculata, Alectona primitiva, Alectona sorrentini, Alectona triradiata, Alectona verticillata and Alectona wallichii. The species most closely related to A. sarai n. sp. is A. wallichii, as the shape of the diactinal spicules is characterised by mushroomlike tubercles (Vacelet 1999), although it differs in the size of these spicules, which are longer in A. wallichii (Bavestrello et al. 1998; Vacelet 1999), and in the shape and arrangement of the tubercles, which are flat and closely spaced in A. wallichii. The latter species is also characterised by fusiform amphiasters with two regular verticils different in shape from those of A. sarai n. sp. Moreover, the smooth diactines regularly present in A. wallichii have never been observed in A. sarai n. sp. The other species known in the area are also different in the shape and size of the spicules (Table 1). Alectona microspiculata has short, sausage-shaped diactines (30– 60 · 7–10 lm) and two kinds of ramose and short amphiasters (nodular and oxyamphiasters). Alectona primitiva has longer, spined diactines (680 · 30 lm) and short amphiasters with tylote extremities (nodular amphiasters) (Vacelet & Vasseur 1971). Alectona sorrentini has diactines covered with short spines and spiny, ramose amphiasters (oxyamphiasters), different in shape from those of A. sarai n. sp. Alectona triradiata has spiny triactines and two kinds of non-fusiform amphiasters, very different in shape from those of A. sarai n. sp. Alectona verticillata is characterised by three kinds of amphiasters (nodular, fusiform and oxyamphiasters) and by the lack of diactinal spicules (Calcinai et al. 2004b). This new species is also similar to A. millari from the Mediterranean Sea and North Atlantic Ocean for the size of the diactinal spicules and fusiform amphiasters. Alectona sarai n. sp. differs for the presence of tubercles instead of long spines on the diactinal spicules and the presence of irregular amphiasters; moreover, we have not observed the modified diactines that are always present in A. millari. This new record brings the number of Alectona species to 10 (Table 1) and we propose the following key for their identification: 276 (1) Triactinal spicules Not as above (2) Diactinal spicules absent (no macroscleres) Diactinal spicules present (3) Diactinal spicules covered with mushroom-shaped tubercles Not as above (4) Diactinal spicules long (>300 lm), up to 800 lm; fusiform amphiasters with regular verticils; smooth diactines may be present Diactinal spicules short (<300 lm), irregular fusiform amphiasters, no smooth diactines (5) Very short diactinal spicules (<100 lm), sausage-shaped Not as above (6) Only fusiform amphiasters as microscleres Not as above (7) Fusiform amphiasters characterised by long rays bearing small spines at the top Not as above (8) Only nodular amphiasters Only oxyamphiasters (9) Long diactines (>400 lm), no polyactinal spicules, large pits (up to 100 lm) Shorter diactines (<400 lm), polyactinal spicules, small pits (about 50 lm) Alectona triradiata (2) Alectona verticillata (3) (4) (5) Alectona wallichii Alectona sarai n. sp. Alectona microspiculata (6) (7) (8) Alectona jamaicaensis (9) Alectona primitiva Alectona sorrentini Alectona mesatlantica Alectona millari Ecological Remarks In the Mediterranean, during a series of surveys conducted on different red coral populations, 14 boring species were identified (Melone 1965; Barletta & Vighi 1968; Corriero et al. 1988; Calcinai et al. 2002, 2004a, 2007). The same number was recorded during our studies (Bavestrello et al. 1995, 1996, 1998; Calcinai et al. 2001, 2002, 2004b) on the Pacific species of precious corals. Twenty six species were involved in boring activity in the precious corals, with only two species, Dotona pulchella Carter, 1880 and A. verticillata, being common to both geographic areas, and Delectona ciconiae Bavestrello, Calcinai & Sarà, 1996 being recorded only in the Mediterranean coral (Fig. 2, Table 2). There are eight genera involved: Holoxea, Alectona, Delectona, Thoosa, Spiroxya, Cliona, Dotona, Aka. The most represented genera are Alectona and Spiroxya (family Alectonidae), both with six species, whereas Cliona (family Clionaidae), which is very common in calcareous, organic substrata such as coral reefs, is poorly represented. There are two species of Alectona in the Mediterranean and five Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd Species Diactinal spicules (lm) Alectona jamaicaensis Alectona microspiculata Alectona mesatlantica a) 186–237 · 9.6–12 b) 186–287 · 6.1–8.5 30–60 · 7–10 Alectona millari 215–370 · 15–20 Alectona primitiva 170 · 8 Alectona sorrentini Alectona triradiata 148–239 · 14–18 Alectona wallichii a) 380–500 · 40–50 b) 280–490 · 10–28 Poliactinal Nodulose Fusiform spicules amphiasters amphiasters (lm) (lm) (lm) 31–74 · 1.4–4.3 7–10 · 1 410–530 · 21–28 127 · 6.3 12–14 · 5 23–37 49–58 15–35 7.5–13 32.5–70 · 3.7–7.5 20–40 · 2.5–3.7 Substratum Chambers Chamber size (mm) shape Depth (m) Ref. Jamaica Porites furcata 5.8–7.9 14 (7) Philippines Disticophora sp. not stated (1) Equatorial organic, up to 50 Mid-Atlantic calcareous rock Ridge Mediterranean, calcareous rock, North Atlantic, coral, Red Sea Corallium rubrum 10–70 · 1–5 7–10 · 2.5 Distribution 15–30 (rays) Australia Haliotis excavata (Cangaroo Island) · 1.5–3 Japan Corallium elatius ·2 New Caledonia, Corallium elatius; Japan calcareous conglomerate limestone rocks, Indian Ocean, South Corallium elatius Australia · 2.5 Mediterranean, Corallium elatius Japan Japan Paracorallium japonicum 0.6–0.8 spherical, ellipsoidal hemispherical single, 2030 subspherical, irregular 500–660 (3); 202 (8); 70–120 m (2); shallow water (9, 10) small, closed 3 (11) spaced (1) (1), (2), (3), (8), (9), (10) (1), (11) large, irregular 1–6 irregularly · 1.5–3.5 spherical, deep 5–6 large, subspherical, irregular 20 · 25 large, rounded not stated 290–350 (1) (1), (4) 10; 146–182 (1), (5) 195–300 (6) not stated not stated not stated (12) (1) Bavestrello et al. 1998; (2) Maldonado 1992; (3) Topsent, 1920; (4) Lévi & Lévi, 1983; (5) Vacelet 1999; (6) Calcinai et al. 2004b; (7) Pang 1973; (8) Carter, 1879; (9) Sarà, 1959; (10) Rützler, 1966; (11) Vacelet & Vasseur 1971; (12) present paper. 277 Sponges boring into precious corals Alectona sarai n. sp. 175–250 · 12.5–20 20–30 · 1 20–60 · 3–7 79–84.6 · 28 Alectona verticillata Oxy amphiasters (lm) Calcinai, Cerrano, Iwasaki & Bavestrello Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd Table 1. Principal features of the species of the genus Alectona (Bavestrello et al. 1998 mod.). Calcinai, Cerrano, Iwasaki & Bavestrello Sponges boring into precious corals Table 2. Boring sponge species recorded in species of the coral genus Corallium from the Mediterranean Sea and the Pacific Ocean. Species Fig. 2. Number of species per genus recorded in the precious corals from the Mediterranean Sea (white bars) and Pacific Ocean (grey bars). in the Pacific, and three species of Spiroxya both in the Mediterranean and in the Pacific Ocean (Table 2). Alectonidae (Alectona, Dotona, Delectona and Spiroxya) account for 14 species, suggesting a strong affinity of this family for the compact calcite of the precious coral scleraxis. The affinity of the species of this family for precious corals could be related to their typical bathophily. The deepest known boring sponges are Spiroxya levispira Topsent 1898 (2165 m, Topsent 1928) and Alectona mesatlantica, recorded at a depth of 2030 m (Vacelet 1999). Pang (1973) underlined that records for A. millari were generally for greater depths and that all the shallow-water records were from underwater cavities or shaded sites. This is also true for the shallow records of A. wallichii (Vacelet 1999). It is possible to hypothesise that species of this genus, together with the genus Spiroxya, are strongly stenoecious, limiting their presence to relatively stable environments such as marine caves and deep bottoms. Nevertheless, whereas Spiroxya is a genus with a strong affinity for precious corals (with six species out of nine recorded in coral colonies), Alectona is common in other organic carbonates. This is also indicated by the geographic distribution of these genera; Alectona has a circumtropical distribution and the genus Spiroxya has exactly the same Tethyan relict distribution as the coral genus Corallium (Calcinai et al. 2004b). Acknowledgements This paper is dedicated to Klaus Rützler to remember his fundamental contribution to the taxonomy and ecology of boring sponges both in the Mediterranean and in the tropics. He has highlighted, more than any other, the great 278 Genus Holoxea Holoxea furtiva Topsent, 1892 Holoxea excavans Calcinai et al., 2001 Genus Alectona Alectona millari Carter, 1879 Alectona wallichii Carter, 1874 Alectona triradiata Lévi & Lévi, 1983 Alectona verticillata (Johnson, 1899) Alectona sorrentini Bavestrello et al., 1998 Alectona sarai sp. n. Genus Delectona Delectona ciconiae Bavestrello et al., 1996 Genus Cliona Cliona janitrix Topsent, 1932 Cliona lobata Hancock, 1849 Cliona desimoni Bavestrello et al., 1995 Genus Spiroxya Spiroxya acus (Bavestrello et al., 1995) Spiroxya corallophila (Calcinai et al., 2002) Spiroxya heteroclita (Topsent, 1988) Spiroxya levispira (Topsent, 1898) Spiroxya macroxeata (Calcinai et al., 2001) Spiroxya sarai (Melone, 1965) Genus Thoosa Thoosa armata Topsent, 1888 Thoosa bulbosa Hancock, 1849 Thoosa midwayi Azzini et al., 2007 Thoosa mollis Volz, 1939 Genus Dotona Dotona pulchella Carter, 1880 Genus Aka Aka labyrinthica (Hancock, 1849) Aka coralliirubri Calcinai et al., 2007 Aka insidiosa (Johnson, 1899) Mediterranean Sea Pacific Ocean X X X X X X X X X X X X X X X X X X X X X X X X X X X X ecological importance of boring sponges, focusing the attention of new generations of researchers on this fascinating group. The authors wish to thank Mr Yoshihico Niiya, Kosaitsuno, Kochi, Japan for sampling the material. References Azzini F., Calcinai B., Iwasaki N., Bavestrello G. (2007) A new species of Thoosa (Demospongiae, Hadromerida) excavating precious coral Corallium sp. from Midway. Italian Journal of Zoology, 74(4), 405–408. Barletta G., Vighi M. (1968) Ricerche sul corallo rosso: 5 – poriferi perforanti. Rendiconti Istituto Lombardo Accademia di Scienze e Lettere, Milano. Scienze Biologiche e Mediche, 102, 145–159. Bavestrello G., Calcinai B., Sarà M. (1995) Two new species of Cliona (Porifera, Demospongiae) boring the scleraxis of Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd Calcinai, Cerrano, Iwasaki & Bavestrello Corallium elatius from the western Pacific. Bollettino di Zoologia, 62, 375–381. Bavestrello G., Calcinai B., Cerrano C., Pansini M., Sarà M. (1996) The taxonomic status of some Mediterranean clionids (Porifera, Demospongiae) according to morphological and genetic characters. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique (Ser. Biologie), Bruxelles, 66, 185–195. Bavestrello G., Calcinai B., Cerrano C., Sarà M. (1998) Alectona species from Western Pacific (Demospongiae: Clionidae). Journal of the Marine Biological Association of the United Kingdom, 78, 59–73. Bramanti L., Magagnini G., De Maio L., Santangelo G. (2005) Recruitment, early survival and growth of the Mediterranean red coral Corallium rubrum (L. 1758), a 4-year study. Journal of Experimental Marine Biology and Ecology, 314, 69–78. Calcinai B., Bavestrello G., Cerrano C., Sarà M. (2001) Boring sponges living into precious corals from the Pacific Ocean. Italian Journal of Zoology, 68, 153–160. Calcinai B., Cerrano C., Bavestrello G. (2002) A new species of Scantiletta (Demospongiae, Clionaidae) from the Mediterranean precious red coral with some remarks on the genus. Bulletin of Marine Science, 70, 919–926. Calcinai B., Bavestrello G., Cerrano C. (2004a) Bioerosion micro-patterns as diagnostic characteristics in boring sponges. In: Pansini M., Pronzato R., Bavestrello G., Manconi R. (Eds), Sponge Science in the New Millennium, Bollettino dei Musei e degli Istituti Biologici dell’Università di Genova, 68, 229–238. Calcinai B., Azzini F., Bavestrello G., Iwasaki N., Cerrano C. (2004b) Redescription of Alectona verticillata (Johnson) (Porifera, Alectonidae) boring into Japanese precious coral. Italian Journal of Zoology, 71, 337–339. Calcinai B., Cerrano C., Bavestrello G. (2007) Three new species and one re-description of Aka. Journal of the Marine Biological Association of the United Kingdom, 87, 1355–1365. Cicogna F., Bavestrello G., Cattaneo-Vietti R., Eds (1999). Red coral and other Mediterranean octocorals, biology and protection. In: Red coral in the Mediterranean Sea: Art, History and Science. Ministero per le Politiche Agricole, Alimentari e Forestali, Roma. Corriero G., Pansini M., Sarà M. (1988) Boring sponges (Demospongiae, Clionidae) perforating Corallium rubrum in the Mediterranean sea. Fao Fishery Reports, 413, 73–78. Cortesogno L., Gaggero L., Bavestrello G., Cerrano C., Cattaneo-Vietti R. (1999) Struttura, mineralogia, minerochimica e chimismo del corallo rosso. In: Cicogna F., Bavestrello G., Cattaneo-Vietti R. (Eds), Biologia e tutela del corallo rosso e di altri ottocoralli del Mediterraneo. Ministero per le Politiche Agricole, Almentari e Forestali, Roma: 83–97. Sponges boring into precious corals Liverino B. (1983) Il Corallo. Ed. Analisi, Bologna: 229 pp. Maldonado M. (1992) Demosponges from the red coral bottoms of the Alboran Sea. Journal of Natural History, 26, 1131–1161. Melone N. (1965) I poriferi associati a Corallium rubrum (L.) della Sardegna. Annali del Museo Civico di Storia Naturale Giacomo Doria Genova, 75, 344–358. Pang R.K. (1973) The systematics of some Jamaican excavating sponges (Porifera). Postilla of the Peabody Museum of Yale University, 161, 1–75. Parrish F.A., Baco A.R. (2007) State of deep coral ecosystems in the U.S. Pacific Islands Region: Hawaii and the U.S. Pacific Territories. In: Lumsden S.E., Hourigan T.F., Bruckner A.W., Dorr G. (Eds), The State of Deep Coral Ecosystems of the United States, Chapter 4. U.S. Dept. Commer., NOAA Tech. Memo CRCP – 3, Silver Spring, MD, USA: 365 pp. Roark E.B., Guilderson T.P., Dunbar R.B., Ingram L. (2006) Radiocarbon-based ages and growth rates of Hawaiian deep-sea corals. Marine Ecology Progress Series, 327, 1–14. Rützler K. (1966) Die Poriferen einer sorrentiner Höhle. Zoologischer Anzeiger, 176, 303–319. Rützler K. (2002) Family Clionaidae D’Orbigny, 1851. In: Hooper J.N.A., Van Soest R.W.M. (Eds), Systema Porifera: A Guide to the Classification of Sponges, Vol. 1. Kluwer Academic ⁄ Plenum Publishers, New York: 173–185. Sarà M. (1959) Specie nuove di Demospongiae provenienti da acque superficiali del Golfo di Policastro (Mar Tirreno). Annali dell’ istituto e Museo de Zoologia dell’ Università di Napoli, 11, 1–22. Topsent E. (1920) Caractères et affinities de Thoosa Hanc. et des Alectona Cart. Consideration sur leur germes à armure. Bulletin de la société Zoologique de France, 45, 88–97. Topsent E. (1928) Spongiaires de l’Atlantique et de la Méditerranée, provenant des croisières du prince Albert Ier de Monaco. Résultats des Campagnes Scientifique du Principe Albert Ier de Monaco, 74, 1–376. UICN (2007) Les Analyses UICN ⁄ TRAFFIC des propositions d’amendement aux Annexes de la CITES pour la Quatorzième session de la Conférence des Parties à la CITES. Available at: http://www.iucn.org/themes/ssc/our_work/wildlife_trade/citescop14/cop14analyses.htm. Vacelet J. (1999) Planktonic armoured propagules of the excavating sponge Alectona (Porifera: Demospongiae) are larvae: evidence from Alectona wallichii and A. mesatlantica sp. nov. Memories of the Queensland Museum, 44, 627–642. Vacelet J., Vasseur P. (1971) Èponges des récifs corallines de Tulear (Madagascar). Tethys (Suppl. 1), 51–126. Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 279