Contributions to Marine Science 2012Contributions to Marine Science 2012: 49–56Date of Publication: 29 Sep.2012© National University of Singapore REDISCOVERY OF THE NEPTUNE’S CUP SPONGE IN SINGAPORE: CLIONA OR POTERION? Lim Swee-Cheng Tropical Marine Science Institute, National University of Singapore, 18 Kent Ridge Road, Singapore 119227. Email: [email protected] Karenne Tun DHI Water & Environment (S) Pte. Ltd., 1 Cleantech Loop, #03-05 CleanTech One, Singapore 637141. Email: [email protected] Eugene Goh DHI Water & Environment (S) Pte. Ltd., 1 Cleantech Loop, #03-05 CleanTech One, Singapore 637141. Email: [email protected] ABSTRACT. — The remarkable Cliona patera, or the famous Neptune’s Cup Sponge, captured the world’s imagination two centuries ago with its peculiar morphology and large size. Measuring more than 1 m in height and diameter, the cup-shaped sponge was also the first sponge species to be described from Singapore in 1820. However, the last record of this iconic sponge from Singapore waters was published in 1908, based on two specimens lodged in the Raffles Museum (collection date unknown). Until recently, Cliona patera was widely regarded as extinct due to overfishing until a single specimen was dredged up from the Gulf of Carpentaria (Australia) in 1990. In this paper, we report its presence from Singapore waters, the type locality, once again after more than a century. The generic position of this species in either Cliona or Poterion is uncertain. There are no records of live characters and little is known about the excavation characteristics of the sponge. This study describes C. patera in detail, including in situ observations for the first time. New excavation characters observed under the scanning electron microscope revealed that it is well placed in the genus Cliona, despite its distinct cup-shape and huge size compared to its congeners. KEY WORDS. — Neptune’s Cup Sponge; Cliona patera, Singapore, Porifera INTRODUCTION the second British Resident, Crawfurd (1830: 73): “… those gigantic sponges, which are peculiar to the coast ofCliona patera (as Spongia patera), the Neptune’s Cup Singapore, and which Europeans have called Neptuniansponge, was the first sponge species to be described from cups. The natives brought them to us in great numbers.” ThisSingapore (Hardwicke, 1820; Hardwicke, 1822; Low, 2012). remarkable sponge continued to fascinate even a hundredIt is arguably one of the most iconic and famous sponge years after it was discovered, featuring in a popular series ofspecies in the world. It is shaped like a cup or wineglass, W. D. and H. O. Wills Cigarette cards titled “Wonders of thegrowing to over a meter in height and diameter (Fig. 1) Sea” (Fig. 3) produced in 1928. However, C. patera was notand sufficiently large for a child to sit inside (Fig. 2). This recorded from Singapore waters since the late 19th century.remarkable sponge was much sought after by museums and Two specimens were dredged up in Singapore waters by theprivate collectors in the past, and most major natural history Eastern Telegraph Company, probably in the 1870s whenmuseums have at least one specimen of this iconic sponge. submarine cables were being laid across Singapore StraitThe Zoological Museum Amsterdam (ZMA) alone has over to connect Madras and Darwin. These were lodged in thetwenty specimens and the Natural History Museum in London Raffles Museum (currently Raffles Museum of Biodiversityhas some ten specimens. Research) and reported in Hanitsch (1908). Cliona patera has not been recorded from this region since the early 20thThis sponge was apparently common and abundant in century. The last record known was off the coast of BantamSingapore waters in the early 19th century. According to in West Java, Indonesia, where two specimens were collected 49 The Neptune’s Cup sponge in Singaporein 1907 (Vosmaer, 1908). Subsequently, there were no records (Hooper & van Soest, pers. comm.). It was only till 1990,of the sponge for a long time, leading biologists to speculate that one specimen was dredged up in the Australian watersthat the species may likely be extinct due to “over-fishing” for the first time at the Gulf of Carpentaria (Hooper & Ekins, Fig. 3. Cliona patera featured in a popular series of W.D. and H.O. Wills Cigarette cards titled “Wonders of the Sea” produced in 1928.Fig. 1. First drawing of C. patera (as Spongia patera) in Hardwicke(1822) description (the scale bar on the left is in inches).Fig. 2. A child bathing in a gigantic C. patera dried specimen. This Fig. 4. Cliona patera exhibit in Museo Civico Di Storiaphoto was taken in Indonesia, possibly in Sumatera, by F.C. Van Naturale (Giacomo Doria), Genoa labeled “Poterion poseidonisHeurn. This sponge is currently lodged in Netherlands Centre for (Singapore)”.Biodiversity (NCB), Naturalis (RMNH) in Leiden. 50 Contributions to Marine Science 2012pers. comm.). It must be noted that there has been a number TAXONOMYof sponge fauna surveys (Hooper et. al., 2000) in this regionbut none of these studies recorded C. patera. CLIONAIDAE d’Orbigny, 1851Two living specimens of C. patera were recently discovered Cliona Grant, 1826off the southern coast of Singapore, the Singapore Strait (Tun& Goh, 2011). This is especially amazing as almost the entire Cliona patera (Hardwicke, 1820)southern coastline of Singapore and many of the 50-odd (Figs. 5–8)islands in Singapore Strait were reclaimed and transformedin the past 50 years (Koninck et. al., 2008). Cliona patera is Material examined. — ZRC.POR. 0272. Singapore, Singaporeiconic but it is also one of the least known sponge species. Strait, Kias on 23 March 2011, 12m; ZRC.POR. 0273. Singapore,There were doubts as to whether it should be placed in the Singapore Strait, Kias on 18 April 2011, 10m.genus Cliona (van Soest, pers. comm.; see also Topsent, Studied for comparison: ZMA.POR.15267. Australia, Gulf1909) as it has a unique cup-shaped top or wineglass like of Carpentaria, W. of Kirk Reef. 31 November 1991; BMNHgrowth form like no other Cliona species. Most Cliona exist 10.10.24.1. Indonesia, Java Sea, off Batavia (now Jakarta).in the alpha and beta form, only a few exhibits the gamma Collection date unknown.form and become massive. However, these are amorphousor irregular in shape, unlike C. patera, which has an upright Description. — Sponge in gamma stage and free-living,stalk with rooting processes extending into the substratum. comprising an oval, shallow, concave disk supported by aA distinct cup forms the main body of the sponge and is stalk with a height of 26 cm (Fig. 5A). The stalk extendssupported underneath by a stalk. Its excavating ability is some 20 cm into substrate after which it branches into 4-6poorly documented, and only Vosmaer (1908) and Annandale rooting processes that extend a further 20 cm in depth into the(1915) reported their observations from dried material. substratum (Fig. 5B). Colour of living sponge was white to yellow. The upper part of the disk is mostly white, becomingA detailed description of C. patera from its type locality more yellowish towards the exterior and the undersides of(Singapore), together with new in situ and fresh excavation the disk and a more intense yellow towards the base of thecharacters are described in detail for the first time since the stalk. The consistency of the sponge is tough, particularly atlast account by Annandale (1915). The contentious issue ofthe genus placement of this species is also discussed in thelight of these new characters. MATERIAL AND METHODSThick sections of freshly collected material were made with asurgical blade, soaked in Xylene-Phenol (1:1) solution beforemounting on glass slides in DPX® and examined under thelight microscope at 100–400 X magnification. Dissociatedspicule suspensions were obtained by boiling in concentratednitric acid (siliceous spicules) and subsequently mounted onbrass stubs for examination under a JEOL scanning electronmicroscope (SEM). Residues of the spicule suspensions werealso mounted on glass slides in DPX® measurements ofspicule dimensions under the light microscope.Spicule size data are given as minimum–average–maximumbased on a minimum of 25 measurements for each spiculetype. Images of specimens, sections and SEM preparationswere obtained digitally. Systematic and the order of treatmentof the families and genera follow the Systema Porifera(Hooper & van Soest, 2002) and the World Porifera Database(van Soest et al., 2012). Fig. 5. Cliona patera. (A) in Singapore waters observed in March 2011; (B) Stalk and rooting processes in substratum exposed. 51 The Neptune’s Cup sponge in Singaporethe base of the stalk. The oval disk has a diameter of 38 cm tylostyles lying tangentially beneath (Fig. 7B). The corticalon the long side and 28 cm on the short side, with a depth of skeleton is an extension of the tylostyle tracts from the5 cm. Rim of disk is not level (Fig. 5). Thickness of disk is choanosomal skeleton. In the choanosomal skeleton, vagueapproximately 1.5 cm at the rim, becoming gradually thicker pauspicular tracts, loose spicules and collagen surroundedtowards the center. Surface of disk is covered with papillae. the thick and widely spaced tylostyle tracts (Fig. 7C).Only oscular papillae are present on the upper surface of thedisk, and most papillae are 1-2 mm apart from each other. Spicules. — Tylostyle megascleres are long, slim andThese oscular papillae are circular in outline with a diameter sometimes bent. Subterminal swellings are observed in someranging between 1-3 mm and are typically 0.1 mm in height tylostyles (Fig. 7D). Tylostyles in: 1) disk, 300–420.7–490 μm(Fig. 6A). Ostial papillae are found only on the underside of x 6–10.1–13 μm; 2) stalk, 205–377.4–480 μm x 8–11.5–13the disk and cover the entire undersurface of the disk. These μm; 3) rooting process, 210–358.4–480 μm x 7.5–10.3–13ostial papillae are typically 1-2 mm apart from each other, μm. Microscleres absent.and they extend downward to the upper part of the stalk.The ostial papillae are 3-5 mm in diameter and 2-3 mm in Ecology. — The two Cliona patera in this study were foundheight (Fig. 6B). The stalk is 13 cm in height, cylindrical in at 10-12 m depth, about 10 m apart from each other, andshape, with a diameter of 4 cm at the middle of the stalk. were in an area subjected to strong tidal currents of up toThe rooting processes are about 1 cm in diameter, and these 3 knots. The sponge stands upright with the disk facing up,are filled with materials from the surrounding substratum, supported by a stalk. The main body was firmly anchoredconsisting mainly of sand, mollusc shells, coral rubble and by rooting processes and basal part of the stalk was buriedforaminifera. in the substratum.Skeleton. — Consists of a thick cortical skeleton, 2.2 mm The rooting processes contain a high composition of thein width, with numerous thick branching and anastomosing surrounding substratum, bound together firmly by spongespicule tracts (1-1.5 mm) (Fig. 7A). The ectosomal skeleton materials, typically 0.5–1 mm in width, in between orcomprises of dense tylostyle palisade supported by dense enveloping the substratum materials. The substrata comprised of intact and broken mollusc shells, coral rubble, sand, and foraminifera. All calcareous material extracted from one piece of rooting process at the basal end, about 10 × 1 × 1 cm in size, were examined under a microscope but none exhibited the typical three dimensional network of excavation chambers below the substrate surface associated with Cliona species in the alpha and beta stage (Rützler, 2002). However, these calcareous materials were not large, ranging from 2–20 mm in width. Ten pieces of these smaller calcareous materials were randomly selected for examination under the SEM (Fig. 8). Seven pieces exhibited fresh excavated marks and slits on the surface (Fig. 8A & B). Excavated chip marks were irregularly circular, their diameters ranging between 15 and 30 μm. The slits were approximately 2 μm in diameter (Fig. 8C) and varied widely in length. Processes from etching cells (Rützler & Rieger, 1973; Pomponi, 1979; Schonberg, 2008) were observed in the slits (Fig. 8C).Fig. 6. Cliona patera. (A) Oscular papillae on the top surface of Distribution. — Only two Cliona patera individuals weredisk. . Scale bar = 1 cm; (B) Ostial papillae on the under side recorded in this study. Historical records have shown, sincesurface of disk. Scale bar = 1 cm. the publication of C. patera species, that they occurred in the Malacca Strait (off Sumatra), South China Sea (from Gulf of Thailand, Singapore Strait, Java Sea off Java) and Gulf of Carpenteria, off Australia (Hardwicke, 1820; Schlegel, 1867; Harting, 1870; Vosmaer, 1908; Sawangwong et al., 2008; van Soest et. al., 2012). However, many specimens lodged in natural history museums around the world do not have locality information and collection date (Harting, 1870). For example, the former Zoological Museum Amsterdam (ZMA, now merged with the National Centre for Biodiversity (NBC), Naturalis) in the Netherlands, has some 20 whole dried specimens, and the Natural History Museum (NHM) in London with some ten whole specimens, but most of them lack locality and collection information. Interestingly, a C. patera labeled “Poterion poseidonis (Singapore)” is being 52 Contributions to Marine Science 2012exhibited in Museo Civico Di Storia Naturale (Giacomo (Koninck et. al., 2008). All previous marine biodiversityDoria) in Genoa but no information of collection date and surveys carried out in the past did not report the presencespecific locality (Fig. 4). of C. patera despite its large size (Chuang, 1961, 1973, 1977; Chou and Wong, 1985; Hooper et al., 2000; Lim et.In 1974–76, surveys of fishes in Java and South China Sea by al., 2008; De Voogd & Cleary, 2009; Lim et. al., 2012). ThePauly et al. (1996) revealed large mushroom-shaped sponges discovery and the presence of two young C. patera reported(tentatively identified as Poterion nautilus) in almost every in this study may be indicative that more could present withinhaul. The P. nautilus looks very much like C. patera from the area, but more importantly, points to the possibility ofa photo (Fig. 9) taken by Pauly. However, there is no way adult populations present within Singapore’s coastal waters.to verify the identity of P. nautilaus as no specimen was It alludes to the presence of pockets of favorable habitatscollected during that survey. with suitable environmental conditions that can serve as refugia for the continued recruitment and survival of thisRemarks. — The rediscovery of the Cliona patera in rare sponge species.Singapore after more than a century is especially significantconsidering the extensive modification of its coastline. The two C. patera in this study, unlike, all those documentedSingapore has developed rapidly since its founding by Sir in the past (Hardwicke, 1822; Crawfurd, 1830; Schlegel,Stamford Raffles in 1819, transforming the quiet island 1858; Harting, 1870; Vosmaer, 1908; Topsent, 1909) did notto a busy port in a few years. Now, Singapore is a highly possess the cup or wineglass shaped main body characteristicdeveloped and urbanized city country, and have one of the of this species. They are probably the smallest and youngestbusiest ports in the world. The coast where C. patera used specimens documented although there is no doubt that theto be found in abundance (Hardwicke, 1822; Crawfurd, two specimens are indeed C. patera. They have similar1830) has been greatly transformed. Almost the entire south spiculation and skeletal structure as C. patera as indicatedcoastline of mainland Singapore and the 50-odd islands in the literature (Harting, 1870) and material from both ZMAhave been reclaimed and developed in the past 50 years and NHM show the distinct stalk and basal rooting processesFig. 7. Cliona patera. A. Skeletal cross-section of disk. Scale bar = 3 mm; B. Tylostyle tract extending to the cortex. At the surface is apalisade of tylostyles with the needle end of tylostyle facing out. Scale bar = 200 μm; C. Vague paucispicular tylostyle tracts in spacebetween the thick spicule tracts. Scale bar = 200 μm; D. Tylostyles. Scale bar = 100 μm. 53 The Neptune’s Cup sponge in Singaporeof C. patera that are also present in the specimens examined. sponge in the substratum. Only a few clionaids have beenThe most closely related species, C. celata, do not exhibit reported in gamma stage, e.g., Cliona celata Grant, 1826,stalk and rooting processes (Rützler, 2002; Rosell & Uriz, C. viridis and C. californiana de Laubenfels, 1932. They are2002; Schönberg et al., 2006). just massive (reaching one meter in width) and amorphous but nothing remotely as structured as C. patera. Beside theirCliona patera has the most advanced gamma form amongst distinct form, C. patera can be distinguished from most of theall clionaids in having a cup or wineglass shaped main body 81 valid Cliona species worldwide (van Soest et al., 2012)supported by a stalk with rooting processes anchoring the in having only tylostyles megascleres and the absence of microscleres. Only 16 species have been reported to contain only tylostyles (Schönberg et. al., 2006), and only three species of these exhibit gamma form, namely C. arenosa (Schmidt, 1870), C. californiana and C. celata. Cliona patera can distinguished from them in having larger tylostyles and thicker tylostyle tract (750–1000 μm), in addition to the unique gamma form. Vosmaer (1908) was the first to report that C. patera (as Poterion patera) was a boring sponge based on a Voluta (neogastropod) shell with numerous holes “found between the roots”, “dried sponge substance in the interior of Voluta proved that the spiculation closely resembles that of the Poterion”. Annandale (1915) reported “…at least one gastropod shell, which was extracted from the centre of the basal portion of large specimen, is wholly permeated and nearly destroyedFig. 8. Etch marks made by Cliona patera on gastropod shells. Fig. 9. A “Poterion nautilus” (at the top right) trawled up from one(A) Gastropod shell with etch marks made by rooting process of Mutiara surveys in Java and Southern South China Sea betweenof sponge. Scale bar = 200 μm; (B) Etch marks and slits on a 1974-1976. Specific collection date and locality not recorded. Photogastropod shell. Scale bar = 20 μm; (C) Processes of etching cells used with permission from Gustav Pauly.in slit. Scale bar = 2.5 μm. 54 Contributions to Marine Science 2012by excavations filled with sponge substance”. Both of them ACKNOWLEDGEMENTSobserved the typical cavities made by clionaids in alpha andbeta stages, which are visible to the naked eye. However, such We thank the two anonymous reviewers for their constructivecavities were not observed in all calcareous material (over comments and Dr Tan Koh Siang, the editor for this volume,100 pieces, ranging from 2-20 mm in width) extracted from for the valuable assistance given to us. Dive trips were madea piece of basal rooting process (10×1×1 cm in size) from possible from funds provided by National Biodiversitythe Singapore material. Instead, freshly etch marks (15–30 Centre, National Parks Board. We are grateful to Ms Lindaμm), slits and etching cells processes in the slits (Rützler & Goh, Mr. Jeffrey Low, and Mr. Lionel Ng for valuableRieger, 1973; Pomponi, 1979) are reported for the first time assistance rendered in this study.in C. patera. All the seven etch calcareous material examinedunder SEM showed extensive chipped marks on the surface LITERATURE CITEDbut no cavity. Fragments of gamma stage C. celata have beenshown to be capable to etch and excavate holes on calcareous Annandale, N., 1915. Indian boring sponges of the family Clionidae.materials within two months (Hartman, 1958). Cliona patera Records of the Indian Museum, 1: 1–24.is observed to be able to etch calcareous materials but morestudies are required to prove that C. patera really excavates Chou, L. M. & F. J. Wong, 1985. Reef community structure ofand make cavities like a typical clionaid. There are no Pulau Salu. In: Delesalle, B., R. Galzin & B. Salvat (eds.),reports of alpha and beta stage excavation characters of C. Proceedings of the Fifth Coral Reef Congress, Tahiti. Antennepatera. Interestingly, it might be difficult to distinguish the Museum-EPHE, Moorea. Pp. 285–290.alpha stage of C. celata from C. patera as both have similarspiculation and skeletal structure. Chuang, S. H., 1961. On Malayan Shores. Muwu Shosa, Singapore. 343 pp.Cliona patera was placed the genus Spongia when firstdescribed by Hardwicke (1820). Spongia was diagnosed by Chuang, S. H., 1973. Life of the seashore. In: Chuang, S. H. (ed.),Linnaeus in 1759 and used for the archetypal Greek bath Animal Life and Nature in Singapore. Singapore Universitysponge, Spongia officialis, in his classification. Over the Press, Singapore. Pp. 150–174.following 100 years or so, authors assigned almost all spongespecies to Spongia (Cook & Bergquist, 2002). Thirty-eight Chuang, S. H., 1977. 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Kluwer Academic/ Plenumfasciculated.” without reference to Schlegel (1858). Topsent Publishers, New York. Pp. 847–863.(1909) proposed the transfer of P. patera to the genusCliona after Vosmaer (1908) reported that C. patera is a Crawfurd, J., 1830. Journal of an Embassy from the Governor-boring sponge. He argued that the distinct cup-shaped top General of India to the courts of Siam and Cochin China. 2ndof C. patera does not warrant a separate genus. However, Edition. Hendry Colburn & Richard Bently, London. 475 pp.Vosmaer (1911) did not agree with Topsent (1909) andpreferred to retain Poterion as a genus. He also said “there De Voogd, N. J. & D. F. R. Cleary, 2009. Variation in spongeare sufficient anatomical grounds to keep Poterion and Cliona composition among Singapore reefs. The Raffles Bulletin ofapart” but did not provide supporting evidence. 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