Demospongiae: Chondrosida: Chondrillidae) in Australia
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DOI: 10.18195/issn.0312-3162.24(4).2008.469-486 I\e(ords of tile Westcm AlIstmllilll /\:1/"C/1I1I 24: 469-"-4K6 (200K) Species of the sponge genus Chondrilla (Demospongiae: Chondrosida: Chondrillidae) in Australia 1 2 4 5 J. fromont , K.L. Usher , D.e. Sutton', S. Toze and J. Kuo 'Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 69K6, Australia :'CSIRO Land and Water, Private Bag no.5, Wembley, Western Australia 6913, Australia 'School of Biomedical, Biomolecular and Chemical Sciences, University of Western Australia, Crawley, Western Australia 6009, Australia 'CSIRO Water for a Healthy Country, Queensland Biosciences Precinct, St Lucia, Queensland 4067, Australia 'Centre for Microscopy, Characterisation and Analysis, University of Western Australia, Crawley, Western Australia 6009, Australia Abstract - A new sponge species, Cholldrilla linnaei, from Australia is described. Chondrilla alls/mliensis Carter 1873 and C. seclInda Lendenfeld 1885 are redescribed with reference to type and new material. The synonymy of C. corlicata and C. papillata with C. allstralicnsis is confirmed. Two species previously described from Australia, C. mix/a and C. nllclIla are discussed, with the conclusion that C. Ill/cilia is unlikely to occur in Australia. The descriptions of the species incorporate previous molecular results. Information about symbiont relationships and the biogeographical distribution of the species is discussed. Keywords: taxonomy, systematics, sponges, Porifera, Cholldrilla INTRODUCTION the presence of three species: C. IIl/slraliel1sis Approximately twenty species of Clzol1drilla and two unidentified species described here. A Schmidt 1862 appear to be valid worldwide specimen collected from Australia and previously (Boury-Esnault 2002). However, the morphology identified as C. Ill/Cl/la was examined and found and skeletal structures of this genus and its to be the new species described in this paper. sister genus (CllOl1drosia) are remarkably uniform, Although fieldwork was undertaken at Christmas with few characters useful for differentiation Island C. //lixla was not found. This study reports at the species level. Consequently at least two three valid species of CllOl1drilla from mainland species of Clzol1drilla have been considered to be Australia: C. IIl/stmlicl1sis, C. secl/l1da and C. lillllllei. globally widespread. The type species, C. Ill/Cl/la The occurrence of C. //lixta in Australian waters Schmidt 1862 was thought to have a cosmopolitan was not resolved. distribution, and C. al/stralieJlsis Carter 1873 Species of Clzolldrillll have been reported was reported with a widespread lndo-Pacific from shallow waters « 50 m depth) in tropical, distribution (Hooper and Wiedenmayer 1994). subtropical and temperate zones, but rarely in Recent genetic studies have reduced the known deeper waters (Boury-Esnault 2002). Many species distribution of C. Ill/Cl/la to the Mediterranean Sea are considered to be cryptic, occurring on vertical (Klautau et al. 1999). walls, at cave entrances, and under rocks (Boury- Six species of the genus Clzolldrilla have been Esnault 2002). In south Western Australia C. reported from Australian waters. These include IIl/stmliellsis is a major space occupant of many C. al/straliclIsis, C. secl/llda Lendenfeld 1885 temperate limestone reef habitats, and forms large and C. Ill/Cl/la from mainland Australia, and C. encrustations in shallow depths (1-20 m) in full //lixla Schulze 1877 from Christmas Island, an light and shaded environments (Usher et Ill. 2001). Australian Territory in the Indian Ocean. "rwo Individuals of this species have been reported to others, C papillllta Lendenfeld 1885 and C. corlicala provide refuge for a wide range of invertebrates Lendenfeld 1885, are regarded as synonyms of C. including brittle stars, molluscs and shrimp Ill/slmliellsis (Burton 1924). On the basis of DNA (Edgar 1997), and to be a food item for species sequence analyses of CllOlldrilla specimens from of the cowrie genus Zoila (Wilson and Clarkson Australian waters, Usher cl al. (2004a) suggested 2004). Clzolldrillll aff. ill/Cilia from Caribbean coral reefs have been found to be a preferred food of 470 J. Fromont, K.L. Usher, D.e. Sutton, S. Toze, J. Kuo the Hawksbill turtle (Meylan 1988) and some double-sided carbon tape attached to SEM stubs, fishes (Randall and Hartman 1968). dried at 70°C and sputter coated with gold prior Species of Chondrilla have been reported to to examination with a Philips SEM 505 or a Zeiss have abundant populations of symbiotic bacteria 1555 SUPRA Variable Pressure SEM operating at in their mesohyl (Boury-Esnault 2002), and C. 15 kY. Images were recorded electronically. allstraliensis has been found to have both bacteria The skeleton was prepared for examination by (Dey et al. 2004) and symbiotic cyanobacteria cutting a representative section at right angles (Usher et al. 2001, 2004c). Cyanobacterial symbionts to the surface of the sponge. The section was are thought to aid in the rapid growth of larvae at dehydrated through an ascending ethanol series, settlement (Wilkinson 1992), and to provide an cleared in xylene and infiltrated in paraffin wax advantage to adult sponges in competition with (Shandon Histoplast) using an automatic tissue algae and other photosynthetic organisms for processor on a nine hour cycle. The sponge substrate in high light areas (Wilkinson 1983). In tissue was further infiltrated with paraffin February 1998 a bleaching event was reported for under a vacuum of 635 mm Hg for 30 min prior a population of C. al/straliensis at Fremantle (south to embedding. Blocks were sectioned at 90 J.1m Western Australia), which coincided with a global thickness with a Leitz slide microtome, and hard and soft coral bleaching event (Fromont and section rolling was eliminated by placing filter Garson 1999). paper, moistened with distilled water, on top of Recently, Chondril/a, Chondrosia and two other the paraffin block. Sections were placed on a glass genera were united in a monophyletic order, slide smeared with egg albumin for adhesion, Chondrosida, based on molecular data (Boury- dried overnight at 60°C, and dehydrated in two Esnault and Lopes 1985), and containing a changes of xylene. Sections were mounted in single valid family, Chondrillidae Gray 1872. Shandon EZ-Mount and examined using light Prior to 1985 the family was located in the order microscopy. Images were recorded with a Leica Hadromerida but its affinities to this order were DFC420 camera on a Leica DME microscope and not clear. saved electronically. This study documents species of Chondrilla All sequences are available on GenBank recently collected from Australian waters. Type (accession numbers, 02 region: AY190190- material of species reported from Australia has AY190224, ITS region: AY190225-AY190239). been examined and reallocated where necessary. Methods to determine sequence comparisons Species currently known to occur in Australia and phylogenetic trees are detailed in Usher et al. are described and a preliminary assessment of (2004a). the biogeography of each species is provided. Abbreviations used in the text: AM, Australian The study also draws on previously published Museum, Sydney, Australia; BMNH, Natural molecular data sets (Usher et al. 2004a) and History Museum, London, United Kingdom; various biological characters such as symbiont LMJG, Landesmuseum Joanneum, Graz, Austria; relationships, reproductive biology and ecological NMY, National Museum of Victoria, Melbourne, distributions. Australia; NTM, Northern Territory Museum of Arts and Sciences, Darwin, Australia; SAM, MATERIALS AND METHODS South Australian Museum, Adelaide, Australia; Preserved material from various museums WAM, Western Australian Museum, Perth, (listed at the end of this section) was examined Australia; 2MB, Museum fuer Naturkunde, Berlin, during the course of this study. Collected Germany. specimens were preserved in 70°!c) ethanol. Skeletal structure and spicule morphology were examined SYSTEMATICS using light microscopy and scanning electron microscopy (SEM). Spicules were prepared by Order Chondrosida Boury-EsnauIt and Lopes 1985 boiling small pieces of sponge (including the ectosome and choanosome) in concentrated nitric Family ChondrilIidae Schmidt 1862 acid, followed by two consecutive washes with Chondrilla Schmidt 1862 both distilled water and absolute alcohol. The resulting spicule extracts were dried on a glass Type species slide and mounted in Shandon EZ-Mount (Thermo Chondrilla I1l/cl/la Schmidt, 1862 (subsequent Electron Corporation). Spicule dimensions were designation by de Laubenfels, 1936). determined by measurement of 20 randomly CllOndrilla linnaei sp. novo selected spicules per specimen using an eyepiece Figures la, 1b, 2-4 graticule with an Olympus BX50 microscope. Clean spicules were spread on coverslips or Australian C/lOlldrilla species 471 Figure 1 Whole specimen images of Australian Cholldrilla. A. C. lillllaei ill situ. B. Holotype (WAM 213267) of C. Iilll1aei after preservation in ethanol. C. C. secullda ill situ. D. C. seclI/lda after preservation in ethanol (WAM 213262). E. C. austrnliellsis ill sitl/. F. C. al/strnliellsis ill situ, maroon form. 472 J. Fromont, K.L. Usher, D.e. Sulton, S. Toze, J. Kuo Material examined General organisation: (Figure 2a, b). Ectosome: oxysphaerasters form a single layer at the surface Holotype or are sparsely distributed throughout this Australia: Western Australia: 5 pieces, in cave, layer. This region is 320-1000 ).lm thick