(P:art 1 of 3). Revision of Microcionidae (Porifera: Demospongiae), with description of Australian Species.

John N . A . Hooper

A phylogenetic revision of the poecilosclerid Microcionidae is based on type material, the worldwide literature, and comprehensive Australian collections. Of 73 available generic names 7 genera and 12 subgenera are recognised here. Of 561 available species names 459 are considered valid (10 virtually unrecognisable), including 52 new species. The Australian fauna, including Australian Antarctic Territory, contains 148 species (31 new), many new records, most are new combinations, and many illustrated for the first time. A synonymy of world species is provided. Valid taxa include: Clathria (with 7 subgenera: C. (Clathria) (with 31 Australian species, 82 other species worldwide, with new species murphyi, noarlungae, biclathrata, borealis, burtoni , sarai, saraspinifera), C. (Wilsonella) (6, 8, abrolhosensis, ensiae, lindgreni spp. nov.), C. (Microciona) (5, 91, illawarrae, lizardensis, simae, brondstedi, campecheae, claudei, hen tscheli, leighensis, stephensae, tunisiae, urizae, vacelettia spp. nov.), C. (Dendrocia) (7 species endemic to Australia), C. (Axociella) (6, 6, patula , fromontae, georgiaensis spp. nov.), C. (Isociella) (4, 1, selachia, skia spp. nov.), and C. (Thalysias) (36, 53, aphylla, craspedia, darwinensis, fusterna, hallmanni, hesperia, lematolae, phorbasiformis, styloprothesis, tin gens, wesselensis, amiranteiensis, hechteli spp. nov.); Antho (with 3 subgenera: A. (Antho) (12, 10), A. (Plocamia) (2, 17) and A. (Isopeneclya) (3,1, punicea, saintvincenti spp. nov.); Echinoclathria (14, 15, bergquistae, levii, notialis, parkeri, riddlei spp. nov.); Holopsamma (9 species endemic to Australia, 1 indeterminate species); Echinochalina (with 2 subgenera: E. (Echinochalina) (10, 2, felixi sp. nov.), E. (Protophlitaspongia) (8 species endemic to Australia and New Caledonia, collata, favulosa, isaaci, tuberosa spp. nov.)); Artemisina (4, 10); and Pandaros (incertae sedis) (0, 2). Generic keys are provided. Morphometric characters of primary importance include the origin, geometry and distribution of structural megascleres within the skeleton, modification of megascleres to monactinal or diactinal forms, the presence or absence of a specialised ectosomal skeleton, presence of detritus incorporated into spongin fibres, and overall skeletal structure (including compression of the axial skeleton and differentiation of axial and extra-axial regions). Brief zoogeographical comparisons are made between continental Australian and adjacent Indo-west Pacific faunas. Australian species comprise about 32% of the world's microcionid diversity; about 75% of species are endemic for the Australian region, and temperate species (81%) have higher levels of endemism than tropical species (59%).

MEMOIRS OF THE QUEENSLAND MUSEUM BRISBANE © Queensland Museum PO Box 3300, South Brisbane 4101, Australia Phone 06 7 3840 7555 Fax 06 7 3846 1226 Email qmlib@qm.qld.gov.au Website www.qm.qld.gov.au National Library of Australia card number ISSN 0079-8835 NOTE Papers published in this volume and in all previous volumes of the Memoirs of the Queensland Museum maybe reproduced for scientific research, individual study or other educational purposes. Properly acknowledged quotations may be made but queries regarding the republication of any papers should be addressed to the Editor in Chief. Copies of the journal can be purchased from the Queensland Museum Shop. A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum REVISION OF MICROCIONIDAE (PORIFERA: POECILOSCLERIDA: DEMOSPONGIAE), WITH DESCRIPTION OF AUSTRALIAN SPECIES. JOHN N.A. HOOPER Hooper, J.N.A. 1996 07 01: Revision of Microcionidae (Porifera: Poecilosclerida: Demospongiae), with description of Australian species. Memoirs of the Queensland Museum 40: 1-626. Brisbane ISSN 0079-8835. A phylogenetic revision of the poecilosclerid Microcionidae is based on type material, the worldwide literature, and comprehensive Australian collections. Of 73 available generic names 7 genera and 12 subgenera are recognised here. Of 561 available species names 459 are considered valid (10 virtually unrecognisable), including 52 new species. The Australian fauna, including Australian Antarctic Territory, contains 148 species (31 new), many new records, most are new combinations, and many illustrated for the first time. A synonymy of world species is provided. Valid taxa include: Clathria (with 7 subgenera: C. (Clathria) (with 31 Australian species, 82 other species worldwide, with new species murphyi, noarlungae,biclathrata, borealis, burtoni , sarai, saraspinifera), C. (Wilsonella) (6, 8, abrolhosensis, ensiae, lindgreni spp. nov.), C. (Microciona) (5, 91, illawarrae, lizardensis, simae, brondstedi, campecheae, claudei, hen tscheli, leighensis, stephensae, tunisiae, urizae, vacelettia spp. nov.), C. (Dendrocia) (7 species endemic to Australia), C. (Axociella) (6, 6, patula , fromontae, georgiaensis spp. nov.), C. (Isociella) (4, 1, selachia, skia spp. nov.), and C. (Thalysias) (36, 53, aphylla, craspedia, darwinensis, fusterna, hallmanni, hesperia, lematolae, phorbasiformis, styloprothesis, tin gens, wesselensis, amiranteiensis, hechteli spp. nov.); Antho (with 3 subgenera: A. (Antho) (12, 10), A. (Plocamia) (2, 17) and A. (Isopeneclya) (3,1, punicea, saintvincenti spp. nov.); Echinoclathria (14, 15, bergquistae, levii, notialis, parkeri, riddlei spp. nov.); Holopsamma (9 species endemic to Australia, 1 indeterminate species); Echinochalina (with 2 subgenera: E. (Echinochalina) (10, 2, felizi sp. nov.), E. (Protophlitaspongia) (8 species endemic to Australia and New Caledonia, collata, favulosa, isaaci, tuberosa spp. nov.)); Artemisina (4, 10); and Pandaros (incertae sedis) (0, 2). Generic keys are provided. Morphometric characters of primary importance include the origin, geometry and distribution of structural megascleres within the skeleton, modification of megascleres to monactinal or diactinal forms, the presence or absence of a specialised ectosomal skeleton, presence of detritus incorporated into spongin fibres, and overall skeletal structure (including compression of the axial skeleton and differentiation of axial and extra-axial regions). Brief zoogeographical comparisons are made between continental Australian and adjacent Indo-west Pacific faunas. Australian species comprise about 32% of the world's microcionid diversity; about 75% of species are endemic for the Australian region, and temperate species (81%) have higher levels of endemism than tropical species (59%). El Porifera, Demospongiae, Poecilosclerida, M icrocionidae, family revision, new species, taxonomy, biogeography, Australia. John N.A. Hooper, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101, Australia; received] December 1995. INTRODUCTION Microcionidae is one of the largest families of Demospongiae, comprising about 8% of all described (extant) Porifera species (Hooper & Levi, 1993a). The family has contained at one time or another about 70 genera and 550 species, although fewer than these are now recognised as valid. The family has a worldwide distribution and it is found from the intertidal zone to depths exceeding 2000m. It is clearly one of the more important, ecologically successful groups of Porifera. Within the Indo-Australian region microcionids are particularly abundant, with some species being dominant components of the shallow water macrobenthos. Previous works describing this fauna (and other literature containing extra-limital records of Australian species) include: Lamarck (1814, 1815, 1816), Gray (1858, 1867, 1869, 1870), Bowerbank (1864, 1875, 1877), Barnard (1879), Kent (1871), Ridley (1884a), Ridley & Dendy (1886, 1887), Lendenfeld (1888, 1889a), Kieschnick (1896, 1900), Thiele (1898, 1899, 1900, 1903), Kirkpatrick (1900a, b), Whitelegge, (1901, 1902), Hentschel (1909, 1911, 1912), MEMOIRS OF THE QUEENSLAND MUSEUM 2zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Hallmann (1912, 1914a-c, 1916a-c, 1920), Dendy & Frederick (1924), Topsent (1897b, 1930, 1932, 1933), Burton (1934a), Bergquist & Tizard (1967), Kelly-Borges & Bergquist (1988), Bergquist & Fromont (1988), Wiedenmayer (1989), Hooper (1990b), Hooper et al. (1991, 1992), Hooper & Levi (1993a, 1994). A brief synopsis of the fauna is given by Hooper & Wiedenmayer (1994), although some of the nomenclature and synonymies contained in that earlier work are revised here. Prior to the present study more than 200 species of Microcionidae had been described in the Australian fauna (including its territorial waters), but many of these were found to be either composite (consisting of several sibling species), or synonyms of other species. This study 1) describes 148 species (31 new), many new locality records for Australia and new taxonomic combinations; 2) provides an annotated synonymy for 311 other species worldwide (including 21 new species); 3) revises the morphometric characters used for classification and population variability for particular species; and 4) determines levels of endemism amongst provincial faunas. MATERIALS AND METHODS COLLECTION AND HISTOLOGICAL TECHNIQUES. Material examined in this study was predominantly collected using SCUBA (0-40m depth) or dredging and trawls (30-360m depth). Seasonal sampling for reproductive periodicity was conducted over two years in the Darwin and Cobourg Peninsula regions, NT. Immediately after collection specimens were either fixed in 80-100% methylated ethanol or frozen (which to some extent fixes the pigments), and later preserved in 70% alcohol. Reproductive products were searched for in fresh or frozen tissue. Nitric acid spicule preparations, thick-sections and thin-section mounts were routinely made as follows. Fragments of each sponge, including ectosomal and choanosomal regions, were heated directly on a glass microscope slide in several drops in nitric acid (the solution was evaporated rather than boiled, using low heat), and mounted in Canada balsam once completely dry, and cooled. Thick, hand-cut sections were made perpendicular to the surface, soaked in a saturated solution of phenol and xylene (for approximately 24 hours), and mounted in Durcupan (ACM Fluka Products) using glass slivers or card to raise the coverslip level. Phenol-xylene precluded the necessity for dehydration through an alcohol series. Some microtome sections cut at 30-35m were made for each species. Fragments were passed through a dehydration series, cleared in toluene or Histosol, and wax embedded for at least 2 hours. Sections were cut from trimmed wax blocks (cutting from the centre of the block to the exterior so as to include both the outer surface and inner skeleton relatively intact), placed in clearing agent for an adequate period to dissolve wax and/or dewaxing on a hot plate, then soaked in ethanol until perfectly clear, floated onto albumen-coated slides, orientated and flattened, stained with basic fuccsin and mounted. Fragments of dry specimens (e.g., type material) were reconstituted in 5% buffered fonnalin for 12 hours, which produced rehydration of the mesohyl and enabled cleaner histological sections to be made. MORPHOMETRIC ANALYSES. Spicu les were measured with a stage micrometer, either directly through a microscope or computer digitiser. Twenty five spicules, of each spicule category, in all specimens were measured. Acanthostyle width measurements were taken immediately below the base. Toxa lengths refer to chord length; isochelae are measured from apex of alae; width measurements of other spicules refer to maximum width. Spicule dimensions were sorted and statistically compared for various parameters (e.g., season, locality, depth), including one- and two-way ANOVs with replication, two-way ANOV with unequal replication, means differentiated using two-tailed t tests. Line-drawings were made using a calibrated camera lucida, and microphotographs were taken with an Olympus microphoto system. Taxonomic keys were constructed using ordered multistate, disordered multistate, morphometric and binary characters, utilising the DELTA system (Dallwitz & Paine, 1986). SCANNING ELECTRON MICROSCOPY. Sections were prepared as follows: 1) Cut at 1-1.5mm thick, ensuring that both the ectosome and choanosome were represented. 2) Placed in a cavity block and covered with several drops of sodium hypochlorite to etch the mesohyl matrix from the skeleton. The etching process was monitored through a dissecting microscope in order to prevent the skeleton falling apart. Delicate structures (plumose, halichondroid, hymedesmoid skeletons) only required a few seconds treatment with bleach; robust skeletons (reticulate, fibrous, articulated REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 3 skeletons) required several minutes; but generally 30 seconds was adequate. 3) Bleach was pipetted off at the appropriate time and 70% ethanol immediately added. Sections were left to stand for several minutes to ensure bleach was completely neutralised. 4) Steps 2-3 were repeated, without removing section from cavity block, substituting concentrated hydrogen peroxide in place of sodium hypochlorite, finally rinsing in ethanol. The hydrogen peroxide step was omitted for very delicate sections. 5) Sections were placed on clean microscope slides and let dry completely. 6) Sections mounted on SEM stubs using double-sided tape, copper dag, or `Supa Glue' (Supa Glue, Selleys Chemical Company, Padstow). An alternative method used to fix samples to stubs was to cover stub with `Aquadhere' wood glue (Aquadhere, Selleys Chemical Company, Padstow), let dry completely (usually several days), then prior to use exposed dry glue to vigorous steam (which softened the set glue), and placed the section on top of the stub (it would sink in a short way but was bonded reasonably well to the stub, and had the advantage of producing a perfectly smooth background). 7) The stub was sputter-coated well to ensure that all fibres were well coated to reduce 'charging'. In some cases uncoated sections could be viewed successfully under low accelerator voltage, but better results were generally obtained on coated specimens at higher voltage. Typical viewing conditions used were 25kV, at close working distance to provide best depth of field and focus, and at low magnifications. Spicule were prepared as follows: 1) Thinly cut sections including both ectosome and choanosome were placed in a durham tube (micro-test tube), to which drops of concentrated nitric acid are added, using drop-by-drop addition so as to control the oxidation reaction and production of by-product oxides. 2) Upon completion of acid digestion the durham tube was half filled with acid and gently heated over an alcohol flame, ensuring that only small bubbles form (low heat, no boiling), for 1-2 minutes. 3) Solution was let stand to cool, then centrifuged (approximately 4000rpm for 30 seconds). 4) Nitric acid was pipetted off leaving a spicule mass at the bottom of the tube, undisturbed. 5) Spicules were resuspended in fresh nitric acid and gently stirred using clean, fine, glass rod. 6) These steps were repeated if any collagen remained. 7) Spicules were resuspended firstly in demineralised water, 70% ethanol, then two series of 100% ethanol solutions, centrifuging and decanting the supernatant between each change of solution, finally ending with suspended spicules in a solution of absolute ethanol. 8) A micro-cover glass was adhered to an SEM stub using double-sided tap or copper dag, several drops of suspended spicules placed onto the cover glass, the alcohol-spicule solution ignited and spread across the glass with a glass rod or forceps until all ethanol was vaporised. Spicules bond to glass relatively firmly, but excess spicules could be blown off glass using compressed air, or spread out over the glass by adding further ethanol and igniting. The distribution of spicules on the cover glass was monitored under compound or dissecting microscope (magnification depending on spicule size). More drops of spicule solution added and this step repeated if too few spicules were present, ensuring not to overcrowd field of view for SEM photographic purposes. 9) An alternative method was used to produce a perfectly smooth background, using an `Aquadhere' glue-coated stub, dried for several days then softened with steam, and spicules placed directly onto soft glue (in this case ethanol was not burnt but evaporated). Single spicules would sink into glue too far if it was too soft (i.e., left in steam too long). 10) Spicule coated stubs were sputter coated briefly and viewed at 25kV, minimum working distance and smallest apperture for best resolution. ABBREVIATIONS AAT, Australian Antarctic Territories; ABIP, Centro de Estudios Avanzados de Blanes, Institut° de Investigaciones Pesqueras Barcelona, Aquarium de Blanes, Gerona; ABRS, Australian Biological Resources Survey, Canberra; AFZ, Australian Fishing Zone; AHF, Alan Hancock Foundation, University of Southern California, Los Angeles; AIMS, Australian Institute of Marine Science, Townsville; AM, Australian Museum, Sydney; AMNH, American Museum of Natural History, New York; BMNH, The Natural History Museum, London; BPBM, Bernice P. Bishop Museum, Honolulu; CP, Cobourg Peninsula, NT; CPMNP, Cobourg Peninsula Marine National Park, NT; CSIRO, Commonwealth Scientific and Industrial Research Or- MEMOIRS OF THE QUEENSLAND MUSEUM 4^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ganisation, Marine Laboratories, Hobart, Cleveland and Perth; DAR, Darwin region, NT; DELTA, Description Language for Taxonomy computer software (Dallwitz & Paine, 1986); EIS, Environmental Impact Study; CSIROEMG, CSIRO Food Research Laboratory, Division of Food Processing, North Ryde, Sydney. EMU, Ensight (formerly Environmental Management Unit), Water Board (Sydney, Illawarra, Blue Mountains), Sydney; EPA, Environment Protection Authority, Sydney; EPALR, East Point Aquatic Life Reserve, Dudley Point, Darw;n Harbour, NT; FNQ, far northern Queensland (Cooktown to Torres Straits); FUB, Freie Universitat Berlin; GBR, Great Barrier Reef, Queensland; HNUK, Natural History Museum, Ham Nam University, South Korea; ICBUC, Instituto Central de Biologia, Universidad de Concepcion, Chile; ICZN, International Code of Zoological Nomenclature (see Anonymous, 1985); IM, Indian Museum (Zoological Survey of India), Calcutta; IMZUB, Istituto e Museo di Zoologia ed Anatomia Comparata della Universita di Bari, Bari; IMZUN, Instituto e Museo di Zoologia dell'Universita di Napoli, Naples; INM, National Museum of Ireland, Dublin; IZUG, Museo Civico di Storia Naturale di Genova, Genova; JCU, James Cook University of North Queensland, Townsville; KFAU, Zoologischen Sammlung der Universitat Erlangen-Niirnberg, Erlangen; LFM, Merseyside County Museums (formerly Liverpool Free Museum), Liverpool; LMJG, Abteilung fiir Zoologie am Landesmuseum Joanneum (Landes Museum Jubileum Graz), Graz; MABA, Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia', Eaenos Aires; MCNP, Div. Invest. del Museo de Ciencias Naturales de la Plata, Argentina; MCZN, Museum of Comparative Zoology, Harvard University, Cambridge (Mass.); MEQ, mid eastern Queensland (Gladstone to Bowen); MHNG, Museum d'Histoire NatureIle de Geneve, Geneve; MLUM, Marine Laboratory of the University of Miami, Miami; MMBS, Mukaishima Marine Biological Station, Faculty of Science, Hiroshima University, Onomichi; MNHN, Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, Paris (DT, Topsent collections; DCL, Levi collections; DJV, Vacelet collections; DNBE, Boury-Esnault collections); MOM, Musee Oceanographique de Monaco, Monaco; MRAC, Koninklijk Museum voor MiddenAfrika, Tervuren; MRHN, Musee Royal d'Histoire Naturelle de Belgique, Bruxelles; MTQ, Queensland Museum, Museum of Tropical Queensland, Townsville; NCIQ66C-, United States National Cancer Institute, Australian Institute of Marine Science shallow water collection contract (1984-91), Townsville (primary voucher samples now lodged in QM, others in NTM and USNM); NC1OCDN-, United States National Cancer Institute, Coral Reef Research Foundation shallow water collection contract, Chuuk State (voucher samples lodged in QM and USNM); NEQ, northeast Queensland (Bowen to Cooktown); NM, Natal Museum, Pietermaritzburg; NMB, Naturhistorisches Museums zu Basel, Basel; NMCIC, National Museum of Natural Sciences, National Museums of Canada, Ottawa; NMNZ, National Museum of New Zealand (formerly Dominion Museum), Wellington; NMV, Museum of Victoria (formerly National Museum of Victoria), Melbourne; NSM, National Science Museum, Tokyo; NSW, New South Wales; NT, Northern Territory; NTM, Northern Territory Museum of Arts and Sciences, Darwin; NTU, Northern Territory University, Darwin; NWS, Northwest Shelf region, Western Australia; PAUP, Phylogenetic Analysis Using Parsimony (see Swofford, 1991); PIBOC, Pacific Institute of Bio-organic Chemistry, Far East Scientific Centre, Academy of Sciences of the USSR, Vladivostok; PMJ, Phyletisches Museum, Jena; PNG, Papua New Guinea; QFS, Queensland Fisheries Service, Department of Primary Industries, Brisbane and Cairns; QLD, Queensland; QM, Queensland Museum, Brisbane; QVML, Queen Victoria Museum and Art Gallery, Launceston; RMBS, Roscoff Marine Biological Station, Roscoff, France; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden; RRIMP, Roche Research Institute of Marine Pharmacology, Sydney (discontinued; sponge collections now held in AM); RSME, Royal Scottish Museum, Edinburgh; SA, South Australia; SAM, South Australian Museum, Adelaide; SEQ, southeast Queensland (Tweed River to Gladstone); SM, Musee Zoologique, Strasbourg; SME, Station Marine d'Endoume, Marseille; SMF, Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt; TAS, Tasmania; TM, Museo e Istituto di Zoologia Sistematica dell'Universita di Torino, Torino; TMAG, Tasmanian Museum and Art Gallery, Hobart; UAZD, University of Auckland, Zoology Department, Auckland; UB, Ubersee-Museum, Bremen; UCT, South African Museum of Natural History, Cape Town; UQ, University of Queensland, Brisbane; USC, University of Southern California, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 5 Los Angeles; USNM, National Museum of Natural History, Smithsonian Institution, Washington DC; UZM, Zoologisk Museum, Universitetsparken, Copenhagen; VIC, Victoria; WA, Western Australia; WAM, Western Australian Museum, Perth; YPM, Peabody Museum of Natural History, Yale University, New Haven (Conn.); ZIL, Zoological Institute of Leningrad, Academy of Sciences Museum of Zoology, St Petersburg; ZMA, ZoOlogisch Museum, Universiteit van Amsterdam, Amsterdam; ZMB, Museum ftir Naturkunde an der Humboldt-Universitat zu Berlin, Berlin; ZMC, Zoologisk Museum, Copenhagen; ZMH, Zoologisches Institute und Zoologisches Museum der Universitat Hamburg, Hamburg; ZMUU, Uppsala Universitets Zoologiska Museet, Zoologiska Institutet, Uppsala; ZSN, Aquarium e Museo della Stazione Zoologica di Napoli, Naples; ZRS, Zoologiska Rijkmuseum, Stockholm. ACKNOWLEDGEMENTS Patricia Bergquist (UAZD) encouraged this study and provided inspiration to persevere with the long learning curve associated with sponge taxonomy. Felix Wiedenmayer (NMB) provided access to his numerous unpublished personal notes on museum collections (Sponge Archives, NMV; Wiedenmayer, 1989; Hooper & Wiedenmayer, 1994). Rob Van Soest (ZMA) provided numerous discussions on sponges, alternative views on the diagnostic importance and polarity of characters, and possible relationships between sponge groups. Michelle Kelly-Borges (BMNH) and Peter Jell (QM) provided many positive comments on the manuscript. I am particularly grateful to Claude Levi (MNHN) for providing a post-doctoral fellowship at the MNHN and giving me access to its type collections, including all the Lamarck material; the Sir Winston Churchill Memorial Trust (Canberra), the Australian Biological Resources Study (Canberra), Klaus Riitzler (USNM), Willard Hartman (YPM), Joachim Reitner (FUB, now Gottingen) and the Trustees of the Queensland Museum (QM) for providing grants at various times, enabling me to examine major Museum collections and to interact with colleagues at several international forums (from which many of the ideas in this present volume matured). The Northern Territory University Planning Authority (1983-1985), the Museums and Art Galleries Board of the Northern Territory (1983-1989) and Heritage Commission of the Northern Territory (Darwin) provided additional funding to visit remote localities and the scattered collections in Australian museums. I am particularly grateful to Leonie Hooper for line drawings and John Kennedy for many of the SEMs. I also thank Bob Hardy (UQ), Charles Webb (NTU), and Clive Wilkinson (AIMS) for providing assistance with SEM photography. For financial or logistic assistance with fieldwork I acknowledge: George Elyakov, Valodya Krasochin, Y. Yakovlev, USSR RV `Akademik Opafin' (PIBOC); Alice Kay, Lester Cannon (QM) and Queensland Fisheries Service; Mick Ready (FV 'Hydronaue); Peter Murphy, Martin Riddle, Shirley Sorokin, Rob McCauley and other members of the NCI team (AIMS); Danny Roberts (EPA); Scott Chidgey, Calwell Connor and Associates; Patricia Byers (FV `Skeleton'); Bill Rudman (AM); Ian Poiner and Trevor Ward (CSIRO Fisheries, RV `Soela', RV 'Sprightly', FV 'Clipper Bird', RV 'Southern Surveyor'); Martin Riddle and Lisa Miller (EMU); Rob Capon (University of Melbourne); Alan Butler (University of Adelaide); Clay Bryce (WAM); Neville Coleman (Australian Marine Photographic Index, Brisbane); Conservation Commission of the Northern Territory, Darwin; Darryl Grey, Dave Ramm and Anne Coleman (NT Fisheries Darwin); Neil Smit (NT University Darwin) Barry Russell and Helen Larson (NTM); C.C. Lu (NMV); Cecile Debitus, George Bargibant, Jean-Louis Menou, Pierre Laboute (ORSTOM Noumea); Pat and Lori Colin (NCI CRRF Chuuk and Palau); and Ian and Pam Low (FV 'Rachel'). I am grateful for competent field assistance and laboratory technical assistance, during various parts of this study, from Jodie Baxter, Steven Cook, Lisa Hobbs, Alen Howard, Cathy Johnston, John Kennedy, Daniel Loy Choy, Anne-Marie Mussig, Paula Tomkins and Rex Williams. I also thank Phil Alderslade (NTM) for assistance in developing computer digitising software, and Russell Hanley (NTM) for identifying commensal polychaetes. I also thank many people for providing material for examination, or for other information cited in the text: Penny Barents (AM), Nicole Boury-Esnault (SME), Beatrice Burch (BPBM), Susan Chambers (RSME), Frank Climo (NMNZ), Ruth Desqueyroux-Faundez (MHNG and LMJG), Jane Fromont (JCU), Manfred Grasshoff (SMF), Jan Den Hartog (RMNH), Takomura Hoshino (MMBS), Frank von Knorre (PMJ), Deiter MEMOIRS OF THE QUEENSLAND MUSEUM 6zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Kuhlmann (ZMB), Vladimir Krasochin (PIBOC), Romely Lockyer (Cootamundra Shoals Survey team, UK), Susan Boyd (NMV), Liz McCaffrey (UQ Brisbane), A.K. Mandal (IM), Loisette Marsh (WAM), C. O'Riordan (INM), Shane Parker (SAM), David Parry (NTU Darwin), Urs Rahm (NMB), Martin Riddle (EMU), Frank Rowe (AM), Klaus Riitzler and Kathleen Smith (USNM), Shirley Stone (BMNH), B.R. Stuckenberg (NM), Ole Tendal (ZMC), Jean Vacelet (SME), Clare Valentine (BMNH), Clive Wilkinson (AIMS), and Wolfgang Zeidler (SAM). DEFINITION OF CHARACTERS MINERAL SKELETON. The form, composition and division of the skeleton remains the most important character for classification of Demospongiae. Recent attempts at higher systematics of Demospongiae based on non-skeletal characters (e.g., Simpson, 1968a; Bergquist & Hartman, 1969; Bergquist, 1980a; Lee & Gilchrist, 1985; Hooper et al., 1992) have had only limited success because in many instances they are unable to corroborate all skeletal and non-skeletal evidence into a single systematics. In some cases amongst Demospongiae (e.g., Verongida), non-skeletal evidence has been decisive and to some extent well correlated with other characters. In other cases (e.g., Axinellida) that evidence has merely highlighted inadequacies in systematics based solely on skeletal morphology (Simpson, 1968a; Bergquist & Hartman, 1969; Bergquist, 1980a; Vacelet, 1985; Hooper et al., 1992). COMPOSITION OF THE SKELETON. All microcionids are siliceous with discrete, free spicules. So far no desma-bearing species or hypercalcified 'relict' species are known. Many species undergo secondary acquisition, loss or reduction of spicule mineralisation, particularly when displaced by arenaceous particles (e.g., Holopsamma). C. (Wilsonella) is partly defined by this feature, with various degrees of arenaceous development among species; this trend is widespread throughout the Poecilosclerida. ORGANISATION OFTHE SKELETON. Gross organic and inorganic skeletal architecture, structural differentiation of the inorganic skeleton, and distribution of mineral components in that structure are primary diagnostics (Levi, 1960a, 1973; Bergquist, 1978a; Hartman, 1982). However, when used alone skeletal architecture is not necessarily a reliable indicator of relationships. Hooper (1991, herein) noted that the so-called typical, compressed, axinellid-like skeleton of many Raspailiidae (Raspailia pinnatifida; Hooper, 1991: Fig.2b) also occurred in C. (Axociella) (Figs 7G, 119A). It is also probable that skeletal structure is influenced to some extent by environmental conditions, and there is some evidence to suggest that flexible, compressed axial skeletons are produced in response to high energy environments (e.g., Palumbi, 1984). Similarly, skeletal characters such as those found in encrusting species have obviously evolved independently in many (otherwise unrelated) taxa. Review of microcionid skeletal structures showed that species which were similar in spicule geometry had different skeletal architectures (Hooper, 1988). Architectural types amongst the Microcionidae include: 1, hymedesmoid (with thin layer of basal spongin lying on the substrate containing erect megascleres (Fig. 7H); 2, microcionid (with a compressed basal spongin, producing ascending fibre nodes and plumose spicule columns) (Fig. 100F); 3, renieroid reticulate (consisting of a rectangular basal reticulation of uni- or paucispicular tracts fully enclosed within spongin fibres or cemented at their nodes by loose collagen) (Fig. 7C); 4, isotropic reticulation (a disorientated, random uni-, pauci- or multispicular reticulation in erect or massive forms, in which there is no distinction between primary or secondary tracts (not figured; seen only in nominal genus Qasimella); 5, isodictyal reticulation (reticulation with triangular meshes formed by uni- or paucispicular tracts of spicules, cemented at their nodes by collagen or fully enclosed within spongin fibres) (Fig. 131A); 6, regularly or irregularly reticulate (with large multispicular tracts and/or fibres forming irregular oval or rectangular meshes (Fig. 7A-B); 7, plumo-reticulate (producing ascending and consecutively diverging tracts and fibres, forming pauci- or multispicular primary lines, and interconnected by transverse uni- or paucispicular tracts and fibres) (Fig. 7D); 8, dendro-reticulate (similar to the preceding, but where ascending tracts are sinuous and more obviously diverging and branching than the less conspicuous transverse elements) (Fig. 231C); 9, plumose (with ascending and diverging primary lines that are not connected by transverse elements) (Fig. 7E); 10, axially or basally compressed (having a skeleton clearly divided into a compressed central or basal core of fibres and/or REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 7 12 of types: 1, principal spicules (Fig. 2A), robust, straight or slightly curved primary styles, subtylostyles, tylostyles or SUBECTOSOME quasi-monactinal forms generally (but not invariably) confined within fibres; 2, auxiliary spicules (Fig. 2B), CHOANOSOME more slender, curved, sinuous 10 or straight styles, subtylostyles, tylostyles or quasi-diactinal forms most commonly located outside fibres in the ectosomal or subectosomal skeletons, or dispersed throughout the mesohyl; 3, accessory spicules SPONGIN FIBRE (Fig. 4), styles, acanthostyles or modified quasi-diactinal 4568 forms generally echinating fibres. Thus, most species have FIG. 1. Idealised microcionid skeletal structure. 1, Echinating acanthostyles. principal spicules coring fibres 2, Reticulate fibre skeleton; 3, Isotropic extra-fibre skeleton. 4, Detrital (=choanosomal principals), entrapping fibres. 5, Renieroid reticulate secondary fibre skeleton. 6, some (e.g., Echinochalina) `Microcionid' radial fibre skeleton. 7, `Spicate' spicule skeleton. 8, Hymedesmoid spicule skeleton. 9, Plumose/dendritic fibre skeleton. 10, have coring spicules derived Coring principal spicules. 11, Subectosomal auxiliary spicules. 12, from auxiliaries (=choanosomal auxiliaries), and some (e.g., Ectosomal auxiliary spicules. Clathria (Dendrocia)) lack spicules, forming tight anastomosing meshes, principal spicules altogether. and from which arise plumose or plumoreticulate Ectosomal Skeleton. Within Microcionidae the extra-axial (subectosomal) fibres and/or spicules) ectosomal skeleton has been attributed greater (Fig. 7G). Some species have combinations of importance (Van Soest, 1984b) than over elethese skeletal structures with different structural ments such as choanosomal architecture and types found in different parts of the skeleton (e.g., growth form (e.g., Levi, 1960a; Berquist & axis and periphery). Fromont, 1988), microsclere diversity (e.g., de STRUCTURAL REGIONS OF THE SKELETON. The Laubenfels, 1936a), or megascleres echinating skeleton may be divided into three main struc- fibres and choanosomal spicules (e.g., Hallmann, tures or regions (Fig. 1): 1, ectosomal (outer 1912, 1920). Within the family ectosomal perforated epithelium comprised of single elon- development ranges from membraneous, without gate flattened pinacocytes, including an ec- an ectosomal skeleton (Fig. 94F); membraneous, tosomal skeleton, where present, arising from the with a specialised tangential, reticulate fibre mesohyl directly below the exopinacoderm); 2, skeleton (Fig. 255C); with an ectosomal tangenchoanosomal or axial core (central or basal region tial reticulation of detritus (Fig. 92A); with subcontaining choanocyte chambers, fibres and ectosomal spicules erect, paratangential or spicule tracts and the mesohyl ground substance, tangential to surface (Fig. 65F), or forming disin addition to the diverse and mobile cell types); crete bundles (Fig. 59E); with special ectosomal 3, subectosomal or cortical zone (strictly part of spicules erect or paratangential to surface, formthe choanosome since it lies within the bounds of ing a continuous palisade (Fig. 155G) or discrete the external epithelial barrier, but is immediately bundles (Fig. 151F). subdermal, and the delineation of this region from Subectosotnal Skeleton. The region between the choanosome proper is based on the develop- choanosomal and ectosomal layers may or may ment of an extra-fibre in some species structurally not be differentiated into a subectosomal (subderdifferentiated from the deeper choanosomal or mal or extra-axial) structure. This region may occupy a small proportion of the peripheral superficial ectosomal skeletons). This system is used in conjunction with spicule mesohyl (e.g., Holopsamma; Fig. 269D), or it nomenclature (Hallmann, 1912), based on origin may comprise the majority of sponge diameter ^ MEMOIRS OF THE QUEENSLAND MUSEUM 8zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 2. Major structural megascleres. A, Principal spicules (style, subtylostyle, acanthostyle subtylostyle, hastate style). B, Auxiliary spicules (style, subtylostyle, rhabdose subtylostyle, spined subtylostyle, asymmetrical styloid, tornostyle, quasistrongyle, mucronate styloid). C, Bases of principal spicules (style, subtylostyle, hastate style, acanthose subtylostyle, vestigial basal spination). D, Bases of auxiliary spicules (pointed subtylostyle, tylostyle, style, acanthose style, terminal spines, tornostyle). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 9 FIG. 3. Terminations of structural megascleres. A, Fusiform (pointed and rounded). B, Telescoped. C, Hastate (pointed and rounded). D, Spined (granular, tuberculate, terminal sharp, subterminal sharp, mucronate, bifurcate). (e.g., C. (Axociella); Fig. 118F). When present, subectosomal extra-fibre development may be completely disorganised or it may be well organised into radial non-plumose bundles of spicules (Fig. 127B). The composition and geometry of spicules within the subectosomal skeleton varies from species lacking any form of subectosomal spicules (e.g., C. (Microciona); Fig. 102F), those with a single form of auxiliary spicule throughout the skeleton (e.g., C. (Dendrocia); Fig. 112D), those with two categories of structural megascleres, one in the ectosome and one in spongin fibres (e.g., C. (Clathria); Fig. 73F), to those with three forms of structural megascleres, all of which occur to some extent in the subdermal region (e.g., C. (Thalysias); Fig. 197G), and spicules may be monactinal (for most microcionid species; e.g., Fig. 28A-B), or quasi-diactinal (Fig. 298A-B), or truly diactinal (Fig. 307A-B) as seen in E. (Protophlitaspongia). Choanosomal Skeleton. In Microcionidae skeletal structures are diverse (listed above), and traditionally important in classification. In terms of differentiated regions within the choanosomal skeleton there are species with simple reticulate fibres without any axial or extra-axial differentiation (Fig. 7C), those with marked axial and extraaxial differentiation of the choanosomal and subectosomal skeletons, which are structurally distinct from the peripheral skeleton (Fig. 7G), and those in which the choanosome is simply reduced to a basal layer of spongin lying on the substrate (Fig. 7H). Similarly, choanosomal skeletal tracts may be diversified into more than one element, including those with no obvious division of primary or secondary skeletal lines (Fig. 159F), those with more-or-less well differen- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 10zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 4. Echinating and acanthose principal megascleres and spination. A, Microspined echinating acanthostyles (typical recurved spines, typical erect, vestigial, poorly silicified, hastate smooth, entirely smooth). B, Myxillid-like echinating acanthostyles (aspinose apex, clavulate apex, entirely spined, bare neck). C, Modified principal spicules (three forms of intermediate principal echinating spicules, acanthostrongyles). D, Spine geometry (greatly recurved spines, slightly recurved spines, erect spines, granular spines, tuberculate spines). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 11 tiated primary (usually ascending) and secondary (usually transverse) lines of skeletal fibres or tracts (Fig. 48E), and those in which secondary (choanosomal) connecting tracts may be absent, leaving only radial primary lines (Fig. 227C). DISTRIBUTION OF SPICULES WITHIN THE SKELETON. Coring spicules. (Fig. 2) Spicules coring spongin fibres or forming tracts within the skeleton (bound by loose collagen) range from principal megascleres (e.g., C. (Clathria); Figs 2A, 28A), auxiliary monacts (e.g., C. (Wilsonella); Figs 2B, 83A), auxiliary diacts or quasi-diacts (e.g., E. (Echinochalina); Figs 2B, 280B), hastate oxeas (e.g., E. (Protophlitaspongia); Figs 2B, 296A), secondarily incorporated echinating spicules (e.g., C. (Thalysias)phorbasifonnis; Fig. 183G), acanthose principal styles, strongyles or rhabdostyles ('plocamiform' species; e.g., A. (Plocamia) ridleyi; Fig. 218C), coring spicules partially or completely absent replaced by foreign particles (detritus, algal filaments) (C. (Wilsonella); Fig. 91F; Holopsamma; Fig. 257D), primary fibres cored by principal styles, whereas secondary (connecting) fibres contain auxiliary megascleres (e.g., C. (Thalysias) mutabilis; not figured), primary fibres contain auxiliary spicules whereas secondary fibres mostly devoid of any spicules (e.g., C. (Clathria) noarlungae sp. nov.; Fig. 55F), axial fibres cored by auxiliary megascleres whereas peripheral fibres heavily arenaceous (e.g., C. (Clathria) echinonematissima; not figured). Echinating spicules. (Fig. 4) The presence, absence, distribution, geometry and secondary modification (including ornamentation) of echinating spicules (accessory spicules in the terminology of Hallmann, 1912) have been used as relatively important taxonomic characters by some authors (e.g., de Laubenfels, 1936a), whereas their importance in the supra-specific classification of the microcionids has been recently questioned (Simpson, 1968a; Van Soest, 1984b). The presence of echinating megascleres in the two families Microcionidae and Raspailiidae indicate the retention of an ancestral character by both families, interpreted as a synplesiomorphy (Hooper, 1991). In Microcionidae echinating spicules may be absent (e.g., C. (Axociella), Fig. 124F; C. (lsociella), Fig. 134D; presumed to be a secondary loss), echinated sparsely by acanthostyles or styles (e.g., C. (Clathria) nexus; Fig. 53E), heavily echinated by acanthostyles or smooth styles (e.g., C. (77talysias)procera; Fig. 187G), echinated by a second category of (acanthose) principal spicule (e.g., A. (Antho) tuberosa; Fig. 214F), or with both acanthose principal spicules and true echinating spicules present on fibres (e.g., A. (Plocamia) ridleyi; Fig. 218H). Similarly, echinating spicules may be confined to the exterior surface of fibres (most microcionids), or clumped on basal spongin only and absent from ascending skeletal lines (e.g., C. (Thalysias) tingens sp. nov.; Fig. 201G), concentrated in tufts at fibre nodes or fibre junctions (C. (Thalysias) spinifera; Fig. 197G), concentrated on exterior edges of fibres, or exclusively on primary fibres, and/or in the peripheral skeleton only (e.g., C. (Thalysias) abietina; Fig. 138F), or secondarily incorporated into fibres (e.g., C. (Dendrocia) myxilloides; Fig. 112D). Extra-fibre spicules. (Fig. 2) The distribution of structural megascleres within the choanosomal skeleton is certainly a feature relied upon by earlier authors (e.g., Carter, 1885 et seq.; Hallmann, 1912), but its value to the supraspecific classification has been recently questioned (Van Soest, 1984b). In most species these megascleres are confined within choanosomal fibres, lying parallel to fibres (e.g., C. (Clathria) raphana; Fig. 67D), or in the case of encrusting species, embedded in basal spongin (e.g., C. (Microciona) illawarrae sp. nov.; Fig. 100F). In other species they may core choanosomal fibres but also occur as plumose brushes on fibre endings (e.g., C. (Thalysias) spinifera; Fig. 197G). Choanosomal spicules may also poke out of fibres, much like echinating spicules (termed `spicate'), forming plumose tracts near the surface (e.g., C. (Thalysias) major; Fig. 181A), or they may be absent from the fibre core, standing perpendicular to the nodes or fibre junctions (e.g., C. (Clathria) biclathrata; Fig. 30G), strewn in loosely aggregated, halichondroid, extra-fibre tracts (e.g., E. (Echinochalina)anomala; Fig. 277A), form a renieroid structure, without a fibre component, but merely bound at the nodes by loose collagen (e.g., C. (Clathria) angulifera; Fig. 26F). SPICULE GEOMETRY. Spicule geometry is an important, sometimes ultimate (e.g., de Laubenfels, 1936a), feature of existing sponge classifications, based largely on the great diversity of megascleres and microscleres throughout the Porifera. This theoretical basis of sponge systematics using the mineral skeleton rests heavily on the assumption that morphogenesis of megascleres and microscleres is a function of the genotype (e.g., Reid, 1963). Whilst there is no ^ MEMOIRS OF THE QUEENSLAND MUSEUM 12zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 13 evidence to reject this hypothesis there is certainly some experimental data to show that spicule geometry and morphogenesis is at least partly influenced by environmental perturbations (e.g., Hartman, 1981; Jones, 1991), including examples from the Microcionidae (e.g., influence of seasonality (Simpson, 1978) and geographical distribution (Hooper et al., 1990) on spicule size and geometry). But the extent to which these phenotypic modifications occur within natural populations has not yet been examined rigorously. In general, however, these features appear to be relatively stable across wide geographical ranges as shown by studies on raspailiids (Hooper, 1991) and microcionids (Hooper & Levi, 1993a) from east and west coasts of Australia and the western Pacific. Hartman (1981) and Simpson (1990) outline the various theories on the functional significance, process of silicification and evolution of demosponge spicules. STRUCTURAL MEGASCLERES. Spicule axes. Microcionidae have exclusively monaxonic spicule axes. Megascleres are usually monactinal, although some may have modified secondary axes (i.e., anisoxeote diactinal modifications to styles), and a few appear to have true diactinal forms (E. (Protophlitaspongia)). Unlike Trikentrion and Cyamon in the allied Raspailiidae (Hooper, 1991) there are no tetraxonic spicule modifications in this family (triacti nal, tetractinal or polyactinal forms). Furthermore, the Microcionidae have a comparatively small range of structural megasclere types in the skeleton, whereas some raspailiids have many. Major types of structural megascleres are illustrated in Figs 2-4. These range from hastate styles or tylostyles (Fig. 87A), fusiform styles or tylostyles (Fig. 77A), asymmetrical styloid, rounded, quasi-diactinal or strongylote spicules (Fig. 280B), and oxeote megascleres (Fig. 296A). Spicule ornamentation. Spines on megascleres are of dubious importance to supraspecific clas- sification (e.g., Simpson, 1968a), although they have been used frequently in the past to define genera (e.g., de Laubenfels, 1936a). Microcionid structural spicules frequently have basal microspines (Fig. 2C-D), occasionally with spines on shafts (Fig. 180B) or points of spicules (Fig. 3D). Spicule ornamentation ranges from entirely smooth (Fig. 28A), smooth shafts with acanthose bases (Fig. 30A), vestigial spination on the proximal portions of shafts only (Fig. 153A), acanthose on both bases and points (Fig. 83A), or entirely acanthose (Fig. 98A). ECH1NATING MEGASCLERES. There is a diverse range of echinating spicule geometries in Microcionidae, although not as great as in Raspailiidae. Major types (Fig. 4) include: evenly spined (granular), claviform or stump-like acanthostyles; acanthose styles with aspinose bases; acanthose styles with aspinose points; acanthose styles with aspinose 'necks' (i.e., area proximal to the basal swelling); acanthostrongyles; entirely smooth styles identical in geometry to principal megascleres; derived oxeotes; or entirely smooth stylotes of different geometry than principal spicules. MICROSCLERES. The geometry, ornamentation and modification of microscleres is an important character for classification (Dendy, 1921), although it has probably been overemphasised by some authors (de Laubenfels, 1936a) and its primary importance has been questioned (Van Soest, 1984b). Within Microcionidae there are two forms of diactinal microscleres: meniscoid forms (chelae) and toxas. Other poecilosclerid microscleres (microxeas, raphides and meniscoid forms such as true sigmas) are not present. Microcionids show many modifications to both chelae and toxas, the latter sometimes resembling microxeas, and frequently microscleres are lost altogether. FIG. 5. Chelae geometry (A-H,J-N, Microcionidae; I,O-U, Other poecilosclerids). A, Palmate (C. australiensis). B, Palmate, reduced alae (C. australiensis). C, Palmate sigmoid, vestigial alae (C. hesperia sp.nov.). D, Palmate, arcuate-like alae with straight shaft (C. oxyphyla). E, Palmate, fused alae (C. curvichela). F, Palmate, contort (C. abietina). G, Palmate, arcuate-like fusion (C. macropora). H, Palmate, central wing on shaft (C. toxipraedita). I, Palmate, anisochelate (Mycale). J, Palmate, arcuate-like alae, fluted alae (C. macropora). K, Arcuate-like, fusion of alae, curved shaft, alae practically fused together (C. grisea). L, Arcuate-like, unguiferous, detached alae (C. scabida). M, Anchorate-like, unguiferous, tooth-like alae (C. scabida). N, Arcuate-like, unguiferous, tooth-like alae (C. myxilloides). 0, True arcuate (Ectyodotyx). P, Arcuate, unguiferous (Ectyodoryx). Q, Arcuate, unguiferous, tooth-like alae (Cretin). R, Palmate unguiferous (Crella). S, Palmate, unguiferous, vestigial alae (Hamigera). T, Arcuate, unguiferous, tooth-like alae (Monanchora). U, Anchorate, unguifcrous (Monanchora). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 14zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 6. Toxa geometry. A, Oxhorn. B, Wing-shaped (intermediate oxhorn-accolada. C, U- and V-shaped. D, Oxeote. E, Accolada. F, Sinuous and raphidiform. CHELAE. These are autapomorphic for Poecilosclerida, although Hajdu et al. (1994b) suggested that the Isodictya might belong to Haplosclerida (with chelae retained as an ancestral feature), although it is more likely that its niphatid-like isodictyal skeleton is merely convergent on the haplosclerid condition. There are many geometric modifications possible to chelae (Fig. 5) which Hajdu et al. (1994a) proposed could be accomodated into three main lines: pal- mate, arcuate and anchotrate (the latter two also termed bidentate- or tridentate-derived). These authors hypothesised further that chelae geometry might be an absolute indicator of phylogeny for the Poecilosclerida based on the assumption that they might be less susceptible to phenotypic modifications than structural spicules, following the arguments of Dendy (1921, 1922). This interpretation is based on parsimony and to some extent is corroborated by REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 15 other evidence (such as congruence of structural features). There are, however, some anomolous examples of chelae that fall between these three categories (see Discussion). Palmate. (Fig. 5A-B) This is the simplest form with 'typical' morphology consisting of straight shaft, front ala completely free and well developed, and lateral alae more-or-less completely fused to the shaft along its longest dimension. Most microcionids have unmodified 'typical' palmate isochelae. Modifications to this 'typical' palmate form include: partial reduction of alae (Fig. 5B), nearly vestigial alae producing a sigmoid spicule (Fig. 5C), partial fusion of alae along lateral margins producing spatulae (Fig. 5E), partial fusion (Fig. or complete fusion along medial margins n in which both the front alae meet and/or fuse at the centre producing cleistochelae (Fig. 76G), sculpturing on fluting on alae (Fig. 5J), contortion of the shaft such that alae are at 90 0 to each other at each end of the shaft (Fig. 5F), expansions of the lateral alae fused with the shaft producing wing-like process on the shaft (Fig. 5H), 'crocae' or j-shaped sigmoid forms where the alae are vestigial and asymmetrical, producing a simple hook-like spicule (Fig. 17F), and deep curvature of shaft and reduction of alae to tooth-like structures (termed unquiferous; Fig. 5R-T). Arcuate. (Fig. 50) Here the lateral alae are more fully developed than in palmate forms and become almost completely detached from the shaft, and the shaft is usually prominently curved and thickened. However, there is no clear transition between the palmate and arcuate forms, whereby an increase in curvature and thickening of the shaft (Fig. 5D-E) and partial detachment of lateral alae (Fig. 5J-K) extend along a continuum from true palmate to true arcuate (compare Fig. 5D, G, J-L, N-Q, T). Somewhere along this continuum chelae are deemed to be arcuate (Fig. 50-P). Anchorate. (Fig. 5U) Further along the continuum are anchorate chelae, in which all three alae are fully formed, the lateral ones completely detached from the shaft, and there are also lateral ridges on the shaft. In this study I use the terms 'arcuate-like' or 'anchorate-like' for modified chelae although it is equivocal whether these spicules are truly arcuates or anchorates. TOXAS. Toxas are found in only a few families of poecilosclerids but also known from Haplosclerida. There is also some evidence to suggest that they may be particularly common in young or larval tissue (e.g., Simpson, 1968b). Eight major morphotypes are delineated here (Fig 6), although intermediates are also possible: 1, Oxhorn toxas (wide central curve, reflexed arms and greatly recurved points; usually thick) (Fig. 6A); 2, Wing-shaped toxas (sharply curved at centre, with recurved arms and reflexed points; usually thick) (Fig. 6B); 3, U-shaped toxas (with wide central curvature but lacking reflexed arms) (Fig. 6C); 4, V-shaped toxas (pinched hairpin-like central curvature, straight arms running more-orless vertical, and slightly reflexed points; usually thick) (Fig. 6C); 5, Oxeote toxas (virtually straight shaft and points) (Fig. 6D); 6, Accolada toxas (wide or slightly pinched central curvature, strait arms running more-or-less horizontal, and strait points; usually thin) (Fig. 6E); 7, Raphidiform toxas (sharply angular central curvature, straight arms and straight points; very thin, hair-like) (Fig. 6F); and 8, Sinuous toxas (asymmetrical, sinuous, raphidiform; very thin, hair-like) (Fig. 6F). The presence or absence of microspines on toxas was at one time considered to be an important supraspecific character (e.g., de Laubenfels, 1936a), but these have since been found in many genera and may not be important above the species level. SIZE OF SPICULES. Variation of spicule size has also been an important diagnostic criterion, but this has been applied mainly at the species level of classification. Numerous (possibly a majority) of taxa have been erected solely on the basis of megasclere and microsclere dimensions, but only a few studies have investigated the statistical variability of spicule size or commented on the effects of physico-chemical factors on that variability (e.g., Hartman, 1958, 1981; Jones, 1984). There is some evidence to show that intraspecific variability can be significant for a given taxon, and spicule size-ranges can span across several closely related taxa which were otherwise erected solely on that basis. Hooper et al. (1990) demonstrated that two sibling species of Clathria (Thalysias) could not be reliably distinguished by their absolute spicule sizes, and only statistical comparisons between these species were of any value in this regard. Thus spicule dimensions used as diagnostic characteristics are of most significance at the species level of classification, and consequently their application is generally comparative rather than absolute. ORGANIC SKELETON. The development of the organic skeleton, the amount of spongin it contains, its architecture and foreign inclusions contained within it, are diagnostic features for the Demospongiae in general. The organic skeleton ^ MEMOIRS OF THE QUEENSLAND MUSEUM 16zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 7. Examples of microcionid skeletal structures. A, Regularly reticulate (C.(C.) noarlungae sp.nov.). B, Irregularly reticulate (C. (C.) multipes). C, Renieroid reticulate (A. (1.) chartacea). D, Plumoreticulate (C. (T.) reinwardti). E, Plumose (C. (T.) procera). F, Arenaceous (C. (W.) tuberosa). G, Axially compressed, extraaxially radial (C. (A.) canaliculata). H, Hymedesmoid (C. (M.) tingens). can be differentiated into two components: a fibre system and a collagenous mesohyl. Spongin fibres. These may be well developed, present but relatively lightly developed, secondarily lost, with or without spicule tracts, or even replaced partially or entirely by algal filaments (e.g., Anomoclathria (Fig. 9A-B)). Simpson (1984) elucidates two morphological types of spongin fibres (= type A spongin of Bergquist, 1978a): those that are cored by siliceous spicules and those that incorporate foreign particles. Detrital-entrapping fibres are most prevalent in the 'keratose' sponges (Dictyoceratida, Dendroceratida, Verongida), Haplosclerida (Arenosclera), and Poecilosclerida of the Desmacididae (Psammascus, Desmapsamnia), as well as several species of Microcionidae (i.e., C. (Wilsonella) and Holopsatntna). In these species fibres may be totally void of true megascleres (e.g., Holopsamma laminaefavosa), or they may have a combination of both foreign particles and indigenous megascleres (e.g., C. (Wilsonella) tuberosa). The independent occurrence of detrital-entrapping fibres throughout the various orders of Demospongiae indicates that it is a character of ecological importance rather than of any phylogenetic significance. Spiculose fibres are typical for microcionids, and the form, size, orientation and origin of megascleres coring the organic skeleton, and the structural complexities of the fibres themselves are important diagnostic characters for this family. In encrusting species that have a basal layer of spongin anchoring themselves to the substrate this spongin is continuous with the spiculated fibres, and as such both actually lie outside the living organism (Weissenfels, 1978). Mesohyl. The development of collagen in the mesohyl matrix, also termed 'interstitial' spongin, extra-fibre spongin, type B spongin (Bergquist, 1978), or ground substance, and including collagenous fibrils, has not been given much value as a systematic character for the Microcionidae, whereas in the 'keratose' orders REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 17 these features have more significance, and in Aaptos (Hadromerida) it has been used to differentiate species through deposition patterns (Kelly-Borges & Bergquist, 1995). However, within the Microcionidae there is evidence to show that the development of collagen throughout the mesohyl varies intra-specifically, especially between specimens in different reproductive condition or as a consequence of overwintering behaviour (e.g., Pandaros acanthifolium; Wiedenmayer, 1977; Van Soest, 1984b; Microciona prolifera; Simpson, 1963, 1968b; Knight & Fell, 1987). Simpson (1968a) attempted to define species and genera of Microcionidae on the basis of the organic skeleton and cytological characteristics but to a large extent his results did not corroborate with a classification based on the mineral skeleton, and in some cases evidence was directly conflicting. GROWTH FORM. The use of external morphology as an important or even crucial diagnostic characteristic has diminished since early systematics (e.g., Lamarck, 1814). Bowerbank (1864), in developing Grant's (1861) scheme for the Porifera, de-emphasised sponge habit in his systematics although he recognised that growth form was related to 'anatomical peculiarities'. That external morphology is often closely linked to the internal architecture and composition of the skeleton has been well documented (e.g., Levi, 1973; Bergquist, 1978). Although there are some groups which are immediately recognisable by their growth form and skeletal architecture (e.g., the honeycomb reticulate structure of Holopsamma), other groups show a higher degree of intraspecific variability in their morphology (e.g., most Clathria), ranging from encrusting to massive forms. Moreover, there is now evidence to suggest that gross morphology is highly plastic, greatly influenced by prevailing environmental conditions (temperature, depth, turbidity, currents, substrate etc.) (e.g., Hartman, 1958; Simpson, 1968a; Fry, 1971; Palumbi, 1984). It is not entirely clear to what extent abiotic factors influence growth form, or the degree to which genotype dictates possible shapes attainable by particular species, but it is becoming more apparent that the sponge 'species' is not as immutable as previously suspected. Palumbi (1984) proposed that sponges have evolved to be capable of producing a quick and decisive response to environmental adversities (unpredictable, high energy environments), and those responses are most readily seen as changes to both growth form and skeletal structure (e.g., the degree to which the skeletal becomes compressed). It is also not clearly understood why some species seem to be highly plastic (e.g., C. (Thalysias) lendenfeldi) (Hooper et al., 1990), whereas others with comparable depth and geographical distributions are much more conservative (e.g., C. (Thalysias) abietina). Growth forms, as characters used in a classification, can be defined as determinate (e.g., Holopsamma, C. (Microciona)) or indeterminate (e.g., most other Clathria, Antho). CONSISTENCY. Sponge texture is a highly subjective characteristic, which is difficult to quantify, but one which may provide clues as to the composition of the skeleton, the amount of spongin present, whether or not detritus is incorporated into the sponge, and silicification of the skeleton (Bergquist, 1978). A description of sponge consistency is usually an integral part of any species description, but its application in systematics has been mostly comparative rather than absolute. More recently de Weerdt (1985) used consistency as an objective feature in the systematics of Haplosclerida. She noted that it was not only useful in characterising particular species but that in broad terms,texture was able to be used at the family level of classification. For the Microcionidae this character does not vary greatly, with most species being firm, compressible, flexible (e.g., Clathria (Thalysias)) or soft, compressible, spongy (e.g., Holopsamma). MACROSCOPIC FEATURES OF THE SURFACE. Surface sculpturing. Macroscopic features of the sponge surface are important for some sponge groups (e.g., Haplosclerida), and surface ornamentation, such as tangential webs of spicules or fibres, perpendicular brushes of spicules, or elevated oscules may be diagnostic for particular genera. Within the Poecilosclerida however, including the Microcionidae, these features are less consistent, and they are usually only reliable in characterising particular species or occasionally genera (Simpson, 1968a). Encrusting species frequently exhibit intricate drainage canals radiating away from oscules, or highly hispid (furry) surfaces (e.g., C. (Thalysias) toxifera), whereas more massive or digitate species may have surface papillae or conules (e.g., C. (Thalysias) abietina), or a surface which is composed of reticulate ectosomal fibres (e.g., Holopsamma globosa). Oscules. The distribution of oscules on the surface may vary considerably between related species, ranging from being confined to distinct MEMOIRS OF THE QUEENSLAND MUSEUM 18^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA pore areas, such as sieve-plates of Echinochalina tubulosa, or restricted to certain regions, such as lateral sides of branches, the tops of digits, or the exterior surface of vases, or scattered indiscriminantly over the surface. Oscules may also be conspicuous, discrete, with a membraneous lip, slightly raised or flush with the surface (e.g., C. (Thalysias) reinwardti), or terminal, raised on the apex of surface papillae or stoloniferous tubes (e.g., C. (Isociella) eccentrica), or scattered, conspicuous, producing a porous reticulate surface (e.g., Holopsamma arborea), or minute and not easily visible optically (e.g., C. (Thalysias) coppingeri). COLOURATION. Sponge pigments are not generally diagnostic (Bergquist, 1978). Some species may show high intra-specific variation in live pigmentation, and this variability may be related directly to microhabitat and depth distribution. The nature of these pigments, their distribution within the mesohyl and their specific cellular association is still poorly known, but it is well established that carotenoids are predominant (Simpson, 1984). Isolation and identification of these pigments is more difficult, as is the determination of whether they are produced or modified by the host, or obtained directly from symbiotic associations (e.g., Litchfield & LiaaenJensen, 1980). The major proportion of sponge carotenoids are metabolised by the sponge (i.e., primary metabolites) and are intracellular (Simpson, 1984), whereas it is suspected that some sponges have a variable proportion (up to 20%) of pigments synthesised by symbiotic algae (e.g., Litchfield & Liaaen-Jensen, 1980; LiaaenJensen et al., 1982). Litchfield & Liaaen-Jensen (1980) studying C. (Microciona) prolifera suggested that the sponge could modify (aromatise) a large proportion of algal carotenoids, and Liaaen-Jensen et al. (1982) divide the classes of carotenoids into a phytoplankton-type, zooplankton-type, bacterial and/or fungal origin, and sponge metabolised (oxidative) groups. These authors found that phytoplankton derived and sponge metabolised carotenoids comprised the major proportion of carotenes in Demospongiae. The Poecilosclerida and Axinellida were found to exhibit the highest capacity for carotenoid accumulation and transformation, explaining their diverse and often brightly coloured pigmentation, and furthermore they possessed a similar carotenoid diversity. Evidence suggests that carotenoid pigments may be photoprotective, in which case it would be expected that intertidal species contain a higher proportion of these pigments than deeperwater species. But it is not clear why some sympatric species have consistent pigmentation (e.g C. (Wilsonella) tuberosa), whereas in others pigmentation is highly variable even in specimens growing side-by-side (e.g., C. (Thalysias) abietina; Plate 4D). Colour consistency is not generally used as a reliable diagnostic character, but it is also true that only very few authors have investigated the intraspecific colour variability of any species. It is therefore advantageous to determine whether live colouration is stable and specific to a species, or has very narrow limits in variation (e.g., C. (Isociella) eccentrica). Alternatively, pigmentation may be highly unstable, not specific and without an accurately definable 'typical' colouration (e.g., C. (Thalysias) abietina). REPRODUCTIVE PRODUCTS AND REPRODUCTIVE CYCLES. Reproductive products and modes of reproduction, as diagnostic characters, have been used predominantly at higher levels of classification (e.g., Bergquist, 1980a), whereas breeding seasons and spawning cycles are most useful for detecting sibling species (e.g., Fromont, 1989). As far as known, within the Microcionidae larvae are viviparous parenchymella with bare posterior poles. The apparent form of sexuality varies from gonochoristic to contemporaneous hermaphroditism (Fell, 1984, 1990; Simpson, 1984). Breeding seasons and/or spawning cycles may be continuous or periodical. CYTOLOGY. Simpson (1984) provided a definitive treatment of sponge cell biology, including a description of diverse cell types and their functional morphology. He suggested that descriptions of characters such as cells with inclusions and the morphology of choanocyte chambers will probably provide further information directly relevant to demosponge systematics. For the Microcionidae, Simpson (1968a) showed that seemingly morphologically convergent genera could be readily differentiated by the presence and morphology of special cell types (gray cells), and that their higher systematic relationships could be defined in terms of cytological characters. However, there were many incongruities between systematics based on skeletal characteristics and those indicated by cytological data. Specific examples of these differences are discussed below in the synopsis of genera, but some general comments are appropriate. Taxonomic groupings indicated by Simpson's microcionid cytological data suggested that many skeletal characters used previously by authors REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 19 had little importance in differentiating genera. These included the presence or absence of palmate isochelae, the presence of acanthose versus smooth echinating megascleres, quantity of spongin in the skeleton, plumose versus anastomosing fibres, megascleres with basal spination or smooth bases, the presence of surface conules and distinct oscules, the production of upright branches, and the presence or absence of an ectosomal skeleton. With the exception of the last feature these conclusions are supported in the contemporary classification of Microcionidae (e.g., Van Soest, 1984b). However, other cytological evidence presented by Simpson (1968a) is more difficult to reconcile with microcionid skeletal data. For example, encrusting species (i.e., the nominal genera Microciona and Ophlitaspongia) were cytologically relatively homogeneous and distinct from ramose forms (Clathria). The cytological characteristics of these encrusting species were more similar to renieroid microcionids (nominal genus Plocamilla) than they were to the ramose forms (nominal genera Rhaphidophlus and Thalysias) which otherwise had the most similar spicule and surface characteristics. Simpson concluded that generic definitions based primarily on spicule types did not lead to natural classifications, and he proposed that these definitions should include skeletal, cytological and histological evidence. He suggested that the numerous classification systems that were based solely on various combinations of skeletal characters, such as those of Vosmaer (1933, 1935a-b), de Laubenfels (1936a) and Levi (1960a), could be defended with equal justification. Although some cytological features have been incorporated into existing systematics (e.g., morphology and arrangement of choanocytes), much of Simpson's (1968a) important work cannot be used in classification based primarily on skeletal characters. SYSTEMATICS Class Demospongiae Sollas, 1885 Order Poecilosclerida Topsent, 1928 Suborder Microcionina Hajdu, Van Soest & Hooper, 1994 Poecilosclerida Topsent, 1928a: 64, 309. REMARKS. This order is the largest and most diverse of Demospongiae (Bergquist 1978). It is characterised by a skeleton of both spicule and spongin elements, usually well developed, sometimes vestigial, in which megascleres are monactinal, diactinal or both, and spongin development varies from well developed horny fibres enclosing spicules to an interspicular collagen cement (Bergquist, 1978; Hartman, 1982). Simpson (1984) suggested that the order is characterised by at least two distinctly localised types of megascleres (with or without distinctive geometry). Those megascleres are choanosomal principal spicules embedded in spongin fibres, and subectosomal auxiliary megascleres which are free in the mesohyl or protrude from spongin fibres in which they are embedded. This definition is consistent with the inclusion of Raspailiidae in the Poecilosclerida as proposed by Hooper (1991). Poecilosclerids usually have an abundantly collagenous mesohyl matrix, and microscleres may include chelae (apomorphic for the order), although not all taxa have them. Sexual reproduction is predominantly viviparous, oviparous in two families, and in those species incubating larvae they are parenchymella with uniform flagellum size and bare posterior poles. The suborder Microcionina was established to include four families of Poecilosclerida (Microcionidae, Raspailiidae, Iophonidae and Rhatxleremiidae), which have terminally spined ectosomal monactinal megascleres (occasionally modified to quasidiactinal forms), isochelae of palmate origin, diverse forms of toxas, up to five categories of megascleres and lacking sigmas. The other suborders (Myxillina and Mycalina) were also defined by their chelae morphology (bidentate-derived and sigmancistra-derived chelae, respectively), and absence of toxas and presence of sigmas, respectively (Hajdu et al., 1994), but assignment of particular genera to these suborders is still contentious. The number of families recognised in the order varies according to different authors (e.g., Levi, 1973; Wiedenmayer, 1977; Bergquist, 1978; Hartman, 1982; Van Soest, 1984b; Bergquist & Fromont, 1988). Recently Hooper & Wiedenmayer (1994) included 16 families in the order: 12 with chelae microscleres, 3 without chelae, and 1 of uncertain placement, whereas Hajdu et al. (1994) recognise 17: Microcionina (Microcionidae, Raspailiidae, lophonidae, Rhabderemiidae); Myxillina (Myxillidae, Crambidae, Coelosphaeridae, Crellidae, Hymedesmiidae, Anchinoidae, Phoriospongiidae, Tedaniidae); and Mycalina (Mycalidae, Hamacanthidae, MEMOIRS OF THE QUEENSLAND MUSEUM 20zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Desmacellidae, Cladorhizidae, Guitarridae). Latrunculiidae, included in the order by Levi (1973) and Van Soest (1984b) has also been assigned to Hadromerida (Reid, 1968; Bergquist, 1978; Hartman, 1982), but is now considered to be polyphyletic (Kelly-Borges & Vacelet, 1995) with Latrunculia having affinities with Iophonidae and Diacamus, Sigmosceptrella, Negombata more closely related to the Mycalidae. Family Microcionidae Carter, 1875 Microcionina Carter, 1875. Microcionidae Hentschel, 1923; Wiedenmayer, 1977. Clathriidae Lendenfeld, 1884a; Hentschel, 1923; Topsent, 1928a; Levi, 1960a; Simpson, 1968a; Bergquist, 1978; Hartman, 1982; Van Soest, 1984b; Bergquist & Fromont, 1988. Ophlitaspongiidae de Laubenfels, 1936a; Thomas, 1968; Hoshino, 1981. Growth form encrusting, lobate, arborescent or flabellate; skeleton differentiated into choanosomal (axial), subectosomal (extra-axial) and ectosomal regions; axial skeleton formed by unispicular or multispicular tracts of choanosomal (principal) megascleres, typically coring spongin fibres or sometimes simply bound together by collagen; fibres echinated by (acantho-) styles (accessory spicules); skeletal structures include isodictyal, renieroid, reticulate, plumo-reticulate, plumose or hymedesmoid, but never radial; extra-axial skeleton formed by tracts of subectosomal (auxiliary) spicules, usually dispersed outside of fibres, rarely well organised but usually with some degree of difference between axial and extra-axial regions; ectosomal skeleton ranges from membraneous, or with protruding subectosomal (auxiliary) spicules, or with a special category of ectosomal (auxiliary) spicules; principal megascleres monactinal, predominantly smooth or partially spined only, occasionally vestigial or absent completely, or sometimes replaced by detritus in skeleton; auxiliary megascleres usually monactinal, rarely quasidiactinal, smooth shaft and basal spines, more slender than choanosomal spicules; echinating styles or subtylostyles smooth, partially or completely spined; microscleres include toxas of several morphologies (including raphidiform and microxeotes), and isochelae primarily of palmate origin (but occasionally with partial 'arcuate' and `anchorate' modifications); larvae viviparous. REMARKS. There has been disagreement as to which of Microcionidae Carter and Clathriidae Hentschel should be used. Wicdcnmayer (1977: 139) argued that Microcionidae was established in 1875, whereas Clathriidae did not appear until 1884. He noted that under Article 40 of the International Code of Zoological Nomenclature (Anonymous, 1984), it was irrelevant whether or not Clathria Schmidt (1862) had priority over Microciona Bowerbank (1862; apparently published 1863). Conversely, Van Soest (1984b: 89) argued that the priority of Clathria over Microciona did have bearing on the choice of the family name. Although 'Clathriidae' is in current usage by most contemporary workers, its preferred use is in direct contravention with the Code and to long term stability of the group and Microcionidae is used here following Hooper & Wiedenmayer (1994). The definition given above restricts Microcionidae to genera which possess predominantly smooth monactinal ectosomal and choanosomal spicules. It excludes certain microcionid-like genera which have true tylotes or strongylotes as their ectosomal spicules (e.g., Acamus, Megaciella). These taxa are now referred to Iophonidae, as defined by their ectosomal features (Hajdu et al., 1994). However, the definition barely distinguishes species with modified or reduced quasidiactinal (styloid) auxiliary megascleres (e.g., several Echinoclathria, Holopsamma and Echinochalina species), or quasimonactinal (amphistrongylote or tornote-like) auxiliary megascleres (e.g., E. (Protophlitaspongia)). These modified auxiliary spicules are usually asymmetrical and are interpreted here as convergent upon true diactinal spicules. These anomalous microcionids share certain characteristics of both Microcionidae and Desmacididae, and the importance of these characters at higher levels of systematics must therefore be questioned, or a certain level of homoplasy must be accomodated in the phylogeny of the order. Similarly, the definition given above cannot always clearly distinguish some Microcionidae and Raspailiidae, but this is a problem of semantics rather than a biological one. As a general rule most species of Raspailiidae have well compressed axial skeletons, and well differentiated axial and extra-axial skeletons. In contrast, most Microcionidae lack these features or they are only poorly developed and probably convergent, perhaps related to growth form (e.g., Clathria (Axociella)). Nevertheless, there are examples in both families where the boundaries between taxa blur, such as the microcionid-like Raspailia (Clathriodendron) arbuscula (see Hooper 1991: Figs 19- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 21 20), and the raspailiid-like Clathria (Axociella) canaliculata (Figs 118-119)). These families are consistently differentiated by their ectosomal features and microscleres, which appear to be more important characters than skeletal structure. Hajdu et al. (1994) restricted Microcionina (and hence Microcionidae) to taxa with only palmate isochelae, tacitly excluding several microcionid-like genera specifically created for species with bidentate-derived (arcuate or anchorate) chelae. Theoretically this is a viable system for the suprafamily classification of Microcionidae but in practical terms it is not always possible to distinguish between true bidentate-derived chelae and palmate chelae with 'arcuate-' or 'anchorate-like' modifications. These cases are discussed indivually below. REVIEW. There are several problems in the taxonomy of Microcionidae that need to be addressed in order to clearly recognise and define valid genera and produce a phylogenetically valid systematics for the family. 1) The family is large, containing about 540 described species and many other as yet undescribed species known from various collections. 79 nominal genera have been previously included, of which 69 are currently recognised as residing here although fewer than this number are valid. Some of these genera have been merged in others by previous authors (e.g., Levi, 1960a; Simpson, 1968a; Van Soest, 1984b; Bergquist & Fromont, 1988; Hooper, 1990a), but in some cases these synonymies are now deemed wrong and have produced further nomenclatural complexities. Several contemporary studies have attempted partial revisions of Microcionidae (Van Soest, 1984; Bergquist & Fromont, 1988; Hooper, 1990a), but these have mainly focused on smaller regional faunas without consideration of all the higher taxa. In the present work each of these genera is redefined and illustrated from its type species (i.e., strict definition). 2) The literature on Microcionidae is vast, scattered, mostly antiquated (pre-1900), descriptions are far too brief for modern purposes and many taxa have never been illustrated. The present work deals primarily with museum material and living populations of species, and decisions are less reliant on the literature than previous studies. 3) There are many characters in sponges whose expressions (character states) change subtly within populations of supposedly single species and across the whole range of species, usually without clear boundaries between related taxa. Some of these characters have been used as important diagnostic criteria in earlier works. This study has examined large numbers of specimens and species, and documents the range of intraspecific and inter-specific character states in an effort to clearly define taxa and understand relationships between them. Inclusion of nonskeletal evidence into the systematics can further support or refute opinions based solely on skeletal characters and gross morphology (to decide whether one character is more important than another, whether morphological characters are homologous, and whether the observed high levels of homoplasies within most Poriferan classifications are in fact real or acceptable). The previous studies of Hooper et al. (1990) and Hooper (1990a) are preliminary to this study. 4) There are nearly as many subjective interpretations between different authors, as to the phylogenetic importance of one character over another in the systematics, as there are taxa. This has arisen partly as a consequence of over reliance on definitions of type species (and hence nominal genera) from the literature (especially the work of de Laubenfels, 1936a), given that many type species are poorly described, misdescribed or barely differentiated from their cogeners. The present study uses a phylogenetic framework to produce an objective and consistent taxonomy for the family. Two previous studies (Van Soest, 1984b; Hooper, 1990a) partially resolved intrafamily relationships within Microcionidae, both are preliminary to this work. GENERIC NAMES INCLUDED IN MICROCIONIDAE Preoccupied generic names are shown in square brackets. The synonomy lists provided in this section refer to works in which the name is used and in the case of genera considered valid do not include the numerous synonyms. The diagnoses provided in this list are based solely on the type material of the type species unless otherwise stated. [Abila] Gray, 1867 (Fig. 8A-B) Abila Gray, 1867: 539. Not Abila Gray, 1867: 522. TYPE SPECIES. Microciona laevis Bowerbank, 1866: 124 (by monotypy) (holotypc BMNH1877.5.21.1543). Encrusting growth form. Surface hispid, even. Choanosomal skeleton composed of short ^ MEMOIRS OF THE QUEENSLAND MUSEUM 22zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 8. Type species of microcionid genera. A-B, Abila (Microciona laevis Bowerbank, BMNH1877.5.21.424). C-E, Allocia (Spanioplon chel(erum Hentschel, SMF1571). F-H, Anaata (A. spongigartina de Laubenfels, USNM21428). I, Anomoclathria (Alcyonium opuntioides Lamarck, MNHNDT654). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 23 plumose spongin fibre nodes, cored by very long smooth or basally spined choanosomal principal styles, with only slightly swollen bases, and with bases embedded in spongin fibres on substrate and points protruding through ectosome. Echinating acanthostyles erect on substrate. Subectosomal skeleton of a single category of very long, curved, sinuous or straight subectosomal auxiliary subtylostyle, with smooth or microspined bases, forming irregular brushes on ectosome. Special ectosomal spicules absent. Microscleres thick wing-shaped toxas. Isochelae absent. REMARKS. Gray (1867) used the generic name Abila on two occasions, for two separate taxa. The senior name (Gray, 1867: 522) refers to a species originally described in Raspailia (viz. R. freyerii Schmidt), which Strand (1928: 32) unnecessarily proposed a replacement name Abilana Strand and which is a junior synonym of Raspailia Nardo (Strand should have replaced the junior name). The junior name (Gray, 1867: 539) was used fora microcionid, M. laevis Bowerbank, related to Microciona in skeletal structure, and related to Thalyseurypon de Laubenfels in lacking isochelae and having an encrusting growth form. De Laubenfels (1936a: 112) merged Abila with Hymantho Burton (see below), and Van Soest (1984b: 90) subsequently reduced Hymantho into synonymy with Clathria. Re-examination of the type species shows that the genus belongs with Clathria (Microciona). Allocia Hallmann, 1920 (Fig. 8C-E) Allocia Hallmann, 1920: 768; Bergyuist & Fromont, 1988: 95. TYPE SPECIES. Spanioplon chehferum Hentschel, 1 9 1 1: 362 (by original designation) (holotype ZMB4440, paratype SMF1571). Arborescent, foliose, planar growth form. Surface hispid, uneven. Choanosomal skeleton reticulate, with multispicular ascending primary fibres and paucispicular transverse connecting fibres. Spongin fibres heavy, cored by styles and tylotes (also scattered throughout mesohyl), echinated by acanthostyles with spinose shafts and bases, apinose 'necks', and large recurved spines. Subectosomal region with ascending primary fibres cored by choanosomal principal styles, latter protruding through surface. Ectosomal skeleton with both tangential and paratangential layers of auxiliary spicules, latter protruding from peripheral fibres and also lying tangential to surface. Megascleres completely smooth principal styles of a single size category, smooth subectosomal auxiliary stylote spicules, with asymmetrical rounded and microspined ends, and acanthostyles. Microscleres palmate isochelae of two sizes. Toxas absent. REMARKS. Hentschel (1911, 1912) expressed doubts concerning the initial generic assignment of S. cheliferum suggesting it showed certain affinities with Ectodoryx Lundbeck. Hallmann (1920) created Allocia to receive the species, including it in Microcionidae because of its 'typical' microcionid microscleres, echinating and coring megascleres. For similar reasons Dendy (1922: 70) merged the type species with Clathria. Based on its described ectosomal characteristics, supposedly consisting of true tylote spicules, the genus would be assigned to Iophonidae (see discussion for Acarnus), but reexamination of the type material found that these spicules were asymmetrical, modified styles very similar to those found in Clathria (Thalysias) major. Allocia is monotypic and belongs in Clathria (Clathria). The type species is known from the north and southwest coasts of Australia (Arafura Sea and Perth region), Amirante (Indian Ocean), and New Zealand (Bergquist & Fromont, 1988). Anaata de Laubenfels, 1932 (Fig. 8F-H) Not Aaata Semenov, 1906. Aaata de Laubenfels, 1930: 27. Anaata de Laubenfels, 1932: 89, 1936a:109. TYPE SPECIES. Aaata spongigartina de Laubenfels, 1 93 0: 27 (by original designation) (holotype USNM21428). Encrusting growth form. Surface hispid, even. Choanosomal skeleton hymedesmoid, with bases of both echinating acanthostyles and choanosomal principal subtylostyles embedded in basal fibres. Ectosome with dense erect palisade of subectosomal auxiliary subtylostyles overlaying choanosomal megascleres, both protruding through surface. Megascleres principal choanosomal styles with prominently acanthose bases and partially acanthose shafts, echinating acanthostyles with aspinose points, and auxiliary subtylostyles of a single size category, being completely smooth. Microscleres two size categories of palmate isochelae with arcuate modifications (strong curvature, partially detached lateral alae, slightly pointed unguiferous teeth). Toxas absent. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 24zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 9. Type species of microcionid genera. A-B, Anomoclathria (Alcyonium opuntioides Lamarck, MNHNDT654). C-E, Antherochalina (A. crassa Lendenfeld, BMNH1886.8.27.450). F-I, Anthoarcuata (A. graceae Bakus, USNM36284). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 25 REMARKS. De Laubenfels (1932: 90) suggested that Anaata was characterised in part by lacking echinating acanthostyles, although he noted that two size categories of choanosomal megascleres may be present. However, in the holotype the smaller category of spined spicule standing perpendicular to spongin fibres are true echinating acanthostyles, as found in most other microcionids. These smaller spicules are predominantly spined, whereas larger choanosomal spicules are only partially spined and are principals. Anaata was originally assigned to Hymedesmiidae Topsent, compared with genera such as Leptosiopsis Topsent (1927: 13) and Leptosastra Topsent (1904a: 194) (both of which have ectosomal tornotes with polytylote or anisotornote modifications, and anisochelae). All three genera have hymedesmoid architecture, which is persistent in Hymedesmiidae but also seen in many encrusting Microcionidae. Van Soest (1984b) mentioned that Anaata differed from other thinly encrusting microcionids, such as Clathria (Microciona) having arcuate isochelae replacing (or in addition to) palmate isochelae. It is questionable, however, whether these chelae are truly arcuate (as defined by Hajdu et al., 1994), given that their lateral alae are fused to the spicule shaft more than half way along their length. The smaller chelae are more-or-less evenly curved, slightly unguiferous (with reduced, slightly pointed alae), and lateral alae are fused to the shaft for about three-quarters their length. The larger spicules have more pronounced central curvature, they are more unguiferous (alae are reduced, sharper), and lateral alae are fused to the shaft for only about half their length. Anaata is considered here to be a Clathria (Microciona) with hymedesmoid architecture and chelae with slight 'arcuate' modifications. Anomoclathria Topsent, 1929 (Figs 81, 9A-B) Anomoclaihria Topsent, 1929: 26. TYPE SPECIES. Alcyonium opuntioides Lamarck, 1 8 1 5: 164 (by original designation) (lectotype MNHNLBIMDT654). Erect, arborescent or flabellate-digitate growth form. Surface smooth, not hispid. Choanosomal skeleton divisible into two components. Ascending primary skeleton plumose, with spongin fibres cored by paucispicular tracts of choanosomal principal styles and also incorporate algal filaments. Secondary basal skeleton renieroid reticulate, with spongin fibres fibres cored by large acanthostyles also grouped into plumose bundles on surface and secondarily incorporated into ascending primary fibres. Echinating spicules absent. Ectosome with sparse tangential skeleton of subectosomal auxiliary subtylostyles. Megascleres smooth choanosomal principal styles, acanthostyles with large spines in renieroid skeleton, with same morphology as principal spicules, and smooth subectosomal auxiliary subtylostyles. Microscleres palmate isochelae, wing-shaped and accolada toxas. REMARKS. This diagnosis on the lectotype differs slightly from Topsent's (1929, 1932) redescriptions of the species, particularly in the emphasis on the secondary renieroid skeleton overlaying the ascending plumoreticulate primary skeleton. This is a prominent diagnostic feature of Antho. In A. (Antho) the secondary renieroid skeleton is composed of acanthostyles, as described above for A. opuntioides, whereas in another species originally referred to Anomoclathria, Spongia frondifera Lamarck, 1814: 445 (lectotype MNHNLB1MDT565), the spicules forming the renieroid skeleton are acanthostrongyles (diagnostic for A. (Plocamia)). Topsent (1932: 103) considered that S. frondifera a junior synonym of A. opuntioides, whereas Lamarck's type material shows that their differences in a number of characters, including spicule geometry, is sufficient to warrant species level separation. Anomoclathria was erected on the basis of 'styloprothese' (Topsent, 1929), whereby algal filaments are incorporated into spongin fibres, partially or completely replacing the coring choanosomal spicules. But this symbiosis has subsequently been shown to be relatively common in marine sponges (Scott et al., 1984), occurring in many families (e.g., Bowerbank, 1862a; Carter, 1878; Lendenfeld, 1886b; Topsent, 1929, 1932, 1933; Weber-van Bosse, 1910,1921; Sciscioli, 1966; Scott et al., 1984; Price et al., 1984), and is an ecological rather than phylogenetic phenomenon. Although there is some evidence to suggest that particular species of algae are specific to particular sponge species, or at least restricted to a narrow range of taxa (Price et al., 1984)), it is unlikely that this symbiosis can be used as a diagnostic character at the supraspecific level. Nevertheless, it is intriguing to consider that a sponge can shed most or all of its diagnostic characters (spicules), replacing them with organic symbionts (e.g., see Clathria MEMOIRS OF THE QUEENSLAND MUSEUM 26zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ (Thalysias) abietina) or inorganic foreign particles (e.g., see Clathriopsamma, Wilsonella). Antherochalina Lendenfeld, 1887 (Fig. 9C-E) Antherochalina Lendenfeld, 1887b: 741, 786; Burton, 1934a: 558; de Laubenfels, 1936a:112. TYPE SPECIES. Antherochalina crassa Lendenfeld, 1887b:787 (by subsequent designation; Burton, 1934a: 558) (holotype BMNH1886.8.27.450). Erect, thin lamellate growth form. Surface smooth, not hispid. Choanosomal skeleton with differentiated axial (compressed, renieroid reticulate) and extra-axial (loose plumose) skeletons, although no regional differences in spiculation. Spongin fibres heavy, cored by choanosomal principal subtylostyles, echinated by small acanthostyles. Ectosomal skeleton with sparse tangential subectosomal auxiliary subtylostyles. Megascleres include robust, entirely smooth choanosomal principal subtylostyles, lightly spined acanthostyles, and auxiliary subtylostyles with basal spines. Microscleres palmate isochelae and wing-shaped toxas. REMARKS. Burton (1934a) designated A. crassa as type species of Antherochalina, in preference to Lendenfeld's (1887b) first-named species, Veluspa polymorpha var. infundibuliformis Maclay which was unrecognisable. He noted that A. crassa was a synonym of Clathria s.s.. Furthermore, of the eight species placed in the genus by Lendenfeld, only the type species now belongs here, whereas the other species are either unrecognisable or have affinities with Raspailia (Syringe/la), Phakellia, Ophlitaspongia, Cymbastela (A. concentrica; Hooper & Bergquist, 1992) or Ectyoplasia (A. frondosa; Hooper, 1991). Antherochalina also resembles some Desmacellidae (such as Sigmaxinella) and some Axinellidae (such as Axinella) in skeletal structure, having a slightly compressed renieroid axial and plumose extra-axial skeletons. Antho Gray, 1867 (Fig. 10A-C) Antho Gray, 1867: 524; Levi, 1960a: 57,76; Van Soest & Stone, 1986: 42; Bergquist & Fromont, 1988: 96. Plocamilla, in part, Burton, 1935a:402; Pulitzer-Finali, 1973:40 (not Plocamilla Topsent, 1928a: 63). TYPE SPECIES. Myxilla involvens Schmidt, 1864: 37 (by monotypy) (schizotype BMNH1867.3.11.92). Thinly encrusting (s.s.) or erect, arborescent, lamellate or vasiform growth forms. Surface rugose, hispid. Choanosomal skeleton renieroid reticulate with acanthostyles-strongyles coring spongin fibres, or simply united at nodes by variable quantities of spongin, producing triangular and rectangular skeletal meshes. Junctions of skeletal meshes with principal choanosomal styles echinating fibre nodes, standing erect or at oblique angles, in tufts or singly. True echinating megascleres absent (i.e., undifferentiated from choanosomal principal styles). Ectosome contains tangential or paratangential multispicular brushes of subectosomal auxiliary styles protruding through surface. Megascleres acanthose principal styles/strongyles of renieroid basal skeleton, smooth or acanthose principal styles/subtylostyles of the choanosomal skeleton, and smooth subectosomal auxiliary styles, often with basal spines. Microscleres palmate isochelae, wing-shaped and accolada toxas. REMARKS. This diagnosis is based on the type species and primarily on the type material, given the existing confusion about the true identity of A. involvens (cf. its alleged synonym A. inconstans; Ackers, Moss & Picton, 1992). Antho appears to be the earliest available name for a group of myxillid-like plocamiform sponges (sensu de Laubenfels, 1936a) which have microsclere spiculation typical of other Microcionidae. These taxa have an axial or basal skeleton composed of mostly acanthose styles or strongyles, producing a more-or-less regular renieroid reticulation. For this reason Berquist & Fromont (1988) referred Antho to the Myxillidae, but the genus has monactinal rather than specialised tylote ectosomal spicules and does not fit the concept of Myxillidae (Van Soest, 1984b). The potential generic synonymy of Antho is large. Burton (1930a: 501), de Laubenfels (1936a:77), Levi (1960a: 57) and Van Soest ( I 984b) combined species of Diayoclathria Topsent in Antho (see below), and that genus has largely disappeared from current usage. Burton's (1959a: 252) merger of Myxichela de Laubenfels (1935: 331, 1936a: 85) (type species Lissodendoryx tawiensis Wilson (1925: 432)) into Antho is not upheld here, because it has a true myxillid ectosomal skeleton (di actinal (tylote) ectosomal spicules). Renieroid reticulate skeletal architecture is not restricted to Myxillidae, known to occur in other families of sponges (e.g., Chalinidae (Haplosclerida), Iophonidae (Acarnus), Raspailiidae (Amphinomia Hooper, Plocamione Topsent, Lithoplocamia Dendy) and Axinellidae (Pitalia REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 27 Gray)), although it is certainly most common in the Myxillidae. The importance of a renieroid skeleton is interpreted differently by different authors, some giving it primary emphasis (e.g., Bergquist & Fromont, 1988) and others relegating it lesser importance (e.g., Van Soest, 1984b). Several species-groups have been created to accomodate microcionid-like species, with spiculation typical of the family, supplemented by a renieroid (myxillid-like) basal choanosomal skeleton composed of acanthose styles or strongyles. Vosmaer (1935a: 653) called this group Microciona prolifera tropus renieroides and de Laubenfels (1936a) recognised it at the family level (i.e., Plocamiidae sensu de Laubenfels (a junior homonym of Plocamiidae Topsent, 1928a)). However, only three genera appear to be sufficiently different to encompass all these microcionids: 1) Plocamia Schmidt (including Plocamilla Topsent, Dirrhopalum Ridley, and Holoplocamia de Laubenfels), which has predominantly (acantho)strongyles forming the renieroid skeleton (less commonly styles), and echinating acanthostyles overlap the main skeleton; 2)Antho Gray (including Anthoarcuata Bakus, Dictyoclathria Topsent, Isociona Hallmann and Jia de Laubenfels)), which has predominantly (acantho)styles forming the renieroid skeleton (less commonly strongyles), and echinating acanthostyles are absent; and 3) Isopenectya Hallmann, which has an axially compressed and extra-axially renieroid reticulate skeleton composed of two forms of choanosomal spicules inside spongin fibres, overlaid by a second extra-axial plumose skeleton. Several authors (Levi, 1960a; Simpson, 1968a; PulitzerFinali, 1973) were unable to reach a consensus of whether or not Plocamilla and Antho were identical, since they only really differed by the presence or absence of echinating spicules, and the extent to which basal spicules of the renieroid skeleton are styles or strongyles (i.e., intermediates occur). Most authors tentatively retain these two genera; Simpson (1968a) and Van Soest & Stone (1986) suggest edthat any decision on these genera, which differ from Clathria in having a renieroid skeletal architecture, would require thorough re-examination of all type species. This has now been done and the conclusion reached here is that differences between all three species-groups (Antho, Plocamia and Isopenecta) are not as great as the similarities (as conferred by the possession of renieroid skeletal structure), and these differences are emphasised only at subgenus. Anthoarcuata Bakus, 1966 (Fig. 9F-I) Anthoarcuata Bakus, 1966: 431. TYPE SPECIES. Anthoarcuata graceae Bakus, 1966:431 (by original designation) (holotype USNM36284 (161848)). Thickly encrusting, massive, cylindrical growth form. Surface even, microscopically hispid. Choanosomal skeleton with basal renieroid reticulation composed of uni- or paucispicular tracts (occasionally multispicular) of acanthostyles joined at nodes by light collagen. Peripheral ascending spicule tracts terminate in brushes of smooth principal styles, originating in choanosome and protruding through surface. Ectosomal skeleton has a dense, mostly tangential crust of smooth subectosomal auxiliary styles (of a single size category). Megascleres include smooth ectosomal auxiliary styles-subtylostyles, sometimes with mucronate bases and telescoped points, and principal acanthostyles with even spination forming basal skeleton. True echinating megascleres absent. Microscleres palmate isochelae with slight arcuate modifications (curvature, partial detachment of lateral alae from shaft, slightly pointed teeth). Toxas absent. REMARKS. The type species was originally identified as Burtonancora lacunosa (Lambe, 1892) by de Laubenfels (1961: 195), but Bakus (1966) noted that B. lacunosa (sensu de Laubenfels) was neither conspecific with Lambe's species nor referable to Burtonanchora (which in any case seems to be a synonym of Myxilla). Anthoarcuata graceae has the same spicule geometries and is structurally identical to Antho, differing only in the supposed possession of arcuate rather than palmate isochelae. However, in the type material chelae are not truly arcuate but are predominantly palmate with some 'arcuate' modifications including curvature and thickening of the shaft, partial detachment of lateral alae from the shaft (but for less than 20% of alar length), and reduced, slightly pointed, slightly unguiferous alae. They may be classed as palmate on the basis that lateral alae are only partially formed and are fused to the shaft for most of their length. Artemisina Vosmaer, 1885 (Fig. 10D-E) Artemisina Vosmaer, 18856: 25; Ridley & Dendy, 1887: 112; Topsent, 1894: 12; Lundbeck, 1905: 110; Burton, 1930a: 501, 528-531; de Laubenfels, ^ MEMOIRS OF THE QUEENSLAND MUSEUM 28zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 10. Type species of microcionid genera. A-C, Antho (Myxilla involyens Schmidt, MNHNDCL636). D-E, Artemisina (A. suberitoides, ZMAPOR443). F-G, Aulenella (A. foraminifera Burton & Rao, IM P I 167/1). H-I, Axocielita (Microciona similis Stephens, RSME1921.143.1447). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 29 1936a: 117; Levi, 1960a: 61, 83; Ristau, 1978: 585; Van Soest, 1984b: 122, 130; Bergquist & Fromont, 1988: 119. Artenisina;Burton, 1934b: 54 [lapsus]. TYPE SPECIES. Artemisina suberitoides Vosmaer, 1885: 25 (by monotypy) (holotype ZMAPOR443); junior synonym of Suberitesarciger Schmidt, 1870:47 (Burton, 1930a: 528) (schizotype BMNH1870.5.3.90). Massive, subspherical growth form. Surface skin-like, microscopically hispid, with few raised oscules; texture distinctly stringy. Choanosomal fibres indefinite or absent, overall architecture plumo-reticulate, nearly halichondroid in places, composed of multispicular ascending and paucispicular transverse tracts of choanosomal principal styles, bound together within spongin. Echinating megascleres absent. Ectosomal skeleton plumose, composed of single category of subectosomal auxiliary styles forming discontinuous palisade of discrete brushes. Megascleres smooth choanosomal principal subtylostyles and smooth fusiform subectosomal auxiliary styles or subtylostyles. Microscleres palmate isochelae, and wing-shaped toxas with spinous extremities. REMARKS. Defining Artemisina in phylogenetic terms is problematic. The taxon has no real distinctive features, although it differs from other Microcionidae in lacking a distinctive choanosomal skeleton or definite spongin fibres (also found in Qasimella Thomas), lacking echinating spicules, and having a nearly radial ectosomal skeleton reminiscent of some Ceratopsion (Raspailiidae; Hooper, 1991). These characteristics, two of which might be interpreted as reductions or secondary losses and the third as a convergence, are the only definable morphological apomorphies. Moreover, ectosomal structure varies between several species, ranging from the typical condition composed of erect brushes (e.g., A. arciger) to a tangential layer of spicules in criss-cross fashion (e.g., A. melana Van Soest). This variability is equivalent to (or analogues of) the Mycale subgenera Carmia and Aegagropila, respectively (e.g., Topsent, 1924b). At least one species lacks a specialised ectosomal skeleton completely (e.g., A. transiens Topsent). Some species of Artemisina (e.g., A. foliata (Bowerbank)) have honeycombed reticulate growth forms, approaching the characteristic Holopsamma morphology, but there is no consistency or pattern of gross morphologies amongst Artemisina, and in any case it is unlikely that the genus can be solely defined by its growth form. De Laubenfels (1936a) and Ristau (1978b) suggested that the Artemisina was defined by the absence of echinating acanthostyles and by the presence of spinous extremities on toxas. Neither character has much systematic value at the generic level. In the present interpretation echinating acanthostyles represent the retention of an ancestral character, in which case their presence or absence does not constitute a valid reason to define a phylogenetic grouping, and in any event they occur and disappear throughout numerous microcionid and raspailiid taxa. Similarly, toxas with spinous extremities are also known to occur in many Microcionidae, including the type species of Clathria, C. cotnpressa Schmidt, Microciona spinarchus Carter & Hope, M. coccinea Bergquist, M. rubens Bergquist M. spinatoxa Hoshino, Eutypon acanthotoxa Stephens, Stylostichon toxiferum Topsent, Labacea juncea Burton, Plocanzia ridleyi Hentschel, and Ophlitaspongia thielei Burton. They also occur in genera which have an ectosomal structure consistent with the Myxillidae (e.g., Melonchela clathrata Koltun). Several species of Artemisina have smooth toxas (e.g., A. melana), and one (A. archegona Ristau) has oxeote toxas similar to Paratenaciella Vacelet and Vasseur. Thus, in the broad sense Artemisina contains a heterogenous assemblage of species, which prompted Burton (1930a) to divide the group into three sections based on the number of megasclere categories present. The simplest forms have only one category of spicule (choanosomal principal megascleres; e.g., A. transiens Topsent); the typical form has two categories of megascleres (larger choanosomal principal styles and smaller subectosomal auxiliary styles; e.g., A. arciger); and the third form has an incompletely differentiated series of three megasclere types (two choanosomal spicules and one subectosomal spicule; A. plumosa Hentschel). In all these forms species are only really united in their lax choanosomal skeletal structure. The majority of species have been described from Antarctic and Arctic regions. [Aulena] sensu Lendenfeld, 1888 [Aulena]; Lendenfeld, 1888: 228, 1889a: 90; Topsent, 1894a: 19; de Laubenfels, 1936a: 16; Wiedcnmayer, 1989: 58. Not Aulena Lendenfeld, 1885c: 309. TYPE SPECIES. Aulena villosa Lendenfeld, 1885c: 309 (by subsequent designation; de Laubenfels, 1936a: 16) (syntypes AMZ130, G8901). Bulbous, subspherical growth form, consisting of fused digitate projections. Surface highly papillose but not hispid. Choanosomal skeleton MEMOIRS OF THE QUEENSLAND MUSEUM 30^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA regularly reticulate, with heavy spongin fibres cored by sparsely scattered sand grains mostly at nodes of fibres, and with a secondary fibre network between main skeleton. Ectosome lacks sand cortex or any other mineral skeleton. Megascleres and microscleres absent. identical to Clathria (s.s.), lacking only toxa microscleres. Vacelet et al. (1976: 75) correctly synonymised Aulenella and Clathriopsamtna, both of which now belong to C. (Wilsonella) (Van Soest, 1984b; Wiedenmayer, 1989; Hooper & Levi, 1993a). REMARKS. Hallmann (1912:275) merged Aulena and Echinoclathria Carter, whereas Wiedenmayer (1989) noted that in the strict sense (i.e., Lendenfeld, 1885c) it belongs to the Dictyoceratida (possibly related to Coscinoderma). Conversely, most other species referred to Aulena by Lendenfeld (1888) (A. laxa (Lendenfeld), A. gigantea (Lendenfeld), A. crassa (Carter)) belong to Holopsamma. Axocielita de Laubenfels, 1936 (Fig. 10H-I) Aulenella Burton & Rao, 1932 (Fig. 10E-G) Aulenella Burton & Rao, 1932:345. TYPE SPECIES. Aulenella foraminifera Burton & Rao, 1932: 345 (by original designation and monotypy) (holotype IMP1167/1). Flabello-digitate growth form, with subspherical, closely reticulate, honeycomb branching pattern. Surface uneven, not hispid. Choanosomal skeleton irregularly reticulate, with spongin fibres more-or-less fully cored by foreign particles and fewer choanosomal principal subtylostyles. Echinating acanthostyles abundant. Ectosome with single size category of subectosomal auxiliary subtylostyles tangential to surface and also strewn randomly throughout subectosome. Megascleres choanosomal principal subtylostyles with spined and tuberculate bases, echinating acanthostyles with evenly dispersed large recurved spines, and subectosomal auxiliary subtylostyles with or without spines on bases. Microscleres palmate isochelae, toxas absent. REMARKS. The 'honeycombed' reticulate growth form of Aulenella is reminiscent of Holopsamma and some E. (Echinochalina). The incorporation of foreign particles into the skeleton is also seen in species of Holopsamma, Aulena of authors, Clathriopsamma, Fisherispongia, Wilsonella and Anomoclathria. It also occurs in other Poecilosclerida Raspailia (Clathriodendron) (Raspailiidae), and many species of Phoriospongiidae) and many Dictyoceratida, and is considered here to be a successful ecological adaptation independently acquired by several groups. Thus,Aulenella is not defined by any unique features, and furthermore the holotype of A. foraminifera has spiculation Axocielita de Laubenfels, 1936a: 118; Hechtel, 1965: 44; Wiedenmayer, 1977: 140. TYPE SPECIES. Microciona similis Stephens, 1915: 441 (by original designation) (holotype RSME1921.143.1447). Thickly encrusting growth form. Surface even, and hispid. Choanosomal skeleton hymedesmoid, with spongin fibres forming basal layer on substrate and ascending non-anastomosing fibre nodes, each node containing plumose short unbranched tracts of choanosomal principal subtylostyles, standing perpendicular to substrate with only bases embedded in spongin fibres. Echinating subtylostyles erect on basal spongin and also echinating erect plumose brushes of choanosomal megascleres. Ectosomal skeleton with single category of subectosomal auxiliary subtylostyles forming paratangential tracts at surface and plumose brushes extending from ends of choanosomal megascleres. Megascleres smooth or minutely basally spined choanosomal principal subtylostyles, echinating subtylostyles with only bases spined, subectosomal auxiliary subtylostyles mostly smooth, occasionally basally spined, of a single size category. Microscleres palmate isochelae and small wing-shaped toxas. REMARKS. De Laubenfels (1936a) created Axocielita for thinly encrusting sponges resembling Axociella (i.e., lacking spined echinating megascleres). However, this is incorrect, based on misconceptions of both the type species (i.e., the published characters of Microciona similis do not agree with those seen in the holotype), and Axociella by de Laubenfels' (1936a). In the holotype of M. similis, diagnosed above, there is only a single category of auxiliary spicule, whereas Axociella s.s. has two distinct categories of auxiliary spicules in the peripheral skeleton. Thus, on the basis of its ectosomal features Axociella is strictly a Thalysias, whereas Axocielita has an unspecialised ectosomal spiculation typical of Clathria. In having plumose unbranched fibres forming a microcionid skeleton Hechtel (1965: 43) referred Axocielita to Microciona. He supported this ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 31 decision on the basis that M. similis has both spined and smooth styles, which he considered to be diagnostic for Microciona (although in the strict sense that feature is diagnostic for Anaata). Axocielita is referred here to Clathria (Microciona). Axociella Hallmann, 1920 (Fig. 11A-B) Axociella Hallmann, 1920:779; de Laubenfels, 1936a: 113; Wiedenmayer, 1977: 140; Bergquist & Fromont, 1988: 116. TYPE SPECIES. Esperiopsis cylindrica Ridley & Dendy, 1886: 340 (by original designation) (holotype BMNH1887.5.2.96). Arborescent, dichotomously branched, stalked growth form. Surface even, membraneous, transparent, hispid. Choanosomal fibre skeleton compressed at centre of stalk, with heavy spongin fibres forming reticulate axis, with longitudinal primary fibres cored by multispicular tracts and secondary connecting fibres aspicular or paucispicular tracts of choanosomal principal styles. Echinating megascleres absent. Subectosomal extra-axial skeleton plumose or plumoreticulate, well-differentiated from axial core, with light spongin fibres cored by multi- or paucispicular larger subectosomal auxiliary styles and few aspicular transverse fibres. Ectosomal skeleton composed of specialised category of smaller ectosomal auxiliary styles forming continuous palisade or individual brushes of spicules. Megascleres entirely smooth principal choanosomal styles-subtylostyles, and two categories of entirely smooth auxiliary styles-subtylostyles. Microscleres palmate isochelae and oxhorn toxas. REMARKS. Axociella has been misinterpreted by all authors since it was first reviewed by de Laubenfels (1936a). It is similar to Tenaciella Hallmann in lacking echinating megascleres but it has a distinctive, compressed reticulate axis and plumose or plumo-reticulate subectosomal (extra-axial) skeleton, reminiscent of the distinctive extra-axial skeletal architecture seen in Raspailiidae. The structure of the ectosomal skeleton in the type species of Axociella (i.e., the possession of two categories of auxiliary megascleres) is the same as seen in Thalysias, and Van Soest (1984b) merged the two genera on this basis. However, unlike typical species of Thalysias or Clathria, those of Axociella have a distinctive skeletal structure (compressed axis and a radial extra-axial skeleton), which is more- or-less homogeneous amongst the several known species, and this structure is interpreted here as indicative of supraspecific relationships. This interpretation is consistent with the treatment of similar structures in Raspailiidae (Hooper, 1991). In fact, Axociella could be justifiably included in Raspailiidae (Hooper, 1991; Hooper et al., 1992), apart from having isochelae and toxa microscleres and lacking the unique raspailiid ectosomal specialisation (large protruding auxiliary or principal spicules surrounded by bundles of smaller auxiliary spicules). True examples of these ectosomal spicules are not seen in the Microcionidae, although two species have analogues of this ectosomal condition: Esperiopsis canaliculata Whitelegge, Ophlitaspongia thetidis Hallmann — both belonging to Clathria (Axociella). Axociella is convergent upon Raspailiidae in skeletal structure, best developed in three Australian species, Esperiopsis cylindrica, E. canaliculata and Ophlitaspongia thetidis, all of which also have a Thalysias ectosomal skeleton. Another species from the NW Pacific, Microciona lambei Koltun, has a mix of both Raspailiidae and Microcionidae skeletons, being convergent on Endectyon in structure (with a markedly compressed axis, brushes of acanthostyles surrounding the bases of protruding choanosomal principal styles) but it also has palmate isochelae and a tangential layer of auxiliary styles lying tangential to the surface (i.e., the Clathria condition). Axociella is recognised here as a subgenus of Clathria based on its specialised compressed skeletal structure. Axosuberites Topsent, 1893 (Fig. 11C-D) Axosuberites Topsent, 1893a: 179; de Laubenfels, 1936a: 118. TYPE SPECIES. Axosuberitesfauroti Topsent, 1893a: 179 (by monotypy) (portion of holotype MNHNLBIMDT1859). Flabellate, flattened digitate growth form. Surface hispid, conulose. Choanosome with compressed reticulate axis and plumose extra-axial skeleton, with only light spongin fibres. Axial fibres produce close-meshed reticulation of multispicular tracts cored by choanosomal principal subtylostyles forming criss-cross reticulation, tracts plumose near periphery. Echinating megascleres absent. Subectosomal extra-axial skeleton well differentiated from axial region, with ascending plumose columns of larger subectosomal auxiliary subtylostyles arising from ^ 32zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 11. Type species of microcionid genera. A-B,Axociella(A. cylindrica Ridley & Dendy, BMNH1887.5.2.96). C-D, Axosuberites (A. fauroti Topsent, MNHNDT1859). E-G, Bipocillopsis (B. nexus Koltun, BMNH1963.7.29.56). H-I, Clathriella (C. primitiva Burton, BMNH1938.7.4.93). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 33 peripheral choanosomal skeleton. Ectosome with brushes of smaller auxiliary subtylostyles overlaying larger subectosomal spicules. Megascleres entirely smooth choano somal principal subtylostyles-tylostyles, and two size classes of auxiliary subtylostyles-tylostyles, both with smooth bases. Microscleres absent. REMARKS. Redescription of the type species above is based on examination of a slide preparation in the MNHN collection and Topsent's (1893a) description; the corresponding specimen has not yet been seen. Topsent (1893a) initially compared Axosube rites with Caulospongia Kent and Pseudosuberites Topsent in Suberitidae (Hadromefida), noting that it showed a certain level of morphological similarity in choanosomal fibres and geometry of tylostyle megascleres. However, in skeletal structure and spiculation Axosuberites appears to be a microcionid, most similar to Axociella, both genera showing similarities in their ectosomal specialisation, axial and extra-axial differentiation, and absence of echinating megascleres. Van Soest (1984b) also suggested that the genus was probably an Axociella without microscleres, and under his scheme it was therefore referable to Thalysias, but in both genera axial and extra-axial skeletal structures are well developed and closely comparable, and it is suggested here that they both should be included in the same subgenus (i.e., Clathria (Thalysias)). Bipocillopsis Koltun, 1964 (Fig. 11E-G) Bipocillopsis Koltun, 1964a: 79. TYPE SPECIES. Bipocillopsis nexus Koltun, 1964a: 80 (by monotypy) (holotype ZIL10644, paratype BMNH1963.7.29.56). Erect, arborescent growth form with cylindrical reticulated branches. Surface hispid, raised into irregular sharp conules. Choanosomal skeleton subrenieroid reticulate, with ascending spongin fibres cored by multispicular plumose tracts and interconnected by paucispicular transverse tracts of choanosomal principal subtylostyles, and echinated by acanthostyles. Subectosomal region with heavy paratangential bundles of subectosomal auxiliary styles protruding through surface and also scattered between fibres. Ectosomal skeleton without specialised spiculation but with bundles of subectosomal auxiliary styles surrounding protruding choanosomal spicules, and also lying paratangential to ectosome. Megascleres choanosomal principal subtylostyles invariably with basal spines and sometimes with spines on shaft, echinating acanthostyles of similar morphology to principal megascleres but shorter and more extensively spined, and single category of entirely smooth subectosomal auxiliary style. Microscleres isochelae, strongly sigmoid, unguiferous with very reduced pointed alae, possibly anchorate. Toxas absent. REMARKS. Bipocillopsis resembles Damoseni in lacking ectosomal specialisation and having modified sigmoid isochelae, but differs in growth form (arborescent versus encrusting) and subectosomal skeletal architecture (thickly paratangential versus tangential). The genus is monotypic and may be included in Clathria (Clathria) by its skeletal structure, whereas Damoseni has hymedesmoid skeletal structure typical of Clathria (Microciona). Chelae morphology has been described as tridentate sigmoid, allegedly related to the arcuate form, but this is very difficult to tell with certainty given that alae are nearly vestigial. There is no doubt that these chelae are more highly modified than most other microcionids. They are very small, strongly unguiferous (sigmoid curved with small pointed teeth), and lateral alae are completely free of the shaft and undifferentiated from the front ala, suggesting they may be of anchorate origin. However, there is no lateral ridge on the shaft of chelae to indicate an anchorate condition. The derivation of these chelae is indeterminable. Under the scheme of Hajdu et al. (1994) this genus could be included in Myxillina in possessing tridentatederived isochelae, whereas all other features (skeletal structure, principal and auxiliary megascleres and ectosomal skeleton) indicate relationships with the Microcionidae, in which it is retained here tentatively. This decision is supported by the case of Damoseni, discussed below, which have chelae of identical form to Bipocillopsis with the addition of oxhorn toxas (which are not found in Myxillina). Cionanchora de Laubenfels, 1936 (Fig. 12E-G) Cionanchora de Laubenfels, 1936a: 108. TYPE SPECIES. Hymeraphia tuberosocaphata Topsent, 1890b: 68 (by original designation) (fragment of holotype MNHNLBIMDT939). Encrusting growth form. Surface smooth, even, microscopically hispid. Choanosomal skeleton hymedesmoid. Spongin fibres reduced to basal MEMOIRS OF THE QUEENSLAND MUSEUM ^ 34zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA layer lying on substrate, with choanosomal principal subtylostyles erect and forming unispicular ascending columns protruding through ectosome, and echinating acanthostyles standing parallel to these. Ectosome tangential skeleton of subectosomal auxiliary subtylostyles, of a single category, forming brushes surrounding protruding choanosomal principal spicules. Megascleres choanosomal principal subtylostyles with tuberculate bases, echinating acanthostyles with large spines evenly dispersed over entire spicule except for bare point, and subectosomal auxiliary subtylostyles completely smooth. Microscleres anchorate-like isochelae. Toxas absent. REMARKS. The diagnosis is based on a slide of the holotype and Topsent's (1890b) description; the corresponding specimen has not been seen. Cionanchora was erected for thinly encrusting sponges with hymedesmoid skeletal construction, similar to Anaata, but with anchorate-like instead of arcuate-like modifications to isochelae (de Laubenfels, 1936a). Both those genera were merged with Clathria (s.1.) by Van Soest (1984b) on the basis that modified microscleres were a homoplasy throughout the Poecilosclerida, and are interspecific discriminators only. Anchoratelike modifications are also found in Folitispa (both of which were included in Clathria by Hooper, 1990a). Microciona dubia from Christmas Island (Kirkpatrick, 1900a: 136), was referred to Cionanchora by de Laubenfels (1936a: 108) supposedly in having anchorate-like chelae but these are of palmate origin. Both species have skeletal architecture typical of Microciona where they are referred. Clathria Schmidt, 1862 (Fig. I2A-B) Clathria Schmidt, 1862: 57; Ridley, 1884a: 443-449, 612-615; Ridley & Dendy, 1887: 31; Hentschel, 1911: 368; Hallmann, 1912: 205; Dendy, 1922: 64; Dendy, 1924a: 352-354; Wilson, 1925: 439; Topsent, 1925: 645-658; Topsent, 1928a: 62,299; Burton & Rao, 1932: 334-337; Burton, 1932a: 319; Burton, 1934a: 558; Koltun, 1959: 184; Levi, 1960a: 50,52,61; Melone, 1963: 1-8; Sara & Melone, 1963: 362; Sara, 1964: 229; Simpson, 1968a: 102, 104-106; Van Soest, 1984b: 90; Wiedenmayer, 1989: 56; Bergquist & Fromont, 1988: 106. Clathria Schmidt, plus Rhaphidophlus Ehlers; Ridley & Dendy, 1887: 146,151; Topsent, 1894a: 14-15,18. Clatharia;Kumar, 1925: 221 [lapsus]. TYPE SPECIES. Clathria compressa Schmidt, 1862: 58 (by subsequent designation (Schmidt, 1864: 35)) (holotype LMJG15509). Erect, arborescent, thinly lamellate, branching growth form. Surface even, not hispid. Choanosomal skeleton regularly reticulate, with well developed spongin fibres forming regular or irregular anastomoses of differentiated primary and secondary spongin fibres. Fibres cored by choanosomal principal subtylostyles in multispicular ascending tracts and uni- or bispicular transverse connecting tracts, and echinated by acanthostyles perpendicular to or at acute angles to spongin fibres. Ectosomal skeleton with tangential layer of subectosomal auxiliary subtylostyles, of a single size category. Megascleres basally spined choanosomal principal subtylostyles, entirely smooth subectosomal auxiliary subtylostyles, and echinating acanthostyles with even spination. Microscleres palmate isochelae and forceps-shaped or accolada toxas with spinose extremities. REMARKS. Strictly defining Clathria is essential in assigning a vast number of microcionids included in the genus by numerous authors. For example, C. compressa has toxas with spinose extremities, which therefore becomes a character 'typical' of Clathria, whereas earlier authors considered that this feature was diagnostic for Artemisina Vosmaer (de Laubenfels, 1936a). Clathria-like (viz. Labacea de Laubenfels) and Artemisina-like genera (viz. Ligrota de Laubenfels) also have spinous toxas, indicating that this character is homoplasious and probably not important above the species level of classification. In general, the original definition of Clathria (s.s.) is upheld here, as re-examination of Schmidt's syntype confirmed that all published characters cited in the species description (Schmidt, 1862: 58; Topsent, 1925: 647) are present in type material. Topsent (1925: 648) noted that C. compressa has variable spicule dimensions, skeletal architecture, and live colouration. He correlated this variability with the diverse growth forms shown by the species: thinly encrusting examples had a hymedesmoid skeletal architecture, thickly encrusting forms had a plumose skeleton, and erect ramose forms had anastomosing fibres (i.e., encompassing the nominal genera Leptoclathria, Microciona and Clathria). He also found correlation between the size of megascleres and growth form variability, although he could find no obvious trends. Topsent's observations are invaluable in deciding whether to maintain nominal REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 35 encrusting genera and more massive sponges as distinct genera, and whether to differentiate taxa with hymedesmoid, plumose or plumo-reticulate skeletal structure. Clathria compressa is known only from the north Atlantic and Mediterranean regions, but judging by its extensive synonymy it appears to be (or have been) moderately common within those regions. Clathriella Burton, 1935 (Fig. 11H-I) Clathriella Burton, 1935c: 73; Koltun, 1959: 186. TYPE SPECIES. Clathriella primitiva Burton, 1935c: 73 (by original designation) (holotype BMNH1938.7.4.93). Crumpled, irregular, massive growth form. Surface porous, uneven, hispid, with meandering ridges covered by thin transparent dermal membrane. Choanosomal skeleton renieroid reticulate, with spongin fibres forming regularly triangular meshes cored by multispicular tracts of smaller principal rhabdostyles (confined to renieroid network) and larger principal styles (latter producing secondary plumose, subisodictyal skeleton of bi- or paucispicular ascending tracts). Echinating spicules absent. Ectosomal skeleton radially arranged tracts of subectosomal auxiliary styles, of a single size category. Extra-fibre skeleton (apparently) has centrally curved (arcuate) oxeote megascleres scattered throughout mesohyl. Megascleres two categories of choanosomal principal styles, larger smooth with rounded bases, smaller spined with rhabdose bases, and smooth or basally spined subectosomal auxiliary styles-subtylostyles. Microscleres absent. REMARKS. Clathriella primitiva is obviously closely related to Clathria chartacea Whitelegge in its skeletal structure and spiculation, conforming to the definition of Isopenectya (see below). Clathriella also shows similarities to Isociella in having an isodictyal reticulate skeleton and in lacking echinating spicules, although in Isociella all megascleres are smooth and chelae and toxa microscleres are present. The presence of smaller spined rhabdostyles and an isodictyal component of the choanosomal skeleton are reminiscent of Rhabderemiidae, although the absence of thraustosigmata, thraustotoxa and other rhabderemiid microscleres in C. primitiva suggest that these similarities are convergent. Burton (1935c) considered that Clathriella was a primitive member of the Microcionidae, in which styles and acanthostyles had not yet become differentiated or segregated into coring and echinating megascleres. He suggested further that the toxiform oxeas, recorded by both Burton (1935c) and Koltun (1959), were derived from acanthostyles, and that both forms were remnants of a primitive condition. There is no empirical evidence to support either argument, and the present study takes the alternative point of view, that species like Clathriella prinntiva are derived or modified microcionids. Koltun (1959) also suggested that arcuate oxeas of C. primitiva were microxeas, and in that respect the genus should be compared with Paratenaciella. However, in Clathriella these spicules are supposedly large (200x7 p.m), indicating that they are true megascleres, whereas in Paratenaciella microxeas are very small (40-75 x 0.7-311m). In any case, re-examination of the holotype of C. primitiva (above) and more recent material from the Sakhalin Is, NW Pacific collected by PIBOC (QMG300052), did not find any toxiform oxeas although several examples of smaller auxiliary styles were sinuous, and it may be these the authors were referring to. The genus is referred here into synonymy with Atitho (lsopenectya). Clathriopsamma Lendenfeld, 1888 (Fig. 12C-D) Clathriopsamma Lendenfeld, 1888: 227; Topsent, 1894a: 19; Hallmann, 1920: 771; de Laubenfels, 1936a: 98; Levi, 1973: 614; Vacelet et al., 1976: 75. TYPE SPECIES. Clathriopsamma reticulata Lendenfeld, 1888: 227 (by subsequent designation; Hallmann, 1920: 771) (lectotype AMG9 1 35). Erect, anastomosing, arborescent growth form. Surface uneven, arenaceous, microscopically hispid. Choanosomal skeleton irregularly reticulate. Spongin fibres without well developed primary or secondary differentiation, cored by choanosomal principal subtylostyles and abundant foreign debris. In subectosomal skeleton principal subtylostyles also form plumose brushes, protruding through ectosome. Fibres heavily echinated by acanthostyles also associated with ectosomal spicule brushes. Ecto some with paratangential tracts of subectosomal auxiliary subtylostyles, of a single category, usually forming discrete brushes of spicules at surface. Megascleres basally spined, fusi form choanosomal principal subtylostyles, smooth and basally spined subectosomal ^ MEMOIRS OF THE QUEENSLAND MUSEUM 36zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 12. Type species of microcionid genera. A-B, Clathria (C. compressa Schmidt, LMJG15509). C-D, Clathriopsamma (C. reticulata Lendenfeld, AMG9135). E-G, Cionanchora (Hymeraphia tuberosocapaata Topsent, MNHNDT939). H-1, Colloclathria (C. ramosa Dendy, BMNH1921.11.7.64). REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 37 auxiliary subtylostyles, and echinating acanthostyles with large spines evenly dispersed. Microscleres palmate isochelae of two size categories, including contort forms, and sinuousraphidiform or accolada toxas. REMARKS. In Clathriopsamma the ectosome is structurally close to the Thalysias condition, but there is only one category of auxiliary spicule producing surface bundles. The genus is distinguished from most other microcionid genera in having foreign particles incorporated into the skeleton (Hallmann, 1920) (see remarks for Aulenella). Detrital entrapment is also known to occur in other Microcionidae (e.g., some Holopsamma), other Poecilosclerida (e.g., Phoriospongiidae), and it certainly also occurs commonly in other sponge orders (Dysideidae, Thorectidae and Ircinidae in the Dictyoceratida, and many Haplosclerida). This evidence indicates that this feature has arisen independently several times within the Porifera, probably indicative of ecological specialisation, and obviously arisen independently in several groups. However, species of Microcionidae that do incorporate sand appear to be relatively homogeneous in most of their other characters (i.e., there are no other conflicting characters such as presence/absence of ectosomal specialisation, or modifications to chelae), and consequently this specialisation is recognised here at the subgenus level. Vacelet et al. (1976) synonymised Clathriopsamtna with Aulenella, and Van Soest (1984b) merged Clathriopsatnma with Clathria, whereas in this work it is shown that the earliest available name for these species is Wilsonella (see below). Colloclathria Dendy, 1922 (Fig. 12H-I) Colloclathria Dendy, 1922: 74. TYPE SPECIES. Colloclaihria ramosa Dendy, 1922: 74 (by monotypy) (holotypeBMNH 1921.11.7.64). Cylindrical, arborescent, digitate growth form. Surface even, hispid. Choanosomal skeleton reticulate, with slightly compressed axis and slightly more cavernous extra-axial skeleton. Spongin fibres cored by large principal choanosomal styles, forming multispicular ascending tracts interconnected by multispicular transverse tracts. Echinating acanthostyles distributed irregularly over fibres. Subectosomal skeleton reduced to immediate outer edge of skeleton, with plumose tracts of larger subectosomal auxiliary subtylostyles extending through ectosome. Larger auxiliary megascleres also scattered between fibres and sometimes coring fibres. Ectosomal skeleton with smaller ectosomal auxiliary subtylostyle forming discrete brushes overlying subectosomal spicules. Megascleres smooth principal choanosomal styles, echinating acanthostyles with spined bases and points (bare 'necks'), and two sizes of auxiliary subtylostyles, usually with basal spines. Microscleres palmate isochelae, cleistochelae and accolada toxas. REMARKS. Colloclathria has a specialised ectosomal identical to Thalysias with two categories of auxiliary spicules forming surface spicule bundles, and on that basis Van Soest (1984b: 115) suggested the two genera should be merged: it is included here in synonymy with Clathria (Thalysias). The possession of cleistochelae in C. ramosa is not unique to Microcionidae also found in Plocamiopsis, Quizciona and several species of Clathria (e.g., C. (Clathria) toxipraedita). Damoseni de Laubenfels, 1936 (Fig. 13A-C) Damoseni de Laubenfels, 1936a: 110. TYPE SPECIES. Hymeraphia michaelseni Hentschel, 1911:351 (by original designation) (fragment of holotype SMF969T). Encrusting growth form. Surface sparsely hispid, even. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer of spongin lying on substrate, with bases of choanosomal principal subtylostyles and acanthostyles embedded in spongin, standing perpendicular to substrate, spicules protruding through ectosome. Ectosomal skeleton with paratangential, slightly plumose tracts of both larger and smaller auxiliary subtylostyles. Megascleres large principal choanosomal subtylostyles with spined bases, echinating acanthostyles with spined bases and shafts (bare 'necks'), and two size classes of subectosomal auxiliary subtylostyles with either smooth or spined bases. Microscleres strongly unguiferous, sigmoid isochelae with vestigial alae, arcuate- or anchoratelike but of uncertain affinity, and large wing-shaped (oxhorn derived) toxas. REMARKS. Hentschel's (1911) description of Hymeraphia michaelseni does not mention the presence of toxa microscleres, nor that principal spicules protrude a long distance through the surface. The modified unguiferous isochelae (of MEMOIRS OF THE QUEENSLAND MUSEUM ^ 38zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA possible anchorate or arcuate derivation) are identical to those of Bipocillopsis, and the combination of toxas and unguiferous isochelae of possible tridentate-derivation supports the inclusion of such taxa in Microcionidae contrary to Hajdu et al.'s (1994) proposal. Damoseni is similar to other encrusting genera with hymedesmoid architecture and it could be included in Clathria (Microciona) on this basis. However, the specialised ectosomal skeleton composed of smaller ectosomal and larger subectosomal auxiliary spicules indicates affinities to Clathria (Thalysias). Dendrocia Hallmann, 1920 (Fig. 13D-E) Dendrocia Hallmann, 1920: 767; de Laubenfels, 1936a:109. TYPE SPECIES. Clathria pyramida Lendenfeld, 1888: 222 (by original designation) (holotype AMG9047). Massive, lobate-digitate growth form. Surface conulose, with subdermal sculpturing, oscules slightly raised above surface with membraneous lip. Choanosomal skeleton dendritic, slightly plumo-reticulate near axis, without any obvious division of primary or secondary fibres, but with clear structural differences between choanosomal, subectosomal and ectosomal regions. Spongin fibres heavy, meandering, cored by auxiliary styles indistinguishable from those in ectosomal skeleton, and heavily echinated by acanthostyles (sometimes also secondarily incorporated into fibres). Subectosomal skeleton plumose or radial, with spicule tracts composed of auxiliary styles diverging from ends of peripheral choanosomal fibres and supporting overlying ectosomal skeleton. Ectosomal skeleton with continuous, heavy palisade of erect auxiliary styles. Principal spicules absent, and megascleres include only echinating acanthostyles with spined bases and necks but smooth points, and single category of structural spicule (auxiliary stylessubtylostyles), entirely smooth with hastate points and sometimes secondarily pointed bases. Micro scleres modified palmate isochelae with curved, thickened shaft and reduced alae. Toxas absent. REMARKS. This strict diagnosis based on the type species should be widened to include the presence of modifications to isochelae, ranging from typical palmate forms (with straight shaft, completely fused lateral alae, as in Clathria (Dendrocia) abrolhensis sp. nov.), modified palmate isochelae (with greatly curved, thickened shaft, partially detached lateral alae, as in C. (D.) dura), to anchorate-like forms (in which lateral alae are detached from shaft, shaft is curved, thickened and has a lateral ridge, as in C. (D.) myxilloides). One species also has oxhorn toxas (C.(D.) scabida) supporting the hypothesis that Dendrocia has affinities with Microcionidae. Dendrocia, like Wilsonella, differs from other microcionids in having auxiliary styles both coring fibres and forming the ectosomal skeleton. However, whereas Wilsonella has two categories of auxiliary spicules and detritus is incorporated into the skeleton, Dendrocia has only one category of structural spicule throughout the skeleton. Dendrocia also has a characteristic dendritic or plumo-reticulate skeletal architecture, whereas Wilsonella is invariably reticulate. In ectosomal skeletal structure (with a continuous ectosomal palisade of spicules) Dendrocia resembles the Thalysias condition, but it has only one geometric form of auxiliary spicule producing the extra-fibre skeleton (i.e., subectosome and ectosomal regions) more similar to Clathria. For this reason Dendrocia is enigmatic, and Hooper (1990a) maintained it as a separate taxon, whereas earlier Van Soest (1984b) had indicated that it was probably a synonym of Clathria. In this work it is referred to Clathria (Dendrocia). Two Australian species were initially included in Dendrocia by Hallmann (1920), the type species and C. alata Dendy (holotype NMV G2280). Both are undoubtedly synonyms. They apparently differ only in their spicule dimensions (styles: 175-230x2-p,m versus 240-250x4-p,m; acanthostyles: 63-95x3-1p.m versus 79-154x81p.m; arcuate isochelae: 23-2p.m versus 20-2p,m, respectively). In contrast, D. antyaja Burton & Rao, from the Indian Ocean, should be removed from Dendrocia because it has regularly reticulate choanosomal architecture, lacks an ectosomal skeleton, and has palmate isochelae indicating affinities to Clathria. Several other Australian species previously referred to Clathria s.s. should also be referred to Dendrocia on the basis of their skeletal architecture and spiculation. Dictyociona Topsent, 1913 (Fig. 13F-G) Dictyociona Topsent, 1913a: 579, 618; de Laubenfels, 1936a: 110; Levi, 1960a: 60; Bergquist & Fromont, 1988: 104. TYPE SPECIES. Microciona discreta Thiele, 1905: 447 (by monotypy) (holotype ZMB3302). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 39 Thickly encrusting, lobate growth form. Surface composed of interconnected micropapillae. Choanosomal skeleton plumose, becoming plumo-reticulate in thicker sections. Spongin fibres divided into primary and secondary elements, heavier in axis, lighter in subectosomal region where dermal spicules implanted at nodes of peripheral fibres. Fibres cored by multispicular tracts of choanosomal principal subtylostyles, and heavily echinated by acanthostyles projecting from fibres at all angles. Ectosomal skeleton composed of paratangential tracts of subectosomal auxiliary subtylostyles, of a single size category, arising from peripheral fibres and piercing surface. Megascleres choanosomal principal subtylostyles with acanthose bases and acanthose shafts near basal end, echinating acanthostyles with spined bases and points (bare 'necks'), and subectosomal auxiliary subtylostyles with microspined bases. Microscleres palmate isochelae and thin toxas intermediate between wing-shaped and oxhorn forms. REMARKS. Dictyociona resembles Microciona in its plumose skeletal structure but has partially acanthose choanosomal megascleres similar to those found in Anaata and Antho. These spicules are also present in several Raspailiidae (e.g., Hymeraphia, Eurypon), and they are considered here to be of minor diagnostic importance above the species level following Simpson (1968a). Levi (1960a: 60) merged Dictyociona with Clathria, although the species could as easily be placed in Microciona. It is included here within Clathria (Clathria). Apart from the type species, other taxa referred to Dictyociona at one time or another include: Microciona clathrata Whitelegge, M. heterotoxa Hentschel, M. pyramidalis Brondsted, Hymedestnia oxneri Topsent, Eurypon asodes de Laubenfels, E. acanthotoxa Stephens, E. ditoxa Stephens, E. microcizela Stephens, E. tenuissima Stephens, Clathria terra novae Dendy, D. contorta Bergquist & Fromont and D. atoxa Bergquist & Fromont. Dictyoclathria Topsent, 1920 (Fig. 13H-I) Dictyoclathria Topsent, 1920b: 18; Burton, 1930a: 501, 533, 1933: 50; de Laubenfels, 1936a: 77; Levi, 1960a: 80. Dyctioclathria [lapsus]; Ferrer Hernândez, 1921: 172. TYPE SPECIES. Clathria morisca Schmidt, 1868: 9 (by original designation) (schizotype BMNH1868.3.2.21); junior synonym of Antho involvens (Schmidt, 1864) (Levi, 1960a: 57) (schizotype BMNH1867.3.11.92). Arborescent growth form. Surface hispid, uneven. Choanosomal skeleton renieroid reticulate composed of spined acanthostyles forming triangular or rectangular meshes, enclosed within fibres (axis) or bound together at nodes by collagen (near periphery). Echinating megascleres absent. Subectosomal (extra-axial) skeleton plumose, with smooth principal styles standing perpendicular to fibre nodes, individually or in bundles, protruding through surface. Ectosomal skeleton with dense plumose or paratangential bundles of subectosomal auxiliary subtylostyles, sometimes surrounding protruding principal styles. Megascleres shorter lightly spined acanthostyles (renieroid skeleton) and longer completely smooth principal styles (extra-axial skeleton), and smaller subectosomal auxiliary subtylostyles with spined bases. Microscleres palmate isochelae and toxas intermediate between wing-shaped and oxhorn forms. REMARKS. Dictyoclathria is an objective synonym of Antho, since the type species of both genera are conspecific (Levi, 1960a). The type specimen of Dictyoclathria is remarkable in having a nearly raspailiid ectosomal condition with bundles of ectosomal spicules appearing to surround the protruding extra-axial spicules, but this is not as perfectly developed as in many Raspailia. Dirrhopalum Ridley, in Ridley & Duncan, 1881 Plocamia Schmidt, 1870: 62. Dirrhopalum Ridley in Ridley & Duncan, 1881: 477. See Plocamia. Echinochalina Thiele, 1903 (Fig. 14A-B) Echinochalina Thiele, 1903a:961; Hallmann, 1912: 288; Topsent, 1928a: 61; Burton, 1934a: 562; de Laubenfels, 1936a:118; Thomas, 1977: 115, TYPE SPECIES. Ophlitaspongia australiensis Ridley, 1884a: 442 (by subsequent designation (Hallmann, 1912: 288)) (holotype BMNH1881.10.21.299). Massive-digitate, reticulate growth form. Surface with low ridges and interconnected lamellae, producing angular cells with thin dermal membrane stretched between. Choanosomal skeleton irregularly reticulate, with heavy primary spongin fibres cored by paucispicular tracts of auxiliary spicules (tornotes), and heavy secondary spongin fibres cored by uni- or ^ MEMOIRS OF THE QUEENSLAND MUSEUM 40zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA - .01k1... FIG. 13. Type species of microcionid genera. A-C, Damoseni (D. michaelseni Hentschel, SMF969T). D-E, Dendrocia (Clathria pyramida Lendenfeld, AMG9047). F-G, Dictyociona (Microciona discreta Thiele, ZMB3302). H-I, Diayoclathria (Clathria morisca Schmidt, MNHNDT unreg.). ^ 41 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS bispicular tracts of auxiliary spicules. Fibres lightly echinated by principal subtylostyles. Ectosomal skeleton with undulating fibres and loose paratangential tracts of auxiliary spicules (also distributed throughout mesohyl). Megascleres include quasidiactinal auxiliary tornotes with aymmetrical or symmetrical ends (coring fibres and on ectosome), and completely smooth principal subtylostyles-styles (echinating fibres), sometimes modified to oxeotes (asymmetrical with two pointed ends). Microscleres wing-shaped toxas. Isochelae absent. REMARKS. In the type species coring spicules are tornotes, whereas in other species of Echinochalina they range from true styles to oxeotes. Re-examination of the holotype of 0. australiensis, and Thiele's (1903a) specimen from Ternate, Indonesia (SMF1855) also found that echinating principal styles can sometimes be modified to oxeote spicules, providing support for the otherwise tenuous placement of Protophlitaspongia with this group of microcionids. Hallmann (1912: 288) noted that Echinochalina differs from the allied Echinoclathria of authors (= Holopsamma as defined here) in having fibres cored by auxiliary spicules (tylotes or strongyles), which may be modified to quasi-monactinal forms (tylostrongyles, tornostrongyles), and echinated by smooth styles or subtylostyles. By comparison, Echinoclathria in the strict sense (or Ophlitaspongia of authors) has principal choanosomal styles (or modified monactinal megascleres) which both core and echinate fibres, as well as a second category protruding through the surface; Holopsamma has only a single category of coring and echinating principal spicule. As such, Van Soest (1984b: 129) suggested that Echinochalina possibly did not belong with the Microcionidae, and might be more appropriately placed elsewhere within the Poecilosclerida (e.g., Phoriospongiididae). However, examination of a suite of species included here in Holopsamma found that this group also undergoes a reduction in coring spicules (e.g., H. pluritoxa (Pulitzer-Finali)), whilst retaining other characteristics common to the genus, and it is possible that Echinochalina, sensu Hallmann, undergoes similar reduction. Probably of greater importance in determining the appropriate placement of Echinochalina is its lack of any special ectosomal skeleton, and in this respect it is similar to Dendrocia (both with only one form of structural (auxiliary) spicule). Whereas Echinoclathria (s.s.) possesses monactinal auxiliary spicules in the peripheral skeleton (sometimes reduced to quasi-diactinal forms), Echinochalina has spicules which are closer to true diactinals (although sometimes modified to quasi-monactinal forms). Both genera lack definite ectosomal specialisation. Some Echinochalina appear to have affinities with certain species of Echinodictyum (Raspailiidae) and with the Desmacididae, and both Thiele (1903a) and Topsent (1904a) have already noted this resemblance, suggesting that they differ mainly in the geometry, ornamentation and derivation of the echinating megascleres. By comparison, Hallmann (1912) considered that these differences, and the presence of long subectosomal styles in many Echinodictyum species, are sufficient to maintain the two genera as distinct taxa (see Hooper, 1991d). Although Echinochalina usually has smooth echinating megascleres, linking it to Echinoclathria and Holopsamma, there are two species which were previously referred to Echinodictyum (E. ridleyi Dendy and E. spongiosum Dendy), which have acanthose echinating megascleres but otherwise conform to Echinochalina in their spicule geometries and skeletal architectures. Hooper (1991) transferred these species to Echinochalina. Tablis de Laubenfels is an obvious synonym of Echinochalina. The two genera differ only by the absence of microscleres and the presence of a reticulate architecture in Tab/is. Echinoclathria Carter, 1884 (Fig. 14C-E) Echinoclathria Carter, 1884: 204 [nomen nudum]; Carter, 1885E 355; Ridley & Dendy, 1887: 159; Topsent, 1894a: 18; Thiele, 1903a: 962; Hallmann, 1912: 275-276; Dendy, 1922: 71 [in part]; Topsent, I928a: 61; de Laubenfels, 1936a: 119; Wiedenmayer, 1977: 143, 1989: 58. (not Uriz, 1988: 89]. TYPE SPECIES. Echinoclathria tenuis Carter, 1885E 355 (by subsequent designation; Burton, 1934a: 562) (holotype BMNH1886.12.15.147); junior synonym of Spongia leporina Lamarck, 1814: 444 (Topsent, 1932: 101) (holotype MNHNLBIMDT567). Thinly flabellate, flattened palmate, stalked growth form. Surface membraneous, microscopically hispid. Choanosomal skeleton renieroid reticulate, slightly compressed with well developed spongin fibres in axis, more openly reticulate, less compressed and with lighter spongin towards periphery. Axial fibres cored by pauci- or multispicular tracts of smaller principal styles producing rounded or irregularly shaped meshes, and echinated by same spicules. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 42zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Subectosomal (extra-axial) skeleton radial, unior paucispicular, with larger principal spicules erect on terminal fibres and usually protruding through surface. Ectosomal specialisation absent, with bundles of subectosomal auxiliary subtylostyles embedded perpendicularly and forming paratangential brushes surrounding larger principal spicules. Megascleres include smaller, robust, entirely smooth principal subtylostyles (coring and echinating fibres), larger principal subtylostyles of similar geometry (projecting from peripheral fibres and protruding through surface), and smooth subectosomal auxiliary subtylostyles, straight or flexuous. Microscleres absent. REMARKS. There is substantial confusion concerning the precise definition of Echinoclathria, and its relationship with other nominal genera such as Holopsamma, Halme, Aulena and Ophlitaspongia. Consequently the above diagnosis is strict, pertaining only to the type species, and a detailed explanation is justified below. Most authors follow Hallmann's (1912: 275) interpretation of Echinoclathria in which the genus is essentially characterised by 'a honeycomb mass of anastomosing flattened trabeculae', a reticulate skeleton of heavy spongin fibres cored and echinated by smooth monactinal principal megascleres of the same geometry, and with monactinal subectosomal auxiliary styles distributed throughout the mesohyl (and in some species also forming a radial subectosomal skeleton). Hallmann also noted that in some species he assigned to Echinoclathria there are both chelae and toxa microscleres, with quasimonactinal auxiliary megascleres, or they may have their coring megascleres replaced partially or completely by detritus (e.g., Holopsamma laminaefavosa). Hallmann suggested further that Echinoclathria and Ophlitaspongia essentially differed only in growth form, a view perpetuated by Wiedenmayer (1989). This interpretation is emended here. Most species included in Echinoclathria prior to the present study do have the characteristics outlined by Hallmann (1912), and most are relatively homogeneous and easily recognisable in the field by their characteristic 'honeycomb reticulate' growth form. It is therefore unfortunate that Burton (1934a) subsequently designated E. tenuis (a junior synonym of Spongia leporina) as the type species of Echinoclathria, because this species has a flabellate growth form (very dissimilar to 'honeycombed reticulate' species), skeletal architecture consisting of a dif- ferentiated axis, extra-axis, a renieroid skeleton, and a second category of principal spicules protruding through the ectosome. Echinoclathria leporina is typical of most Ophlitaspongia (of authors) (e.g., 0. axinelloides Dendy). Confusing the generic boundaries even further, Spongia leporina closely resembles Antho (Isopenectya) in growth form and gross skeletal construction, but differentiated by their skeletal structures, spicule ornamentation and localisation of particular spicules to certain regions of the skeleton. Nevertheless, it could be argued that Isopenectya could be included in Echinoclathria as equally as in Antho. The former option is tentatively rejected here based on the unequivocal possession of spined (versus smooth) styles composing the renieroid skeleton, and possession of a secondary, longitudinal, subisodictyal secondary skeleton in Isopenectya. Similarities in renieroid skeletal construction may link both these genera into a clade based on secondary reduction or loss. The difficulty in positively assigning Isopenectya is discussed further below. Wiedenmayer (1977: 144) suggested that Echinoclathria should be restricted to IndoPacific species, although there were similarities in skeletal architecture with Pandaros from the West Indies (which lacks microscleres). He noted further that Echinoclathria had mostly smooth choanosomal spicules, except for occasional vestigial spines on the bases of some auxiliary spicules, whereas Pandaros had acanthose echinating spicules. Simpson (1968a) has already demonstrated that the loss of spination on echinating spicules is common amongst microcionids and not correlated with any cytological differences (i.e., of low taxonomic value). Wiedenmayer (1977, 1989) concluded that although two genera intergraded they could not be consistently differentiated, and Pandaros is not included in this group. Thus, on the basis of evidence presented by Hallmann (1912: 275), Burton (1959a: 246) and Wiedenmayer (1989: 58), and re-examination of all nominal species belonging to these groups, it is clear that we are dealing with two distinct, homogenous groups. One, agreeing with the definition of Spongia leporina above, includes the genera Echinoclathria (in the strict sense only, and not of authors), and Ophlitaspongia (of authors, and not in the strict sense). Echinoclathria is the earliest available name for this group, and its characteristics should be taken as those traditionally associated with the concept of Ophlitaspongia (of authors). This genus is REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 43 most closely related to A ntho in its renieroid main skeletal structure, differing in having smooth choanosomal spicules and a single skeletal stucture (i.e., Antho has 2 skeletal structures: a renieroid skeleton composed of spined spicules and a plumose or subisodictyal skeleton composed of smooth spicules). The second group contains honeycombed reticulate species traditionally associated with Echinoclathria (of authors, not in the strict sense), together with Holopsamma, HaIme, Plectispa and Aulena (of authors, not in the strict sense). The earliest available name for this group is Holopsamma Carter (18850. There is no doubt that this honeycombed reticulate group of microcionids warrants inclusion in a separate taxon, although its level of divergence is arguable (cf. Wiedenmayer, 1977, 1989; Hooper, 1991). Its peculiar growth form is consistent for all 12 Australian species (all from southern Australia (Gondwanan) faunas), and 4 non-Australian species (2 from South America (Gondwanan) and 2 from the Indo-west Pacific (Tethyan) faunas). This growth form is correlated with a reticulate skeletal architecture, forming a homogeneous group which is recognised here at the generic level. A honeycombed reticulate growth form is also known for Acamasina de Laubenfels (1936a: 117) in Mycalidae (Van Soest, 1984b) and Panda ros (see below), both known only from the West Indies. Echinonema Carter, 1875 (Fig. 14F-G) Echinonema Carter, 1875: 194 [nomen nudum]; Carter, 1881a: 378; Ridley, 1884a: 615; Topsent, I 894a: 19; Dendy, 1896: 32; Whitelegge, 1901: 80; Topsent, 1928a: 61, 1932: 89, 98; de Laubenfels, 1936a: 112; Levi, 1960a: 56. TYPE SPECIES. Echinonema typicum Carter, 1881a: 377 (by typonymy) (lectotype BMNH1877.5.21.149); junior synonym of Spongia cactifonnis Lamarck, 1814: 440 (lectotype MNHNLBIMDT580). Arborescent, shrubby, lamellate growth form. Surface even, hispid, subectosomal striations. Choanosomal skeleton reticulate, with open rectangular or elongate open meshes although slightly compressed in axis. Primary spongin fibres ascending, heavy, cored by multispicular tracts of choanosomal principal styles, interconnected by pauci- or aspicular secondary spongin fibres. Fibres echinated by acanthostyles heaviest in peripheral skeleton. Subectosomal skeleton plumose, with brushes of larger subectosomal auxiliary subtylostyles erect on peripheral choanosomal fibres. Ectosomal skeleton dense, with smaller ectosomal auxiliary subtylostyles forming a dense palisade on surface. Megascleres include smooth choanosomal principal styles, larger subectosomal auxiliary subtylostyles with smooth or microspined bases, smaller ectosomal auxiliary subtylostyles with smooth or microspined bases, and short thick echinating acanthostyles with spined base and point but bare neck. Microscleres palmate isochelae of two sizes, including contort forms, and thin accolada and asymmetrical toxas. REMARKS. It is confirmed here that Carter's (1881a) Echinonema typicum is identical to Lamarck's (1814) Spongia cactifonnis, and consequently the name cactifonnis has seniority over the better known junior synonym Clathria typica, widely used in the literature. Lendenfeld (1888), Whitelegge (1901) and Hallmann (1912) created many new subspecific names (as varieties) for this species, and Hooper & Wiedenmayer (1994) assigned Lamarck's (1814) specimen to Clathria (Thalysias) cactifonnis cactiformis, and Carter's (1881) specimen to C. (T) cactiformis typica. Examination of type material of all these subspecies (var. typica (SMF1589); var. porrecta (SMF1653); var. brevispinus (AMZ931); var. favosus (AMZ944); var. geminus (AMZ928); var. obesus (AMZ937); var. proximus (AMZ930); var. stelligera (AME648); and three unnamed varieties of Hallmann (1912) (AMZ1158; AMZ1430, G9135, Z938; and AMZ41), showed that they were conspecific with the nominotypical variety (based on spicule size, spicule geometry and skeletal structure), whereas growth forms and surface features varied substantially between each taxon. The recognition of these subspecies, a preoccupation of many earlier authors, does at least demonstrate a high degree of external morphological variability for the species, but is not of particular nomenclatural importance because there are no other morphological characters that correlate with these differences in external morphology. It may be eventually determined from biochemical or genetic data, that this polymorphism is indicative of sibling species relationships, but no studies of this sort have yet been undertaken. De Laubenfels (1936a: 112) suggested that Echinonema was identical to Thalysias (sensu de Laubenfels), whereas Van Soest (1984b) and others placed both genera into synonymy with Rhaphidophlus (see below). The genus is in- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 44zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 14. Type species of microcionid genera. A-B, Echinochalina (Ophlitaspongia australiensis Ridley, BMNHI881.10.21.299). C-E, Echinoclathria (E. tenuis Carter, BMNH1886.12.15.147). F-G, Echinonema (E. typicum Carter, BMNH1877.5.21.149), H-I, HaIme (Holopsamma laminaefavosa Carter, BMNHI886.12.15.312). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 45 eluded here in Clathria (Thalysias) on the basis of its ectosomal specialisation. Fisherispongia de Laubenfels, 1936 (Fig. 15A-C) Fisherispongia de Laubenfels, 1936b: 460. TYPE SPECIES. Fisherispongiaferrea de Laubenfels, 1936b: 460 (by original designation) (holotype USNM22239). Encrusting growth form. Surface tuberculate, arenaceous, hispid. Choanosomal skeleton hymedesmoid, with basal layer of spongin fibre incorporating detritus and bases of larger choanosomal principal subtylostyles and smaller echinating styles, standing perpendicular to substrate, in groups or individually, ascending to but not protruding through ectosome. Ectosomal skeleton plumose, with subectosomal auxiliary polytylostyles, of a single category, arising from ends of choanosomal megascleres in multispicular bundles protruding through surface. Megascleres large choanosomal principal subtylostyles with smooth bases, smaller smooth echinating styles with smooth or microspined bases, and polytylote auxiliary tylostyles with smooth or microspined bases. Microscleres palmate isochelae, including contorted forms, and thick wing-shaped toxas. REMARKS. De Laubenfels (1936b) distinguished Fisherispongia from other microcionids by the polytylote bases on their subectosomal auxiliary styles. In all other respects, however, the type species resembles other encrusting species with hymedesmoid architecture (e.g., Leptoclathria). Polytylote ectosomal megascleres are known in several other species of Microcionidae (e.g., Clathria aceratoobtusa, Paratenaciella microxea), as well as in other Poecilosclerida (e.g., Camptisocale Topsent and Phellodenna Ridley & Dendy; Coelosphaeridae). The incorporation of detritus into the choanosome and fibres is well known for several microcionids and other sponges (see remarks for Aulenella) and on this basis the species is included in Clathria (Wilsonella). Re-examination of the holotype found a marked contrast in size between the smaller and larger (so-called principal) spicules, with no intermediate sizes, and these smaller spicules are interpreted here as being smooth echinating styles. By its toxa morphology Fisherispongia ferrea (from the Atlantic coast of Panama) is very similar to Clathria aceratoobtusa (from the Indo-west Pacific). Folitispa de Laubenfels, 1936 (Fig. 15D-F) Folitispa de Laubenfels, 1936a: 119. TYPE SPECIES. Hymedesmia laevissima Dendy, 1922:81 (by original designation) (holotype BMNH1921.11.7.69). Thickly encrusting growth form. Surface even, slightly hispid. Choanosomal skeleton hymedesmoid, with spongin fibres lying on substrate and bases of choanosomal principal subtylostyles embedded, standing perpendicular to substrate individually or forming short multispicular plumose columns protruding through surface. Echinating megascleres absent. Subectosomal skeleton irregularly plumose, with loosely aggregated bundles of subectosomal auxiliary subtylostyles erect on surface or lying tangential to it. Megascleres include smooth choanosomal principal subtylostyles, and smooth subectosomal auxiliary subtylostyles with mucronate or telescoped points. Microscleres palmate isochelae with anchorate-like modifications (curvature, partially detached alae, continuous ridge on shaft). Toxas absent. REMARKS. The type species of Foliti.spa differs from other encrusting microcionids with hymedesmoid skeletal architecture (e.g., Leptoclathria) in lacking echinating acanthostyles (cf. Axocielita) and having chelae with anchorate-like modifications instead of typical palmate isochelae (cf. Cionanchora). These chelae are strongly curved, with lateral alae partially detached from the shaft and a continuous lateral ridge running the length of the shaft. However, these lateral alae are not fully formed (being about two-thirds the size of the front ala), nor are they completely detached from the shaft (attached for approximately 50% of their length), and consequently they cannot be considered true anchorate chelae but perhaps palmate isochelae with substantial anchorate modifications. The genus is included here in Clathria (Microciona) based on its skeletal structure. [Halme] Lendenfeld, 1885 (Fig. 14H-I) Ha/me Lendenfeld, 1885c: 285, 1889a: 446; de Laubenfels, 1936a: 17; Bergquist, 1980b: 454; Wiedenmayer, 1989: 58 (preoccupied). Not Ha/me Pascoe, 1869. TYPE SPECIES. Holopsamma laminaefavosa Carter, 1885b: 212 (by subsequent designation; de Laubenfels, 1936a: 17) (holotype BMNH1886.12.15.312). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 46zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 15. Type species of microcionid genera. A-C, Fisherispongia (F. ferrea de Laubenfels, USNM22239). D-F, Folitispa (Hymedesmia laevissima Dendy, BMNH1921.11.7.69). G-I, Holoplocamia (H. penneyi de Laubenfels, USNM22460). REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 47 Massive, globular, lobate-digitate honeycombed reticulate growth form. Surface composed of small branches (lacunae') interconnected to form regular network. Choanosomal skeleton irregularly reticulate, with heavy spongin fibres fully cored by both sand particles and other detritus, and with fewer choanosomal principal subtylostyles both coring and echinating fibres. Subectosomal skeleton with peripheral fibres cored and echinated by principal megascleres, slightly heavier, more plumose at periphery than at core, and with subectosomal auxiliary strongyles forming irregular paratangential tracts near surface. Ectosome with external fibre reticulation reinforced by sand. Megascleres vary from common to relatively scarse (or spicules reportedly absent entirely in some specimens), including short entirely smooth choanosomal principal subtylostyles, and smooth sinuous or straight subectosomal auxiliary strongyles or quasidiactinal styles. Microscleres absent. REMARKS. Ha/me (sensu Lendenfeld, 1889b) is virtually identical to Aulena (of authors, e.g., Lendenfeld, 1888, but not Lendenfeld, 1885c), in skeletal construction, growth form and the presence of detritus within the fibre skeleton. In contrast, Hahne (of Lendenfeld, 1885c) differs from Aulena (of authors) by the virtual absence (or inconsistent presence) of proper, heavily mineralised spicules. However, examination of relevant type material, recent material from southern and eastern Australian waters (see below) and the literature (e.g., Wiedenmayer, 1989) shows that H. laminaefavosa is relatively polymorphic in its growth form, surface structure, spongin fibre construction, amount of detritus incorporated into the skeleton and the number and presence or absence of megascleres. Despite this variability, the species is clearly a synonym of Holopsamma, closely related to other honeycombed reticulate species. Of the numerous species referred to Hahne by Lendenfeld (1885c, 1888) many are Dictyoceratids (Bergquist, 1980b), whereas the type species is undoubtedly a microcionid. Unfortunately the name Halme Carter, 1885b is preoccupied by Hahne Pascoe, 1869 (Wiedenmayer, 1989), and Holopsamma Carter is the senior-most available name for this group of honeycomb reticulate microcionid sponges. Heteroclathria Topsent, 1904 (Fig. 16A-B) Heteroclathria Topsent, 1904b: 93; Burton, 1935a: 403. TYPE SPECIES. Heteroclathria hallez; Topsent, 1904b: 94 (by original designation and monotypy) (schizotype MNHNLBIMDT1884). Erect, digitate growth form. Surface even, microscopically hispid. Choanosomal skeleton regularly renieroid reticulate, with heavy spongin fibres well differentiated into primary ascending multispicular fibres, cored by choanosomal principal tylostyles, and secondary transverse uni- or bispicular fibres cored by amphistrongyles. Subectosomal skeleton plumose or paratangential, with subectosomal auxiliary subtylostyles forming poorly developed brushes arising from peripheral primary fibres, and also forming clusters around margins of oscules. Echinating acanthostyles absent, but choanosomal principal styles sometimes echinate primary fibres. Megascleres choanosomal principal tylostyles with microspined bases, large strongyles (dumbell spicules) of renieroid skeleton with spined bases, and subectosomal auxiliary subtylostyles with microspined bases. Microscleres palmate isochelae and thin wing-shaped toxas. REMARKS. Heteroclathria is unusual to the plocamiform group of sponges (de Laubenfels, 1936a), such as Antho and Plocamilla, in having a differentiated primary and secondary fibre network cored by monactinal and diactinal megascleres, respectively. The type species is only known from the holotype which unfortunately lacks collection data. Burton (1935a: 403) referred two other species to the genus, Plocamia karykinos de Laubenfels (1927: 262) and P. manaarensis (sensu Lambe, 1895: 124; holotype USNM6331; not Carter, 1880a: 34), which he renamed H. lambei Burton, and which Bakus (1966: 512) also renamed Plocamilla zhnmeri. De Laubenfels (1936a: 78) suggested that Heteroclathria was a synonym of Plocamia, and this is confirmed here from re-examination of both H. lambei and H. hallezi. Heteroclathria is referred here to Antho (Plocamia) in having (acantho)-strongyles in the renieroid skeleton, although true echinating spicules are absent. Another species which shows some similarities to Heteroclathria in skeletal structure is Stylotella corn uta Topent (1897b: 464), from the Andaman Sea off Malaysia, for which Burton & Rao (1932: 343) created Acanthostylotella. That species lacks the characteristic `dumbell spicules' but has MEMOIRS OF THE QUEENSLAND MUSEUM ^ 48zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA a renieroid skeleton of primary ascending multispicular fibres interconnected by unispicular tracts of smooth styles. It lacks an ectosomal skeleton and lacks microscleres, and it possibly best placed in Iophonidae. to Plocamione Topsent (Raspailiidae; Hooper, 1991), whereas the poecilosclerids (including Holoplocamia) are considered here to belong to Antho (Plocamia) (the latter a senior name for Plocamilla). Holoplocamia de Laubenfels, 1936 (Fig. 15G-I) Holopsamma Carter, 1885 (Fig. 16C-D) Holoplocamia de Laubenfels, 1936a: 75; Levi, 1960a: 80; Little, 1963: 47. TYPE SPECIES. Holoplocamia penneyi de Laubenfels, 1936a: 75 (by original designation) (holotype USNM22460). Thinly encrusting growth form. Surface rugose, microscopically hispid. Choanosomal skeleton hymedesmoid, with basal layer of spongin fibre, principal choanosomal styles embedded in fibre nodes, standing perpendicular to and protruding through surface, and with basal mass of acanthostrongyles forming an irregular renieroid secondary reticulation of spicules around principal spicules, interconnected by sparse collagen at nodes. Smaller acanthostyles also present echinating fibre nodes. Mesohyl incorporates large quantities of detritus and auxiliary spicules. Ectosomal skeleton with tangential or paratangential tracts of subectosomal auxiliary subtylostyles. Megascleres principal choanosomal styles-subtylostyles with either smooth or microspined bases, acanthostrongyles or acanthostyles of basal skeleton more heavily spined at ends than middle, echinating acanthostyles evenly spined, and subectosomal auxiliary subtylostyles with microspined bases. Microscleres palmate isochelae, including contorted forms, and wing-shaped toxas. REMARKS. Holoplocamia was erected for sponges similar to Plocamia Schmidt, but having spiny rather than smooth principal spicules. Levi (1960a) suggested that the genus was a synonym of Plocamilla, whereas Topsent (1928a) and Little (1963) argued that Plocatnilla was different from both Plocamia and Holoplocamia in lacking any differentiation between primary and secondary skeletal tracts. This opinion is not upheld here. De Laubenfels (1936a: 75) referred several `plocamiform' species to Holoplocamia, including the type species of Plocamilla, and it is now generally accepted that Holoplocamia and Plocamilla are synonymous (Bakus, 1966; Simpson, 1968a; Levi & Levi, 1983a; PulitzerFinali, 1983; Van Soest, 1984b). Most of the `plocamiform' species discussed by de Laubenfels (1936a) were subsequently found to belong Holopsamma Carter, 1885c: 211. TYPE SPECIES. Holopsamma crassa Carter, 1885c: 211 (by subsequent designation, de Laubenfels, 1936a: 98) (lectotype BMNH 1886.12.15.313; Hooper & Wiedenmayer, 1994), a senior synonym of Ha/me globosa Lendenfeld, 1885c: 303 (lectotype BMNH1886.8.27.71) (cf. Wiedenmayer, 1989: 63). Subspherical, digitate, regularly 'honeycomb' reticulate growth form. Surface arenaceous, porous, with tympanic membrane-like ectosomal crust stretched across adjacent subdermal cavities. Choanosomal skeleton reticulate, with well developed spongin fibres not well differentiated into primary or secondary elements, although many ascending fibres have core of small quantities of detritus (mostly spicule fragments), whereas other fibres clear of detritus completely. Coring and echinating spicules absent from choanosomal skeleton. Ectosomal skeleton heavily arenaceous, with crust of sand and scattered reticulate (or plumose or paratangential in places) bundles of subectosomal auxiliary strongyles lying tangential on surface crust. Mesohyl matrix relatively heavy between fibres . Megascleres only smooth subectosomal auxiliary strongyles. Microscleres absent. REMARKS. De Laubenfels (1936a: 97) noted that Holopsamma differs from other 'sandy sponges' (i.e., the polyphyletic 'family Psammascidae' de Laubenfels) in lacking microscleres and having both monactinal and diactinal megascleres. It is unfortunate that he designated H. crassa as the type species because in some of the 5 'valid' syntypes the monactinal (principal) styles may be lost completely, and the diactinal (auxiliary) strongyles are vestigial, leaving only heavy spongin fibres (the major ones with a core of detritus), and a heavy ectosomal sand cortex. The status and affinities of this genus are still confused, despite the comprehensive redescription and discussion of the type species by Wiedenmayer (1989). This confusion is due to the fact that no-one had previously nominated a lectotype amongst the 31 syntypes of the type species, which are composite and represent at REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 49 least 6 different species: only 5 of these actually conformed to Carter's (1885c) original description of Holopsamma crassa. Furthermore, the lectotype (BMNH1886.12.15.313) designated by Hooper & Wiedenmayer (1994) and figured by Wiedenmayer (1989: p1.6, fig.7) is identical to Ha/me globosa Lendenfeld, and there is some conjecture as to which name is the most valid. Wiedenmayer (1989: 63) chose to use the name H. globosa over H. crassa, although he admitted that crassa was more senior (apparently by only several months), and therefore under the rules of the ICZN it must take precedence. His arguments in choosing globosa over crassa were that type material of globosa was firmly established whereas the type series of crassa was an unresolvable mess, but this is inrelevant with the subsequent designation of a lectotype for H. crassa by Hooper & Wiedenmayer (1994). Wiedenmayer (1989: 63) provided many further details concerning these species synonymies and the affinities of 'sandy sponges' belonging to the Microcionidae. But more important than the nomenclatural problems associated with the type species (and the genera Ha/me and Holopsamma), there are some biological questions unanswered by Wiedenmayer's (1989) work. Holopsamma crassa is very similar to Holopsamma laminaefavosa, the type species of Ha/me Lendenfeld, 1885 (not Ha/me Pascoe, 1869), and it is possible that in fact the two species are synonymous (given that they are both allegedly very polymorphic). A comparison of type material shows that the only substantial differences are that in H. crassa spongin fibres contain virtually no sand and principal spicules have been lost, whereas in H. laminaefavosa primary fibres are virtually fully cored with sand, and both principal and auxiliary spicules are retained. In dealing with preserved material (i.e., without accompanying field characters), these definitions should be adhered to strictly. Conversely, following Wiedenmayer's (1989) definition, diagnostic characters in each species overlap substantially providing reasonable cause to synonymise the species (and genera Holopsamma and Ha/me). In the present work Holopsamma is the senior name for the group of honeycomb reticulate sponges traditionally known as Echinoclathria (of authors). Hymantho Burton, 1930 (Fig. 16F-G) Hymantho Burton, I930a: 503. TYPE SPECIES. Hymantho normani Burton, 1930a: 503 (by original designation) (holotypeBMNH 1910.1.1.791). Thinly encrusting growth form. Surface even, hispid. Choanosomal skeleton hymedesmoid, with basal layer of spongin on substrate and bases of choanosomal principal subtylostyles and echinating acanthostyles embedded and standing perpendicular to substrate. Ectosomal skeleton with paratangential bundles of subectosomal auxiliary subtylostyles of single size category. Mesohyl matrix with some debris incorporated. Megascleres choanosomal principal subtylostyles with acanthose bases, echinating acanthostyles with spined bases and shafts but aspinose points, and smooth auxiliary subtylostyles or polytylostyles. Microscleres palmate isochelae and thick forceps-shaped or v-shaped toxas. REMARKS. Hymantho normani was originally described with only toxa microscleres but reexamination of the holotype discovered that large palmate isochelae (18-22m), with large alae, are also present. Hymantho was erected by Burton (1930a), being similar to Leptoclathria Topsent in its hymedesmoid skeletal architecture, but supposedly lacking chelae microscleres (Levi, 1960a: 60). In this latter respect the genus was also be compared with Pseudanchinoe and QuercicIona, which Van Soest (1984b) considered to be junior synonyms of Clathria (s.1.), but in any case the discovery of isochelae in the type species negates the concept of the genus. Alander (1942) and Van Soest & Stone (1986) also suggested that the secondary loss of isochelae and the presence of a leptoclathriid skeleton have little generic value in the Microcionidae. The genus is referred into synonym with Clathria (Microciona). Isociella Hallmann, 1920 (Fig. 16H-I) Isociella Hallmann, 1920: 784; de Laubenfels, 1936a: 152; Bergquist & Tizard, 1967: 187; Bergquist & Fromont, 1988: 114. TYPE SPECIES. Phakellia flabellata, in part (sensu Ridley & Dendy, 1886: 478) (by monotypy) (not P. flabellata Carter, 1885f: 363); =Phakellia jacksoniana Dendy (replacement name; Dendy, 1897: 236) (holotype BMNH1887.5.2.9), both junior synonyms of Clathria macropora, in part, Lendenfeld, 1888: 221 (holotype AMZ466) (this work). Erect, stipitate, flabelliform growth forms. Surface hispid, relatively smooth, with ridges and lamellae. Choanosomal skeleton relatively homogeneous, renieroid, composed of primary, multispicular, plumose, ascending tracts of ^ MEMOIRS OF THE QUEENSLAND MUSEUM 50zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 16. Type species of microcionid genera. A-B, Heteroclathria (H. hallezi Topsent, MNHNDT1884). C-D, Holopsamma (H. crassa Carter,BMNH1886.12.15.313). E, Ha/me (H. globosa Lendenfeld, synonym of H. crassa, BMNH1886.8.27.71). F-G, Hymantho (H. normani Burton, BMNHI910.1.1.791). H-I, Isociella (Phakelliaflabellata sensu Ridley & Dendy, BMNH1887.5.2.9). ^ 51 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR choanosomal principal styles, interconnected by secondary, uni- or paucispicular, transverse tracts of same spicules, together forming regular renieroid or sub-renieroid (triangular) meshes; principal spicules bonded together at nodes by collagen or enclosed in relatively poorly developed spongin fibres. Echinating acanthostyles absent. Subectosomal auxiliary styles of a single category, arranged tangentially, paratangentially or in plumose brushes on surface, with some principal styles also protruding through peripheral skeleton singly or in sparse plumose brushes. Megascleres robust smooth choanosomal principal styles, and small smooth or basally spined subectosomal auxiliary stylessubtylostyles. Microscleres palmate isochelae with 'fluted' alae, no toxas (in type species). REMARKS. Isociella contained 3 species prior to the present study (Clathria macropora Lendenfeld, Ophlitaspongia eccentrica Burton (1934a: 560), and !. incrustans Bergquist (1961a: 42)), although Hallmann (1920: 784) suggested that there were some other species included in his concept of Ophlitaspongia (=Echinoclathria as defined in this study) which could also be referred here. The definition above is widened below to include species with toxas and different forms of palmate isochelae. Dendy (1897) renamed the type species jacksoniana because Phakellia flabellata Ridley & Dendy (1886) was preoccupied by Phakellia flabellata Carter (1885f) from Port Phillip Bay, but Hallmann (1920) considered that the 2 species belonged to different genera, and the replacement name flabellata was unnecessary. However, P.flabellata Ridley & Dendy is a junior homonym of P. flabellata Carter, and Dendy's (1897) replacement name P jacksoniana is a valid emendment. Neither species belongs to Phakellia. In any case C. macropora Lendenfeld is identical to, and the senior available name for, P jacksoniana. Isociella is distinguished from other microcionids by its relatively homogeneous, wide-meshed, sub-renieroid reticulation of a single category of smooth choanosomal styles, lacking echinating spicules, and without any differentiation between axial and extra-axial regions (although choanosomal spicules may diverge slightly towards periphery, sometimes becoming plumose on surface). It differs from the two other groups of microcionids that have renieroid skeletal structure. Antho (including Isociona, Plocamia, Plocamilla, Plocamiopsis, and Isopenectya), has two categories of choanosomal megascleres, one acanthose forming a basal renieroid skeleton, and one smooth forming ascending plumose extra-fibre tracts of the true choanosomal skeleton. Some Echinoclathria species (as defined in the present study, including Ophlitaspongia in the sense of most authors), have a renieroid main skeleton of smaller, smooth choanosomal styles and the same spicules echinating fibres, a radial peripheral skeleton in which larger, smooth choanosomal styles are embedded in the subectosomal skeleton and protrude a long way through the surface, and a very well developed spongin fibre reticulation seen in most species, often with some axial compression (reflecting a digitate or flabellate growth form). The most abundant Australian species, Isociella eccentrica, has a choanosomal skeleton reminiscent of Callyspongia (Haplosclerida) and an open, reticulate ectosomal skeleton, and all 3 known Australian species have closest affinities with certain Clathria species (e.g., C. (Clathria) conectens, C. (Thalysias) hirsuta). Thus, the definition of the type species is expanded below to include forms which are predominantly semiencrusting, branching, with rugose reticulate surface sculpturing, well developed spongin fibres (as opposed to only having collagen binding principal spicules together); poorly differentiated primary and secondary skeletal tracts, and to allow for the presence of toxa microscleres. Isociella is included here as a subgenus of Clathria. Isociona Hallmann, 1920 (Fig. 17A-B) Isocionu Hallmann, 1920: 768; de Laubenfels, 1936a: TYPE SPECIES. Lissodendoryx tuberosa Hentschel, 1911: 326 (by monotypy) (holotype ZMB4417). Thickly encrusting, bulbous growth form. Sur- face irregularly microconulose, hispid. Choanosomal skeleton with differentiated primary and secondary skeletons. Secondary skeleton renieroid or subrenieroid reticulate, with acanthose styles forming uni-, pauci-, or less frequently multispicular tracts, bound together by very light fibres or collagen at spicule nodes. Primary skeleton plumose, with choanosomal principal styles forming larger primary ascending tracts irregularly connected by smaller secondary transverse tracts. Subectosomal skeleton plumose, with principal styles protruding through MEMOIRS OF THE QUEENSLAND MUSEUM ^ 52zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ectosome overlayed by erect bundles of subectosomal auxiliary subtylostyles. Echinating megascleres absent. Megascleres large smooth choanosomal principal styles, basally spined or entirely lightly spined styles-subtylostyles of renieroid skeleton, and smooth auxiliary subectosomal subtylostyles. Microscleres palmate isochelae and thick wing-shaped toxas. REMARKS. Isociona tuberosa and Isociella eccentrica appear very similar from published descriptions whereas examination of type material shows that they are different. Isociella eccentrica has a wide meshed renieroid reticulate skeleton composed of smooth principal spicules, whereas I. tube rosa has a close-meshed renieroid secondary skeleton of spined monactinal spicules, overlayed by a plumose primary skeleton of smooth principal spicules. Van Soest (1984b) merged lsociona with Antho, although this relationship is not straightforward given that Antho is usually reserved for forms with diactinal megascleres (in the renieroid basal skeleton). Nevertheless, the two taxa have similar skeletal architecture and Isociona is maintained as a synonym of Antho herein. Isopenectya Hallmann, 1920 (Fig. 17C-D) Isopenectya Hallmann, 1920: 789; de Laubenfels, 1936a: 125. TYPE SPECIES. Clathria (?) chartacea Whitelegge, 1907: 497 (by monotypy) (holotype AMZ436). Thinly flabellate-lamellate growth form. Surface smooth, even. Choanosomal skeleton renieroid reticulate, with differentiated axial and extra-axial regions of skeleton. Axial skeleton with compressed spongin fibres running through centre of lamellae, cored by smooth choandsomal principal styles (marginally smaller than those in surface bundles) forming subisodictyal tracts, overlain by renieroid skeleton of small spined styles. Extra-axial skeleton with more openmeshed, regularly reticulate spongin fibres, cored by uni- or bispicular renieroid tracts of small acanthose styles, and plumose, subisodictyal tracts of longer smooth styles standing perpendicular to axis, both fully enclosed in spongin fibres. Echinating megascleres absent. Subectosomal skeleton plumose, with bundles of smooth choanosomal principal styles protruding through surface and tangential or paratangential tracts of subectosomal auxiliary styles. Ectosome lacks specialised spiculation. Megascleres lightly acanthose styles-subtylostyles forming renieroid skeleton, short and long smooth choanosomal principal styles, and subectosomal auxiliary styles with spined bases. Microscleres absent. REMARKS. Isopenectya is similar to other renieroid `plocamiform' genera (sensu de Laubenfels, 1936a), Antho, Plocamilla, Plocamiopsis, Isociona, Labacea, Panda ros, lsociella and Echinoclathria, in having a primarily renieroid reticulate skeleton. Unlike these other genera, however, Isopenectya has 2 forms of choanosomal megascleres enclosed within spongin fibres. The smaller acanthose styles forming the renieroid structure, whereas smooth styles form the subisodictyal, mostly longitudinal skeleton. These latter spicules also produce the extra-axial plumose tracts that ascend to the surface, and larger, smooth choanosomal styles produce brushes on the ectosome. Although the fibre skeleton is compressed in the axis the renieroid skeleton is barely different between axial and extra-axial regions. Isopenectya chartacea is remarkable in that it closely resembles the type species of Echinoclathria (E. leporina) in growth form, gross skeletal architecture and fibre characteristics. Whereas E. leporina has one category of relatively homogeneous smooth principal styles throughout the choanosomal skeleton, I. chartacea has 2 differentiated structures within the choanosome (renieroid and plumose or subisodictyal skeletons) and 2 geometrically different categories of megascleres forming these skeletal structures. Skeletal architecture is also more regularly renieroid and renieroid fibres/tracts are relatively homogeneous than in most Echinoclathria species, but this is a matter of degree. Echinoclathria has a radial skeleton of larger, smooth principal styles embedded in peripheral fibres, poking through the surface, and it could be argued that this is a vestigial subisodictyal skeleton like that in Isopenectya. But this similarity is inferred and any relationship is equivocal, with emphasis placed here on the possession of spined spicules of the renieroid skeleton and clearly differentiated renieroid and subisodictyal skeletal structures in deciding on affinities of Isopenectya. Nevertheless, these 2 species are certainly remarkably similar and potentially may be confused. Van Soest (1984b) suggested that Isopenectya may be valid, but `plocamiform' microcionid type shows that it is a close relative of this group, of Antho in particular, differing from it only in REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 53 having a unique subisodictyal skeleton of smooth spicules overlaying the renieroid basal skeleton. It is recognised here as a subgenus of Antho. TYPE SPECIES. Clathria juncea sensu Burton, 1931a: 343 (by original designation) (type fragment BMNH1926.2.19.2). Erect digitate, arborescent growth form. Sur- Jia de Laubenfels, 1930 face even, minutely hispid. Choanosomal skeleton reticulate, with well differentiated primary ascending and secondary transverse skeletal tracts. Primary skeleton with well developed thick spongin fibres, ascending to surface, cored by multispicular tracts of choanosomal principal styles, interconnected by thin transverse secondary fibres cored by paucior unispicular tracts of same spicules. Echinating acanthostyles moderately common in axial skeleton, very heavy in peripheral skeleton, forming plumose brushes at surface. Subectosomal skeleton tangential with subectosomal auxiliary subtylostyles lying on surface. Ectosome without specialised spiculation, but with many foreign spicule fragments embedded in outer layer of skeleton. Mesohyl with abundant detritus and auxiliary spicules dispersed between fibres. Megascleres include both smooth and completely lightly spined fusiform choanosomal principal styles, evenly spined echinating acanthostyles, and smooth auxiliary subectosomal subtylostyles. Microscleres large palmate isochelae and accolada toxas with microspined points. (Fig. 17E-F) Jia de Laubenfels, 1930: 28, 1932: 97. TYPE SPECIES. Jia jia de Laubenfels, 1930: 28 (by original designation) (holotype USNM21510). Encrusting growth form. Surface uneven, hispid. Choanosomal skeleton confused renieroid reticulate, composed of smaller smooth or acanthose styles forming basal more-or-less rectangular network, overlayed by larger smooth principal styles standing erect, forming ascending bundles or single spicules projecting through surface. Subectosomal auxiliary subtylostyles paratangential to surface. Ectosome without special spicules. Megascleres larger smooth choanosomal principal styles, smaller smooth or acanthose styles-subtylostyles of basal skeleton, and subectosomal auxiliary subtylostyles with basal spines. Microscleres palmate isochelae, modified J-shaped chelae resembling sigmas (= `crocae'), and wing-shaped toxas with spinous extremities. REMARKS. De Laubenfels (1932) description of the type is incomplete. The essential specific characteristics include the modified (J-shaped) isochelae (crocae of Van Soest & Stone, 1986), and a renieroid albiet ill-defined reticulation. Those characters are contrasted with the Antholike nature of closely related A. (Jia) brattegardi Van Soest & Stone, which has acanthose monactinal and diactinal spicules in the main skeleton, whereas Jia (s.s.) has predominantly smooth spicules. Despite these differences, Van Soest & Stone (1986) justifiably merged Jia and Antho because many other Antho-like species without crocae also have predominantly smooth styles and poorly defined skeletal construction (e.g., A. dichotoma (Esper)). Jia is referred to Antho (Ant ho). Crocae are not unique to Jia; other taxa with similarly modified chelae are Dendoryx luciensis Topsent (Myxillidae) and Zygherpe hyaloderma de Laubenfels (Hamacanthidae) (de Laubenfels, 1932; Bakus, 1966; Van Soest & Stone, 1986. Labacea de Laubenfels, 1936 (Fig. 17G-H) Labacea de Laubenfels, 1936a: 125. REMARKS. Clathria juncea is attributed to Burton (1931a), and a lectotype designated from the syntypes (BMNH1933.7.4.4-7). However, Burton and de Laubenfels (1936a) suggested that Clathria juncea sensu Burton may be synonymous with Alcyonium junceutn Lamarck, 1816. Topsent (1933: 26) merged Lamarck's species as a variety of Anomoclathria opuntioides, but noted that the type was missing from the MNHN. It is still missing, and must now be presumed destroyed. If Lamarck's species is identical with A. opuntioides then C. juncea sensu Burton is quite different (see remarks for Anomoclathria). Burton's (1931a) description, and de Laubenfels' (1936a) interpretation of the type species are both erroneous. Burton's described material exhibits several characters not noted by either of these authors, so that Labacea is incorrectly diagnosed. The skeletal architecture of C. juncea is reticulate, not renieroid as supposed by de Laubenfels, and echinating acanthostyles (a prominent feature of the peripheral skeleton in particular) and palmate isochelae were overlooked. I assign it to Clathria (Clathria). De Laubenfels (1936a) assigned Clathria oxeifera Ferrer Hernandez to this genus; it sup- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 54zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 17. Type species of microcionid genera. A-B, lsociona (Lissodendoryx tube rosa Hentshel, ZM B4417). C-D, Isopenectya (Clathria chartacea Whitelegge, AMZ436). E-F, Jia (J. jia de Laubenfels, USNM21510). G-H, Labacea (Clathria juncea sensu Burton, BMNH1926.2.19.2). I, Leptoclathria (L haplotoxa Topsent, MNHNDT1101). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 55 posedly differed from the type species in having echinating acanthostyles (as distinct from acanthose varieties of choanosomal principal megascleres (Levi, 1960a: 84)) but as noted above these spicules are typical for the genus, and de Laubenfels' observation is redundant. Leptoclathria Topsent, 1928 (Figs 171, I8A) Leptoclathria Topsent, 1928a: 298. TYPE SPECIES. Leptoclathria haplotoxa Topsent, 1928: 298 (by monotypy) (holotype MNHNLBIMDT1101). Encrusting growth form. Surface irregular, hispid. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer of spongin lying on substrate, with bases of choanosomal principal subtylostyles and abundant smaller acanthostyles standing perpendicular to substrate. Subectosomal skeleton with light tangential tracts of subectosomal auxiliary subtylostyles occurring in bundles or individually, forming irregular dermal brushes surrounding protruding choanosomal megascleres. Ectosomal skeleton without specialised spiculation. Megascleres include entirely acanthose choanosomal principal subtylostyles, evenly spined echinating acanthostyles, and subectosomal auxiliary subtylostyles with spined bases and telescoped or mucronate points, sometimes quasidiactinal. Microscleres palmate isochelae and toxas intermediate between wing-shaped and v-shaped. REMARKS. The type species is simply a thinly encrusting Microciona-like species with a hymedesmoid skeletal architecture, but which differs from Microciona (s.s.) in having entirely acanthose principal styles (similar to Dictyociona,Hymeraphia and Anaata). Levi (1960a) proposed that Leptoclathria should be merged with Microciona, and this is certainly true for the type species (referred here to C. (Microciona)), but there are some species which have a clearly differentiated ectosomal and subectosomal auxiliary spicules (e.g., L. lambda Levi), and these species are more appropriately included in C. (Thalysias). Topsent (1928a) overlooked the spined bases and telescoped (or mucronate) points on subectosomal auxiliary subtylostyles which are prominent in the holotype. Furthermore, and of greater taxonomic significance, there are at least a small proportion of auxiliary megascleres with tylote (quasidiactinal) geometry, including basal microspination, which is reminiscent of the specialised diactinal ectosomal megascleres characteristic of Myxillidae and Iophonidae. Only a few other microcionids have this feature, C. (Clathria) chelifera, C. (Wilsonella) australiensis and C. (Thalysias) major (see also Hooper et al., 1990). These quasi-diactinal spicules are analogous to (but not homologous with) ectosomal tylotes found in Myxillidae for example (see remarks for Acamus below). Ligrota de Laubenfels, 1936 (Fig. 18B-C) Ligrota de Laubenfels, 1936a: 125. TYPE SPECIES. Clathria lobata Vosmaer, 1880: 151 (by original designation) (holotype RMNH276). Arborescent growth form with flattened branches. Surface even, microscopically hispid. Choanosomal skeleton regularly reticulate, with poorly developed spongin fibres forming differentiated primary and secondary tracts. Primary fibres ascend to surface, cored by plumose multispicular tracts of choanosomal principal styles; secondary fibres transverse, paucispicular, cored by same spicules. Subectosomal skeleton plumose, with bundles of choanosomal principal megascleres extending into ectosome and protruding through surface. Echinating acanthostyles most abundant in periphery, below surface spicule brushes. Some detritus incorporated into mesohyl, scattered between spicule tracts. Ectosome radially arranged, erect brushes of subectosomal auxiliary subtylostyles of a single size class. Megascleres robust choanosomal principal styles-subtylostyles with smooth bases, echinating acanthostyles with large spines and aspinose points, and subectosomal auxiliary subtylostyles with basal spines. Microscleres palmate isochelae and wing-shaped toxas with spined points. REMARKS. This definition from the holotype differs from descriptions provided by Vosmaer (1880), Ridley & Dendy (1887), and Levi (1963), but corresponds with Stephens's (1915) concept of the species. In particular, the species has a regularly reticulate skeleton with radial architecture, but it lacks a well differentiated axial and extra-axial region as suggested by these authors. Ligrota has spiculation virtually identical to that of Clathria (s.s.), including spines on the points of toxas. It is not a Thalysias, as supposed by Ridley & Dendy (1887), because it has only a MEMOIRS OF THE QUEENSLAND MUSEUM 56^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA single, undifferentiated category of auxiliary subtylostyle and is referred here to Clathria (Clathria). Similarly, de Laubenfels' (1936a) diagnosis of Ligrota is incorrect. Clathria lobata of Stephens (1915) does not have diactinal megascleres or sigmas. Those attributes were described by Vosmaer (1880) for the type specimen, but were discounted by Ridley & Dendy (1887) as being probable contaminants; nor were they observed in type material. Lissoplocamia Brondsted, 1924 (Fig. 18D-E) Lissoplocamia Brondsted, 1924: 470. TYPE SPECIES. Lissoplocamia prima Brondsted, 1924: 470 (by original designation) (holotype in ZMC, not seen; MNHNLBIMDCL637 from South Africa). Digitate, arborescent, flattened or cylindrical branches. Surface prominently hispid, velvetty. Choanosomal skeleton renieroid reticulate, with differentiated axial and extra-axial regions. Axial skeleton slightly compressed, with well developed spongin fibres cored by uni- or paucispicular tracts of tylotes forming renieroid reticulation, overlaid by plumose (or plumoreticulate) extra-axial skeleton composed of diverging single or multiple choanosomal principal styles, echinating (protruding from) spongin fibres and ascending to surface. Spongin fibres in extra-axial skeleton lighter, more-widely spaced. Subectosomal skeleton plumose, with bundles of subectosomal auxiliary styles perched on ends of principal megascleres. Ectosome without special category of megascleres, but isochelae microscleres predominant in peripheral skeleton. Megascleres tylotes ('dumbell-shaped' spicules) of renieroid skeleton with swollen microspined bases, choanosomal principal styles-subtylostyles, with smooth or faintly microspined bases, and smooth subectosomal auxiliary styles. Microscleres palmate isochelae and wing-shaped toxas. REMARKS. This definition is based on a recent redescription of the holotype from New Zealand (Bergquist & Fromont, 1988: 122) and Levi's (1963) specimen from South Africa. Levi's material is more complete, with an intact ectosomal skeleton and containing palmate isochelae, whereas Brondsted's (1924) holotype is poorly preserved, lacking an ectosomal skeleton in which the isochelae are predominantly found (Bergquist & Fromont, 1988). Lissoplocamia belongs to de Laubenfels' (1936a) `plocamiform' group of sponges in having diactinal, quasi-diactinal or monactinal `dumbell-shaped' spicules forming a renieroid basal skeleton (it should be noted that this character is different from (analogous to) similar 'sausage-shaped' diactinal spicules found in the ectosomal skeletons of some genera in families such as Coelosphaeridae and Petrosiidae). Some of these `plocamiform' genera belong to Raspailiidae (Hooper (1991: 1319)), whereas others are closely related to Microcionidae and are referred to Antho (Plocamia). Bergquist & Fromont (1988: 122) mentioned several of these genera (Lissoplocamia, Holoplocamia, Plocamilla, Heteroclathria, and Plocamia), and to this group should also be added Antho and Jia. Axoplocamia, included by them with the microcionids, was shown to be a raspailiid (Hooper, 1991). Bergquist & Fromont (1988) included only 2 New Zealand species in Plocamia, Dirrhopalum novizelanicum Ridley and L. prima, both of which are referred here to Antho. Lissoplocamia differs from most other `plocamiform' microcionids, such as Antho (Antho) in having predominantly smooth 'clumbell-shaped' spicules, similar to the type species of Plocamia, P gymnazusa, but this character is considered insignificant at the generic level given its large variability within the family. Litaspongia de Laubenfels, 1954 (Fig. 18F-G) Litaspongia de Laubenfels, 1954: 162. TYPE SPECIES. Ophlitaspongia arbuscula Row, 191 1: 347 (by original designation) (holotype BMNH1912.2.1.63). Convoluted arborescent growth form. Surface irregularly conulose, arenaceous, minutely hispid. Choanosomal skeleton regularly reticulate, with heavy spongin fibres cored by uni- or paucispicular tracts of choanosomal principal subtylostyles (sometimes fibres aspiculose). Echinating acanthostyles sparse. Subectosomal skeleton plumose, with dense bundles of principal styles (identical to coring spicules) diverging from ends of peripheral fibres and forming discrete brushes on surface. Ectosomal skeleton with sparse subectosomal auxiliary subtylostyles tangential to surface and also dispersed throughout mesohyl. Megascleres thin smooth choanosomal principal subtylostyles, smooth quasi-diactinal subectosomal auxiliary subtylostyles (usually resembling asymmetrical strongyles), and evenly spilled echinating acanthostyles. MICIDscleres wing-shaped toxas. Isochelae absent. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 57 REMARKS. De Laubenfels (1954) stated that Litaspongia was established for sponges like Echinoclathria (as defined here) in having monactinal megascleres, toxa microscleres and arborescent growth form. The holotype also has echinating acanthostyles, previously overlooked by Row (1911). The resemblance between 0. arbuscula and Echinoclathria is here considered superficial, based on the fact that principal spicules in 0. arbuscula are thin, attaining only the thickness typical of auxiliary spicules found in most other Clathria species; auxiliary spicules are quasidiactinal in 0. arbuscula, resembling diactinal ectosomal spicules in some species of Echinoclathria (e.g., E. chalinoides) and Holopsamma (e.g., H. ramosa); and spongin fibres are regularly reticulate, well developed, and tend to dominate skeletal structure over spicule components. But unlike Holopsamma and Echinoclathria which have undifferentiated coring and echinating spicules, Litaspongia has different principal spicules coring fibres from those echinating fibres. I consider it a reduced Clathria (Clathria). De Laubenfels (1954: 162) synonymised 0. arbuscula and 0. horrida Row (1911: 349), and re-examination of both holotypes (the latter BMNH1912.2.1.65) supports this decision. He added Echinoclathria nodosa (which he merged with E. subhispida) to Litaspongia, but both are species of Echinoclathria. Pulitzer-Finali (1982: 105) referred 0. arbuscula and 0. horrida to Kerasemna (Desmacellidae) (Hooper, 1984b), but neither have sigma microscleres and this placement was unjustified. Marleyia Burton, 1931 (Fig. 18H-I) Marleyia Burton, 1931a: 346; de Laubenfels, 1936a: 109. TYPE SPECIES. Marleyia irregularis Burton, 1931a: 346 (by original designation) (holotype NM 1279). Digitate growth form, flattened branches. Sur- face uneven, porous, with specialised reticulate external fibrous skeleton. Choanosomal skeleton reticulate, with subisodictyal reticulation of well developed primary and secondary spongin fibres. Primary fibres ascending, cored by multispicular tracts of both choanosomal principal subtylostyles and subectosomal auxiliary subtylostyles. Fibres heavily echinated by short acanthostyles. Secondary fibres transverse, aspicular, but of similar diameter and density of echinating spicules as primary fibres. Abundant detritus scattered through mesohyl, but not incorporated into fibres. Ectosome with tangential reticulation of spongin fibres, more closely reticulate and slightly thinner than choanosomal fibres, lightly cored by both subectosomal and choanosomal subtylostyles and echinated by abundant acanthostyles. Megascleres smooth robust choanosomal principal subtylostyles, smooth straight or sinuous subectosomal auxiliary subtylostyles, and robust echinating acanthostyles with bare neck. Microscleres absent. REMARKS. The holotype is dry and lacks a well preserved ectosomal skeleton). Burton suggested that Marleyia differs from all other microcionid genera in having a special ectosomal skeleton formed by a tangential reticulation of spongin fibres, which are thinner and more closely compacted than choanosomal fibres. This feature was not as remarkable as we are led to believe from the original description (although the holotype is not well preserved), nor is it unique in the Microcionidae (also seen in Echinochalina (Protophlitaspongia) labouti Hooper & Levi) or other families (e.g., Callyspongiidae (Haplosclerida)). In gross morphology and fibre characteristics Marleyia is similar to several Holopsamma species, but it has different spicules coring and echinating spicules (whereas Holopsamma has only one category of principal spicule performing these functions). Burton (1931a) made further casual comparisons between Marleyia and certain Dictyoceratida, based on fibre characteristics and a greatly reduced skeleton. De Laubenfels (1936a: 109) suggested that Marleyia may have affinities with Acantheurypon, representing a more mature form of that genus, but he noted that Marleyia had a remarkable external resemblance to `keratose' sponges. On the basis of its unusual ectosomal fibre characteristics Van Soest (1984b) suggested that Marleyia might be a valid genus of Microcionidae, but re-examination of type material indicates that it clearly belongs to Clathria (Clathria). Marleyia is monotypic, and known only from the Durban region, Natal coast, South Africa. Microciona Bowerbank, 1862 (Fig. 19F-G) Microciona Bowerbank,1862b: 1109; Topsent, 1894a: 18; Dendy, 1922:60; Topsent, 1928a: 62; Vosmaer, 1935a: 604 (in part); de Laubenfels, 1936a: 447; Levi, 1956b: 399; Koltun, 1959: 181; Burton, 1959a: 225; Levi, 1960a: 51; Sara, 1963: 210, 1964: ^ MEMOIRS OF THE QUEENSLAND MUSEUM 58zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 18. Type species of microcionid genera. A, Leptoclathria (L haplotoxa Topsent, MNHNDT1101). B-C, Ligrota (Clathria lobata Vosmaer, RMNH276). D-E, Lissoplocamia (L. prima Brondsted, MNHNDCL637). F-G, Litaspongia (Ophlitaspongia arbuscula Row, BMNH1912.2.1.63). H-I, Marleyia (M. irregularis Burton, NM1279). 59 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 230; Bergquist, 1965: 168; Hechtel, 1965: 41; Simpson, 1968a: 93, 102; Levi, 1973: 613; Wiedenmayer, 1977: 140; Bergquist, 1978: 172; Bergquist & Fromont, 1988: 100. TYPE SPECIES. Microciona atrasanguinea Bowerbank, 1862b: 1109 (by subsequent designation (Bowerbank, 1864: 188)) (holotype BMNH 1930.7.3.225). Encrusting growth form. Surface hispid, uneven. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer lying on substrate, bearing erect, non-anastomosing, rarely branching, scattered fibre nodes perpendicular to substrate ('microcionid' fibres), each cored by plumose ascending columns of choanosomal principal subtylostyles, wholly or partly embedded in fibres, with points of spicules usually projecting through ectosome. Echinating acanthostyles also erect on fibre nodes. Subectomal skeleton with tangential layer of subectosomal auxiliary subtylostyles, singly or in bundles on surface. Ectosomal skeleton without specialised spiculation, but choanosomal and subectosomal spicules protude through surface. Megascleres choanosomal principal subtylostyles with smooth or microspined bases, evenly spined echinating acanthostyles, and smooth or basally spined subectosomal auxiliary subtylostyles. Microscleres palmate isochelae and wing-shaped toxas. REMARKS. In many publications, Clathria Schmidt and Microciona Bowerbank have the date 1862. In merging the two genera, Wiedenmayer (1977, 1989) and Van Soest (1984b) note that the former has priority, because Bowerbank's work was not officially published until 1863. This ddefinition of Microciona is compiled from description of the type species (Bowerbank, 1862b, 1864, 1866) and histological preparations of the holotype and paratypes (BMNH1910.1.1.68, 1930.7.3.226). Microciona (s.s.) differs from Clathria (s.s.) in its encrusting growth form, a reduced hymedesmoid skeleton with erect fibre nodes cored by plumose tracts of principal and echinating spicules standing erect on the substrate (= `microcionid' architecture of Levi, 1960a), and smooth toxas. The critical difference between these genera, therefore, is the possession of the plumose, non-anastomosing fibre nodes, whereas cofamilial encrusting genera have hymedesmoid skeletal construction (Leptoclathria, Anaata, Cionanchora, Hymantho) or a basal renieroid reticulation (e.g., some Antho, Plocamilla). By comparison, erect or massive Microcionidae commonly have reticulate or plumo-reticulate skeletons (Clathria, Holopsam- ma), renieroid or subisodictyal skeletons (e.g., Isociella, Isopenectya, Pandaros), or virtually halichondroid skeletal architecture (Artemisina). Many authors maintain the distinction between Microciona and erect non-plumose genera, such as Clathria (e.g., Levi, 1969; Wiedenmayer, 1977; Pulitzer-Finali, 1983; Uriz, 1984a-b; Wintermann-Kilian & Kilian, 1984; Boury-Esnault & Lopes, 1985; Bergquist & Fromont, 1988). Cytological evidence to support this argument (Simpson, 1968a) is difficult to reconcile completely with other morphological features. Moreover, only a few species were examined in this context, and the cytological characters themselves are obscure, not having been tested subsequently in other taxa. Conversely, no evidence was found from biochemistry (protein electrophoresis, free amino acid or carotenoid protein profiles) to support the differentiation of encrusting (hymedesmoid or microcionid) species from erect (reticulate, non-plumose) species (Hooper et al., 1992), but once again only few taxa were examined in this context. Thus, I have little clear evidence from non-skeletal sources on the boundary between these genera. Overlying this classification based on skeletal structure many authors subdivide Microcionalike species groups (at the generic level) according to modifications in microsclere geometry, ornamentation on megascleres, or further reductions in skeletal structure (e.g., Anaata, Axociella (of authors), Axocielita, Dictyociona, Leptoclathria, Pseudanchinoe, Thalyseurypon; e.g., Little, 1963; Koltun, 1976; Alcolado, 1980; Hoshino, 1981; Lee & Gilchrist, 1985; Sim & Bakus, 1986). Undoubtedly, recognising all these genera is a matter of pragmatism, as this scheme offers a convenient and relatively quick means to manage a large and diverse assemblage of microcionids. Many authors have questioned the validity of these genera, debating generic distinction between Microciona and Clathria(Alander, 1942; Hechtel, 1965; Sara & Melone, 1966; Van Soest, 1984b; Wiedenmayer, 1989; Hooper, 1990a). Some empirical evidence supports counterarguments, that these character states intergrade between these genera (ontogenetically and/or phenotypically), but no consensus has yet been reached. Van Soest (1984b) and Hooper (1990a) questioned separation of encrusting genera with hymedesmoid or `microcionid' skeletons (e.g., Microciona) from erect growth forms with reticulate or plumoreticul ate skeletons (e.g., Clathria), because they are clearly linked to some extent by ^ MEMOIRS OF THE QUEENSLAND MUSEUM 60zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 19. Type species of microcionid genera. A-B, Megaciella (Amphilectus pilosus Ridley & Dendy, BMNH1887.5.2.125). C-E, Melonchela (M. clathrata Koltun, BMNH1963.7.29.7). F-G, Microciona (M. atrasan guinea Bowerbank, BMNH1930.7.3.438). H-I, Naviculina (N. cliftoni Gray, BMNH1877.5.21.270). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 61 the ontogeny of the sponge individual (Simpson, 1968a). They argued from a phylogenetic basis that the recognition of this character as being a primary one confers exceptionally high levels of homoplasy within the classification, cutting across lines of apparent evolutionary decent based on a combination of other characters (such as the origin and disposition of structural megascleres within the skeleton, spicule geometry). For example, there are many thinly encrusting species with hymedesmoid architecture and plumose spongin fibre nodes referable to Microciona (e.g., M. maunaloa de Laubenfels, M. microchela Hechtel), but also having two differentiated categories of auxiliary spicules (i.e., a specialised ectosomal skeleton), which is characteristic of Thalysias species. Which genus do these species then belong to, Microciona or Thalysias, or do we recognise a third genus because they have both characters ? This third alternative was adopted by Bergquist & Fromont (1988), largely following de Laubenfels' (1936a), who recognised Axociella for microcionids with Microciona-like fibre skeletons and Thalysiaslike ectosomal skeletons (although, like de Laubenfels (1936a), they misinterpreted Axociella which is neither encrusting nor hymedesmoid). This is a 'convenient classification' for managing species, but it is also responsible for most of the 70 or so microcionid genera that exist today (i.e., approximately 1 genus per 6 species). Few authors agree on the level of taxonomic divergence, or the phylogenetic interpretation of character polarity: is encrusting growth form and hymedesmoid architecture a primitive or derived strategy? Is a `microcionid' architecture (hymedesmoid with ascending plumose fibre nodes) a subsequent developmental stage of hymedesmoid architecture (spongin fibres flat against the substrate)? It is anticipated that evidence from biochemistry or genetic sources will eventually contribute towards the resolution of these conflicts. A resolution is provided here based on phylogenetic interpretation of morphology. We follow Levi's (1960a) arguments in considering three alternatives. 1) One generic taxon may be used to encompass all the skeletal types from leptoclathriid (hymedesmoid), `microcionid', to reticulate. This option was used by Vosmaer (1933, 1935a-b), who suggested that varying grades of skeletal construction (from hymedesmoid, plumose to renieroid reticulate) could be seen within populations of particular species and was apparently largely related to ontogenetic development. Van Soest (1984b), Hooper & Levi (1993a) and others used this ontogenetic argument to merge Microciona and Clathria (as well as Thalysias and Clathria), and Hooper et al. (1992) presented some empirical data to show that there was no homogeneity in biochemical profiles to support the retention of distinct taxa for encrusting versus erect species. 2) Two separate taxa can be recognised for species with hymedesmoid plumose (Microciona) and reticulate (Clathria) architectures. This was the argument accepted by Levi (1960a), Bergquist & Fromont (1988), and others, on the basis that these skeletal structures were consistent for populations of particular species, and therefore represented fixed genetic differences. Simpson (1968a) suggested further that this option had some empirical support from cytological evidence, although (unfortunately) the cytological characters themselves are at the moment not particularly useful taxonomic characters. A more pragmatic argument for the retention of the name Microciona is that it is in current widespread use by sponge biochemists, ecologists and experimental biologists, and retaining this name provides some sort of nomenclatural stability consistent with the previous literature. This argument, unfortunately, has a pragmatic rather than biological basis. 3) The name Clathria could be used for an adult terminal phase of skeletal architecture, related directly to the ontogeny of the sponge, whereas Microciona could be used for the juvenile phase of the same species. Dictyociona is an example of a Clathria with an intermediate Microcionalike skeleton, and Pseudanchinoe is an example of a Microciona verging on a reticulate Clathrialike skeleton. This argument presupposes that plumose skeletons are always precursors of, and juvenile to, reticulate skeletons. But there are several thinly encrusting species which do have reticulate skeletons (e.g., Sara & Melone, 1966), even though most plumose species are also persistently encrusting. In recognising M icrociona at the subgeneric level, Van Soest (1984b) implicitly also recognised a phylogenetic basis for the `microcionid' skeletal specialisation, even though there were no other corroboratory characters, such as unique spicule geometries. The phylogenetic interpretation of this skeletal specialisation taken in the present study is that Microciona is a persistently encrusting sponge which consistently has nonanastomosing basal spongin fibres cored by nonanastomosing plumose spicule tracts or single 62zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM spicules, also lacking any ectosomal specialisation; it is an incompletely differentiated sister taxon of Clathria and recognised here at the subgenus level (Clathria (Microciona)). The Leptoclathria (flat, hymedesmoid) and Microciona (ascending `microcionid' fibre nodes) conditions are not differentiated, these being interpreted as being more likely to be related to the thickness of encrusting growth forms than anything else. Echinoclathria). Wiedenmayer (1989) provided further explanation of these relationships, although his nomenclatural decisions are not entirely correct. Seriatula was erected for Spongia seriata Grant (Gray, 1867) which is conspecific with 0. papilla Bowerbank (e.g., Simpson, 1968a), and therefore Seriatula becomes an objective synonym of Ophlitaspongia. Ophlitaspongia Bowerbank, 1866 (Fig. 20A-B) Pandaros Duchassaing & Michelotti, 1864 (Fig. 20C-E) Ophlitaspongia Bowerbank, 1866: 14, 378; Wieden- Pandaros Duchassaing & Michelotti, 1864: 88; mayer, 1989: 59; Bergquist & Fromont, 1988: 113. Not Ophlitaspongia; Dendy, 1896: 36; Hallmann, 1912: 253; Levi, 1960a: 58; Wiedenmayer, 1977: 140. Seriatula Gray, 1867: 515; de Laubenfels, 1936a: 122. TYPE SPECIES. Ophlitaspongia papilla Bowerbank, 1866: 378 (by original designation) (holotype BMNH1910.1.1.395); = Spongia seriata Grant, 1826: 116 (Simpson, 1968a:37) (holotype BMNH1847. 9.7.14). Encrusting bulbous growth form. Surface microconulose, microscopically hispid. Choanosomal skeleton isodictyal reticulate, with compressed layer of spongin fibre lying on substrate and regularly reticulate spongin fibres arising from base, producing regular isodictyal fibre network divided into primary ascending and secondary transverse fibre elements. Primary fibres cored by plumose tracts of choanosomal principal styles, which also protrude from fibres at acute angles resembling quasi-echinating spicules. Secondary transverse fibres uncored or with unispicular tracts of usually smaller choanosomal principal styles. Subectosomal skeleton paratangential, with scattered subectosomal auxiliary styles lying on or near surface and also dispersed throughout mesohyl. Ectosome without specialised spiculation, but plumose tracts of choanosomal principal megascleres protrude through surface. Megascleres large and small, entirely smooth choanosomal principal styles-subtylostyles, and thin smooth subectosomal auxiliary styles. Microscleres u-shaped toxas. Chelae absent. REMARKS. This definition is based on type material and Simpson's (1968a) description of live populations. Ophlitaspongia papilla, was shown by Simpson (1968a: 95) to be a synonym of the type species of Microciona (M. atrasanguinea). A broader concept of Ophlitaspongia (Wiedenmayer, 1989), (not 0. seriata (Grant)), is identical to Echinoclathria (s.s.) (see remarks for Schmidt, 1870: 59; de Laubenfels, 1936a: 123; Wiedenmayer, 1977: 143; Van Soest, 1984b: 127. TYPE SPECIES. Pandaros acanthifolium Duchassaing & Michelotti, 1864: 90 (by subsequent designation of de Laubenfels, 1936a: 123)) (lectotype TMPOR57). Bushy arborescent growth form. Surface highly conulose, with flattened or lobate lamellae. Choanosomal skeleton reticulate, with welldeveloped flattened spongin fibres (trabeculae) cored by choanosomal principal subtylostyles lying in all directions within fibres (from isodictyal reticulate to echinating) and with sparse acanthostyles echinating or also incorporated into fibres. Subectosomal skeleton radial, reduced to single long subectosomal auxiliary subtylostyles protruding through surface and also scattered throughout mesohyl. Ectosome without special spicules. Megascleres smooth choanosomal principal subtylostyles-tylostyles, often with slightly rhabdose bases and terminal or subterminal basal swellings, long curved or straight subectosomal auxiliary subtylostyles, and lightly acanthose or rarely smooth styles `echinating' fibres. Microscleres absent. REMARKS. This definition is based on the type and a fragment of the type MNHNLBIMDNBE 1309, specimen BMNH1884.7.11.2 and description of live populations by Van Soest (1984b). Important features are: 1) the prominently flattened fibres cored by smooth slightly rhabdose principal subtylostyles-tylostyles (more reminiscent of Rhabderemiidae than of Microcionidae); 2) the sparse, lightly spined styles which more closely resemble a second category of principal spicules than than they do echinating (accessory) spicules typical of other Microcionidae; furthermore, these spicules are only rarely seen echinating fibres, but more commonly they are incorporated into them together with the principal megascleres; and 3) the long subectosomal auxiliary subtylostyles protruding through the ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 63 surface (more similar to a reduced Raspailiidae, such as Echinodictyum or Ceratopsion, than to typical Microcionidae). Pandaros could be legitimately included in either Raspailiidae or Microcionidae. Van Soest (1984b) noted that only the rare echinating acanthostyles in P acanthifolium gives any cause to link it to the Microcionidae at all. He speculated that it might be necessary to erect a separate family for the species, or even remove it from the Poecilosclerida altogether, as it also shows affinities with axinellids such as Ptilocaulis. There are no microscleres to give any further clues as to its affinities. Pandaros is maintained as a separate genus and tentatively included in Microcionidae. Of 12 species referred to Pandaros, only the type clearly belongs here. Raspailia kasumiensis Tanita (MMBS SIS-052) was assigned to Pandaros (Hooper, 1990a). Wiedenmayer (1977) merged Thalyseurypon with Pandaros, because he considered that its type species had architecture closely comparable to P acanthifolium (Hechtel, 1965), but this is not upheld here. The only features these genera have in common is lacking microscleres. Wiedenmayer (1977) also speculated that the genus had a close relationship with Echinoclathria (= Holopsamma as defined here), based on alleged similarities in skeletal architecture, and he suggested that the two genera probably intergrade in habit and spiculation, but these suggested affinities are not evident in relevant specimens. Paradoryx Hallmann, 1920 (Fig. 20E-G) Paradoryx Hamann, 1920: 767; de Laubenfels, 1936a: 109. TYPE SPECIES. Clathria dura Whitelegge, 1901: 83 (by original designation) (holotype AMG3046). Arborescent, flabellate growth form with compressed branches. Surface even, smooth. Choanosomal skeleton reticulate, with heavy spongin fibres forming irregular anastomoses in axis becoming more regular and rectangular towards periphery. Spongin fibres differentiated into primary ascending and secondary connecting components, cored by auxiliary styles and heavily echinated by acanthostyles. Subectosomal skeleton plumose, with bundles of auxiliary styles forming plumose brushes on peripheral fibres. Ectosome fibrous, without specialised skeleton, and peripheral fibres form more-or-less reticulate structure through which choanosomal spicules protrude. Megascleres entirely smooth, hastate or quasidiactinal auxiliary styles or subtylostyles, and echinating acanthostyles with large and even spination. Microscleres palmate isochelae resembling arcuate forms with lateral alae fused to shaft for about 1/2 alae length, and shaft greatly curved, thickened. Toxas absent. REMARKS. Ectosomal fibres of Paradoryx are unspecialised, identical to choanosomal fibres in size and form, unlike the Callyspongia-like ectosome of Marleyia. Hallmann (1920) erected Paradoryx for Wilsonella species (with auxiliary megascleres coring fibres) which had arcuate chelae instead of palmate isochelae. Loss or replacement of principal megascleres varies considerably between otherwise allied microcionid taxa and this feature is considered to be of lesser significance at the generic level than assumed by Hallmann (Hooper, 1990a; Hooper et al., 1990). Similarly, isochelae described by Hallmann (1920) as arcuate in the type species are modified (curved, thickened) palmate forms. Apart from the type species, Hallmann (1920) included: Dictyocylindrus pinifonnis Carter and W oxyphila (both of which belong to Clathria (Clathria)), and Clathria elegantula Ridley & Dendy and Wilsonella curvichela Hallmann (which are referred here to Clathria (Dendrocia)). Paratenaciella Vacelet & Vasseur, 1971 (Fig. 20H-0 Paratenaciella Vacelet & Vasseur, 1971: 103. TYPE SPECIES. Paratenaciella microxea Vacelet & Vasseur, 1971: 103 (by original designation) (holotype MNHNLBIMD.W27). Encrusting growth form. Surface uneven, hispid. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer on substrate and with ascending fibre nodes enclosing bases of choanosomal principal subtylostyles standing perpendicular to substrate. In thicker sections up to 3-4 spicules form ascending plumose tracts, protruding through surface. Echinating megascleres absent. Subectosomal skeleton plumose or paratangential composed of bundles of subectosomal auxiliary subtylostyles often forming brushes around protruding principal spicules and protruding through ectosome. Megascleres entirely smooth choanosomal principal subtylostyles, and subectosomal auxiliary subtylostyles with microspined bases and polytylote shafts. Microscleres palmate ^ MEMOIRS OF THE QUEENSLAND MUSEUM 64zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 20. Type species of microcionid genera. A-B, Ophlitaspongia (0. papilla Bowerbank, BMNH1910.1.1.395). C-E, Pandaros (P. acanthifolium Duchassaing & Michelotti, BMNH1884.7.17.2). F-G, Paradoryx (Clathria dura Whitelegge, AMG3046). H-I, Paratenaciella (P. microxea Vacelet & Vasseur, MNHNDJV27). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 65 isochelae, including contort forms, and oxeote toxas. REMARKS. Paratenaciella has a Microcionalike choanosomal fibre skeleton composed of non-anastomosing, plumose fibre nodes, but lacks acanthostyles (similar to Axociella, Axocielita, Tenaciella), has polytylote modifications to auxiliary spicules (similar to Fisherispongia ferrea de Laubenfels and Microciona aceratoobtusa Carter), and has modified (oxeote) toxas, like those in Artemisina archegona Ristau (although type material shows that they do resemble true microxeas). This latter feature appears to be the only unique character for the type species, and as far as is known it is also unique amongst the Microcionidae. Paratenaciella shows some similarities to Esperiopsis canaliculata Whitelegge (which is referred here to Axociella) in its spiculation, although P. microxea has a microcionid architecture, microxeote microscleres and undifferentiated ectosomal-subectosomal megascleres. The genus is monotypic and is synonymised with Clathria (Microciona). Plectispa Lendenfeld, 1888 (Fig. 21C-D) Plectispa Lendenfeld, 1888: 225; Topsent, 1894a: 19; de Laubenfels, 1954: 164. TYPE SPECIES. Plectispa macropora, in part, Lendenfeld, 1888: 226 (by subsequent designation of Hallmann, 1912: 204) (holotype AMG9159). Lobodigitate growth form. Surface 'honeycomb' reticulate. Choanosomal skeleton reticulate, with well developed spongin fibres forming more-orless regular oval or eliptical meshes, sparsely cored and abundantly echinated by principal subtylostyles, and fewer auxiliary styles-tornotes also in fibres. Echinating megascleres less predominant in periphery than in axis. Some detritus also incorporated into choanosomal fibres. Subectosomal skeleton plumose, with brushes of principal subtylostyles from ascending fibres of peripheral skeleton protruding through surface, surrounded by paratangential loose bundles or individual auxiliary spicules. Megascleres smooth principal subtylostyles, and long curved, setaceous or sinuous auxiliary styles or quasi-diactinal tornotes, both with blackened axial canals. Microscleres palmate isochelae, including contort forms. Toxas absent. REMARKS. This genus is poorly defined due to the dubious status of all of Lendenfeld's syntypes, inaccuracies in Lendenfeld's (1888) original description, and misinterpretation of the species (e.g., Clathria macropora; Whitelegge (1901: 91) is different from C. macropora Lendenfeld (1888)). Hallmann (1912: 165) attempted to sort out these problems but was only partially successful. He rejected Whitelegge's proposed merger of P. macropora with Echinonema levis Lendenfeld (1888), but instead he synonymised the latter with C. macropora (sensu Whitelegge), as Crella incrustans Carter, var. levis Lendenfeld. However, Hallmann's (1912) remarks and assumptions concerning the genus are not entirely correct (see remarks below for Clathria macropora and Holopsamma macropora), and there is no evidence that he actually re-examined relevant type material. His nomenclatural decisions are erroneous and not supported here. There are 4 type specimens and one other specimen in the AM and BMNH which bear the name macropora'. 1) AMZ959 (with the label 'dry, cotype') from Port Jackson, NSW, is the so-called 'type' of Clathria macropora (sensu Whitelegge), and is a species of Crella. It is probably a syntype of Lendenfeld's (1888) Echinonema levis, but that assumption is difficult to corroborate due to the loss of Lendenfeld's 'key-list' from the Australian Museum (E. Pope & E Rowe, pers.comm.). 2) AMZ466 (with label stating 'spirit preserved, ?type') from Port Stephens, NSW is the lectotype of Lendenfeld's (1888: 221) Clathria macropora, later referred to Wilsonella by Hallmann (1912: 240). It is neither a Wilsonella, nor closely related to such species as W australiensis Carter, but is a Clathria lacking echinating megascleres and having an axially compressed skeleton (e.g., Clathria (Axociella)). 3) AMG9159 (with the label stating 'dry, type'), probably from Port Jackson, NSW (i.e., there is a discrepancy between the published locality of Torres Strait, Qld. and the museum specimen label and register), is the holotype of Plectispa macropora. 4) BMNH1925.11.1.555 (with label stating Plectispa macropora, dry, Dendy coll.') from Manly Beach, NSW, is identical to Clathria multipes Hallmann. 5) Another specimen (BMNH1957.8.30.2, with label stating Plectispa macropora, dry') belongs to Clathria lendenfeldi. Consequently, only a single syntype is valid becoming the holotype of P. macropora, and the diagnosis of Plectispa is based on this specimen. Hallmann (1912:205) referred the type species (and genus Plectispa) to Wilsonella, but then returned it to Clathria (Hallmann, 1920: 768). The species has a 'honeycombed reticulate' MEMOIRS OF THE QUEENSLAND MUSEUM 66zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ growth form and smooth echinating spicules (not acanthose as supposed by Lendenfeld (1888) and Hamann (1912)). It also has long, setaceous, quasi-diactinal auxiliary megascleres, which also core fibres, scattered throughout the mesohyl and associated with spicule brushes on the surface. These characters indicate that Plectispa belongs to Holopsamtna, and the species shows some resemblance to H. ramosa (Hallmann, 1912) and H. laminaefavosa Carter, 1885b (sensu Wiedenmayer, 1989). Plocamia Schmidt, 1870 (Fig. 21A-B) Plocamia Schmidt, 1870: 62. Dirrhopalum Ridley in Ridley & Duncan, 1881: 477. spicules are much better developed in Plocamilla. Plocamia is the earliest available name for a group of Antho-like ('plocamiform') microcionids having basal tylotes and echinating spicules (see remarks for Holoplocamia, Lissoplocamia). Plocamilla Topsent, 1928 (Fig. 21E-F) Topsent, 1928a: 63; Levi, 1960a: 80; Levi, 1960b: 760; Pulitzer-Finali, 1973: 40; Simpson, 1968a: 95; Van Soest, 1984b: 26, 29, 125; (not Burton, 1935a: 402). Plocam illa TYPE SPECIES. I sodiciy a coriacea Bowerbank, 1874: 136 (by original designation of Topsent, 1928a: 63) (holotype BMNH1910.1.1.251). Plocam ia gym nazusa Schmidt, 1870: TYPE SPECIES. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA DIAGNOSIS. Encrusting growth form. Surface 62 (by subsequent designation (Burton, 1935a: 401)) uneven, porous, microscopically hispid. (holotype possibly LMJG, schizotype Choanosomal skeleton renieroid, with regular MNHNLBIMDCL1105L). DIAGNOSIS. Encrusting growth form. Surface even, microscopically hispid. Choanosomal skeleton renieroid, with tylotes forming basal reticulation, interconnected at nodes by light spongin fibres, and with plumose columns of choanosomal principal styles and echinating styles embedded in basal renieroid skeleton, individually or in bundles, largest protruding through surface. Subectosomal skeleton with paratangential and erect tracts of subectosomal auxiliary styles. Megascleres robust slightly curved choanosomal principal styles with microspined or smooth bases, robust tylotes with swollen microspined bases, long subectosomal auxiliary styles with smooth or microspined bases, smaller echinating styles with microspined bases. Microscleres palmate isochelae and wingshaped toxas. basal reticulation of acanthose strongyles in unior paucispicular tracts, overlaid by plumose brushes of larger choanosomal principal styles and echinating acanthostyles projecting perpendicularly from nodes of renieroid spicule skeleton. Ectosomal skeleton contains plumose or paratangential brushes of subectosomal auxiliary styles and projecting echinating acanthostyles from peripheral nodes of choanosomal renieroid skeleton. Megascleres acanthose strongyles or tylotes with spines mostly on basal ends, larger choanosomal principal styles-subtylostyles with spined bases, subectosomal auxiliary styles-subtylostyles with smooth or microspined bases, and small echinating acanthostyles with few spines concentrated mainly on basal end. Microscleres palmate isochelae, smaller wingshaped toxas, and larger accolada toxas with spined points. REMARKS. Ridley (in Ridley & Duncan, 1881) suggested that Plocamia Schmidt was preoccupied by Plocamium Lamouroux, 1828, a seaweed, and consequently proposed the replacement name Dirrhopalum Ridley. However, under Article 56b of the ICZN (Anonymous, 1985) this is an unjustified emendation and Plocamia stands. Bergquist & Fromont (1988) also used Plocamia over Dirrhopalum. This definition is based on slides of the holotype and published descriptions (Schmidt, 1870; Ridley in Ridley & Duncan, 1881). The type species is essentially the same as Plocamilla in structure and spicule geometry, but spines on both the basal tylote spicules and echinating REMARKS. This definition is based on the holotype and descriptions of the type species (Topsent, 1928a; Levi, 1960a). Plocamilla has spiculation similar to Plocamia Schmidt, but it has an encrusting growth form and therefore its skeleton is not clearly differentiated into primary or secondary lines (Topsent, 1928a: 63; Levi, 1960a: 80). This skeletal development is probably related to growth form and not an important generic character. The genus was not formally diagnosed until Burton (1935a: 402) differentiated a number of `plocamiform' genera, all of which had acanthostrongyles or acanthostyles forming the basal renieroid skeleton. Burton emphasised the diagnostic value of choanosomal REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 67 acanthostrongyles ('dumbell spicules') which united those genera, but he differentiated them by their skeletal architecture and the presence or absence of various spicule types. Holoplocamia is a synonym of Plocamilla (Levi, 1960a: 80), although Little (1963: 47) argued to the contrary. Levi (1960a) also noted that the skeleton of P coriacea was very close to Antho and Dictyoclathria (both of which are objective synonyms), lacking special echinating spicules and having acanthostyles instead of acanthostrongyles in the basal renieroid skeleton. Levi (1960a) and Pulitzer-Finali (1973: 40) were cautious in interpreting whether Plocamilla was distinct from Antho, because the spined acanthostyles and smooth principal styles could not be consistently differentiated in all taxa. For example, P. elegans (Ridley & Dendy) does not show any clear separation between the smaller category of smooth principal styles and the more sparsely spined acanthostyles (Pulitzer-Finali, 1973). On that basis, Pulitzer-Finali suggested that future studies may show that P. elegans should be referred to Antho, whereas P coriacea has clearly differentiated coring and echinating megascleres, and should remain in Plocatnilla. Levi (1960a), Simpson (1968a), Pulitzer-Finali (1973) and Van Soest (1984b) maintained Plocamilla and Antho as separate genera, the former having echinating acanthostyles and predominantly (acantho)strongyles in the renieroid skeleton. However, all these authors admitted that the two genera may be too similar to maintain sufficient generic separation. Van Soest & Stone (1986) noted that the genus should probably be merged with Antho, together with other plocamiform genera containing renieroid skeleton of acanthose megascleres, and this suggestion is supported here (i.e., Antho (Plocamia)). Simpson (1968a: 95) found that although Plocamilla and Microciona were easily differentiated by their skeletal construction and spicule geometries, they were remarkably similar in their cytological characteristics. These findings contradict the more obvious similarities between microcionid genera, based on spicule geometry and skeleton construction. Microciona, Clathria and Thalysias are a relatively homogenous group in skeletal construction, compared to any inferred relationship between Plocamilla and Microciona for example. Probably Simpson's (1968a) cytological characters have not been incorporated into the classification, nor have the the implications of his findings to phylogeny of demosponges been widely discussed. Plocamiopsis Topsent, 1904 (Figs 21G-H, 22A)zyxwvutsrqponmlkjihgfe Plocam iopsis Topsent, 1904a: 155; Burton, 1935a: 402. TYPE SPECIES. Plocam iopsis signora Topsent, 1904a: 155 (by monotypy) (holotype MNHNLBIMDT947). DIAGNOSIS. Encrusting growth form. Surface even, hispid. Choanosomal skeleton irregularly renieroid, with basal reticulation of acanthostrongyles in uni- or bispicular tracts lying on substrate. Choanosomal principal subtylostyles, smaller acanthostyles and spicules intermediate to both embedded individually in basal skeleton, standing perpendicular to substrate. Subectosomal skeleton plumose, with ascending brushes of subectosomal auxiliary subtylostyles surrounding principal megascleres. Ectosome without specialised spiculation, but with both tangential and plumose tracts of subectosomal auxiliary spicules protruding through surface. Megascleres choanosomal principal subtylostyles with spines on bases and proximal region of shaft, smaller acanthostyles with aspinose points, and subectosomal auxiliary subtylostyles with spined bases. Microscleres cleistochelae and smooth wing-shaped toxas. REMARKS. Plocamiopsis is a member of de Laubenfels' (1936a) plocamiform group of sponges having a basal renieroid skeleton of acanthostrongyles. It differs from others in this group (Dirrhopalum, Heteroclathria, Holoplocamia, Lissoplocamia, P locamilla) having cleistochelae, which are also seen in Colloclathria and Quizciona. Cleistochelae are hypersilicified palmate isochelae with the front alae fused (Fig. 22A) but probably have little phylogenetic significance given the complete transitional series from palmate to cleistochelate (Fig. 76G). Plocamiopsis is referred to Antho (Plocamia). Protophlitaspongia Burton, 1934 (Fig. 22B-C) Prot ophlit aspongia Burton, 1934a: 562. TYPE SPECIES. Siphonochalina bispiculat a Dendy, 1895: 246 (by original designation) (lectotype NMVG2319). Lamellar, tubulo-digitate growth form, with osculum on apex of each tube. Surface uneven, microscopically hispid. Choanosomal skeleton subisodictyal reticulate, with more-or-less regularly anastomosing heavy spongin fibres ^ MEMOIRS OF THE QUEENSLAND MUSEUM 68zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 21. Type species of microcionid genera. A-B, Plocamia (P. gymnazusa Schmidt, MNHNDCL1105L). C-D, Plectispa (P. macropora sensu Lendenfeld, AMG9159). E-F, Plocamilla (Isodictya coriacea Bowerbank, BMNH1910.1.1.251). G-H, Plocamiopsis (P. signata Topsent, MNHNDT947). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 69 forming primary ascending and secondary transverse lines. Primary fibres cored by multispicular tracts of longer auxiliary oxeas; secondary fibres usually aspiculose occasionally with only single coring auxiliary oxeas. Fibres sparsely echinated by smaller, smooth hastate oxeas (presumed to be homologous with principal spicules). Ectosome lacks specialised spiculation, but has plumose erect brushes of oxeas from peripheral skeleton protruding through surface. Megascleres modified diactinal, including longer smooth hastate auxiliary oxeas, and smaller smooth hastate principal oxeas. Microscleres absent. REMARKS. The genus contains 7 previously named species: P bispiculata (Dendy, 1895: 246) from Port Phillip Bay, Victoria, P. oxeata Burton (1934a: 562) from the Papuan Pass, northern Australia, P. ada de Laubenfels (1954: 96) from Ponape, P aga de Laubenfels (1954: 97) from the central west Pacific, P antillana Pulitzer-Finali (1986: 138) from the West Indies, Echinochalina (P.) laboutei Hooper & Levi (1993a: 1277) and E. (P.) bargibanti Hooper & Levi (1993a: 1280) both from New Caledonia. These species resemble Haplosclerida to some extent; P. anti!lanais a haplosclerid, (Niphatidae). Similarly, de Laubenfels' (1954), P ada and P aga, are placed in the Phoriospongiidae leaving 4 known species and 4 new Australasian species described below. The genus has a characteristic three-dimensional ectosomal architecture reminiscent of Hemigellius and Anzphimedon (Haplosclerida: Niphatidae), with only oxeas as megascleres. However, smooth hastate oxeas echinating fibres, with a similar geometry but smaller size than coring spicules, gives some clues as to probable phylogenetic relationships (i.e., Microcionidae, some lophonidae, and Raspailiidae). This pattern of spiculation is seen in Echinochalina, and to a lesser extent Echinoclathria as defined here. It is particularly well developed in E. axinelloides, and for this reason Burton (1934a: 562) tentatively referred Protophlitaspongia to the Microcionidae. Conversely, de Laubenfels (1936a: 54, 1954: 96) suggested that the genus was closer to Guitarra (although lacking their peculiar microscleres), or Liosina, and he referred Protophlitaspongia to the Desmacidonidae, but he was probably referring mainly to his two Micronesian species in doing this. Pulitzer-Finali (1986: 138) followed similar reasoning, placing the genus in Esperiopsidae, but none of these species belong to Protophlitaspongia in any case. Further clues as to the most appropriate placement of this genus can be seen from other characteristics: the palmate isochelae in E. (P.) bargibanti, the accolada toxas in E. (P.) oxeata, and the plumose ectosomal skeleton, composed of auxiliary spicule brushes (typical of Microcionidae, Axinellidae and Raspailiidae) in all species. The present interpretation of Protophlitaspongia suggests affinities with Echinoclathria and Holopsanuna microcionids and the reticulate Raspailiidae, and in the distribution of its megascleres (i.e., auxiliary spicules in fibres and peripheral skeleton, principal spicules echinating fibres) it is assigned to Echinochalina. Pseudanchinoe Burton, 1929 (Fig. 22D-E) Pseudanchinoe Burton, 1929a: 433; de Laubenfels, 1936a: 109. TYPE SPECIES. Stylostichon toxiferum Topsent, 1913a: 621 (by original designation and monotypy) (holotype MNHNLBIMDT1612). Massive subspherical growth form. Surface uneven, apical surface conules. Choanosomal skeleton plumoreticulate, with spongin fibres forming primary plumose ascending columns, partially interconnected by transverse secondary fibres, both cored by multispicular tracts of choanosomal principal subtylostyles and heavily echinated by acanthostyles. Subectosomal skeleton plumose, non-anastomosing, composed of multispicular plumose tracts of choanosomal principal spicules from peripheral fibres surrounded by plumose bundles of subectosomal auxiliary subtylostyles. Ectosomal skeleton with second tier of subectosomal auxiliary subtylostyles forming dense palisade on surface. Megascleres choanosomal principal subtylostyles with or without basal spines, subectosomal auxiliary subtylostyles with microspined bases, and echinating acanthostyles varying from smaller forms with even spination to larger forms with aspinose points. Microscleres oxhorn and accolada toxas. Chelae absent. REMARKS. Burton (1929a) recorded palmate isochelae in the type species but none were found in the holotype nor did Topsent (1913a) record any in his original description. In fact the absence of isochelae and the possession of plumoreticulate skeletal architecture are virtually the only two diagnostic features that distinguish Pseudanchinoe from other Clathria (s.s.). Similarly, Topsent (1913a) stated that there were MEMOIRS OF THE QUEENSLAND MUSEUM ^ 70zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA microspines on the points of toxas, but these were not seen in the holotype. The type species of Pseudanchinoe is intermediate between Clathria and Thalysias, having the ectosomal structure of the latter (i.e., with a thick continuous palisade of erect spicules overlaying a similar subectosomal skeleton), but with spicule composition of the former (single category of auxiliary styles), further supporting the decision to merge these two genera in a single taxon (Clathria s.1.). In skeletal structure and diversity of megascleres the genus could be placed in the series: Dendrocia (single category of structural spicule in the entire skeleton), Clathria (one category of auxiliary spicule and one category of principal spicule), Pseudanchinoe (one category of auxiliary spicule forming a continuous ectosomal palisade), Thalysias (two categories of auxiliary spicules the smaller forming a continuous ectosomal palisade, and one category of principal spicule), and Antho (some of which have two categories of auxiliary spicules and two categories of principal spicules). Burton (1929a) remarked on the apparent similarities between Pseudanchinoe and Anchinoe Gray (= Phorbas Duchassaing & Michelotti; Anchinoidae). The latter genus has plumose columns of intermingled acanthostyles and tornotes (or oxeas), which are echinated by acanthostyles, but these similarities are superficial. Anchinoidae have tangential ectosomal diactinal megascleres and plumose columns of choanosomal megascleres in which acanthostyles predominate (e.g., Van Soest, 1984b: 86). Of the numerous species assigned to Pseudanchinoe (e.g., de Laubenfels, 1936a: 109) most are interpreted here as belonging to Clathria, having secondarily lost their isochelae. Australian examples are C. (C.) caelata Hallmann, C. (C.) costifera Hallmann, C. (C.) inanchorata Ridley & Dendy, and C. (C.) partita Hallmann. Two other species belong to Clathria (Thalysias) (C. dentata Topsent and C. fascicularis Topsent, which are synonymous) (Van Soest, 19846). Of contemporary authors only Koltun (1976) recognises this genus whereas Van Soest (1984b) and (Wiedenmayer, 1989) merged it with Clathria. It is included here in Clathria (Clathria). Qasimella Thomas, 1974 Qasimella Thomas, 1974: 311. TYPE SPECIES. Qasimella indica Thomas, 1974: 311 (by original designation)(holotype CMFRIT84/1 not seen). Specialised tubular growth form, with apical oscules and central cavity running longitudinally through body. Surface smooth, even. Choanosomal fibres apparently absent, and skeletal structure poorly defined with more-orless longitudinal tracts of choanosomal principal subtylostyles bound together at nodes by collagen. Echinating acanthostyles absent. Subectosomal skeleton rudimentary, composed of subectosomal auxiliary subtylostyles lying just below surface, not protruding through it. Ectosomal skeleton absent. Megascleres choanosomal principal subtylostyles with spined bases, and slender smooth subectosomal auxiliary subtylostyles. Microscleres described as 'arcuate' isochelae and wing-shaped toxas. REMARKS. The holotype and paratypes (CMFRI T8412) of Q. indica are housed at the Central Marine Fisheries Research Institute, Mandapan Camp, Cochin, India. A slide of the holotype is housed at the IM (A.K. Mandal, pers. comm.). The genus is monotypic and known only from its original description (Thomas, 1974). It is an unusual, very thin-walled hollow tubular sponge with pseudo-syconoid construction, and attached to the substrate by short peduncles reminiscent of Aulospongus Norman (Raspailiidae). Apart from spicule diversity and geometry, which place the type species in Microcionidae, we know of few other characters that give clues as to its affinities with other microcionids. From its skeletal structure, or lack of it, Qasimella shows similarities to Artemisina. It is also possible that Q. indica is a larval sponge, but until type material is seen its published description suggests that it has affinities to Artemisina. The chelae have been described as 'arcuate', but this cannot be confirmed and must be regarded as suspect (e.g., they may be merely modified, curved, palmate ones). Quizciona de Laubenfels, 1936 (Fig. 22F-G) Quizciona de Laubenfels, 1936a: 111. TYPE SPECIES. Microciona heterospiculata Brondsted, 1924: 465 (by original designation) (fragment of type BMNH1901.12.26.13). Encrusting growth form. Surface hispid, uneven. Choanosomal skeleton differentiated into two components: renieroid basal layer composed of spongin fibres lying on substrate cored by bispicular tracts of acanthostyles forming triangular meshes; basal fibres ascending non-anas- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 71 tomosing fibre nodes echinated by multispicular plumose tracts of both choanosomal principal subtylostyles and echinating acanthostyles (of same morphology as basal renieroid skeleton spicules) embedded and erect on substrate. Ectosomal skeleton paratangential, composed of two size classes of subectosomal auxiliary subtylostyles, forming occasional brushes and scattered in mesohyl matrix near periphery. Megascleres longer choanosomal principal subtylostyles with heavily microspined bases, shorter acanthostyles with aspinose points (forming both renieroid basal skeleton and echinating fibre nodes), and two size classes of auxiliary subtylostyles with smooth or spined bases. Microscleres palmate isochelae, cleistochelae and small accolada toxas. REMARKS. The original description differs slightly from the definition above. In particular, the very small, thin toxas were overlooked, as was a renieroid basal skeleton typical of de Laubenfels' (1936a) plocamiform sponges. Brondsted's (1924) description of Quizciona suggests that it is close to Microciona with a hymedesmoid basal skeleton and plumose ascending fibre nodes but having cleistochelae in addition to palmate isochelae. The type material shows the species more closely related to Antho with a renieroid basal skeleton. Smaller acanthostyles and larger principal styles echinating basal fibres is similar to Plocamilla, whereas acanthostyles rather than acanthostrongyles coring the basal fibres is similar to Antho, and this mix of characters illustrates the difficulty in separating these two taxa. Cleistochelae have been recorded in several microcionids (Microciona cleistochela Topsent, M. clathrata Whitelegge, M. chelifera Levi, Clathria simpsoni Van Soest, C. toxipraedita Topsent and Colloclathria ramosa Dendy). Alander (1942) and Van Soest (1984b) suggested that cleistochelae are modified palmate isochelae (with fused chelate teeth) which have arisen several times independently within the Microcionidae, and consequently they do not have primary taxonomic significance. Interestingly, Brondsted (1924) also remarked on the similarities between cleistochelae of Microcionidae and clavidiscs of the hypercalcified sponge Merlia Kirkpatrick, supporting current theories on the origin and affinities of the `sclerosponges' with the demosponges (e.g., Vacelet, 1985). Bergquist & Fromont (1988) merged M. heterospiculata with Clathria mortensenii and suggested that Quizciona should be abandoned on the basis that they did not find cleistochelae in their material, contrary to de Laubenfels' (1936a) description. This synonymy is not upheld here, although Bergquist's (1961a: 39) record of M. heterospiculata from northern New Zealand may be Clathria mortensenii. I interpret Quizciona, based strictly on the type specimen, as a synonym of Antho (Antho). Ramoses de Laubenfels, 1936 (Fig. 22H-I) Ramoses de Laubenfels, 1936a: 109. TYPE SPECIES. Clathria pauper Brondsted, 1927: 3 (by original designation) (fragment of type BMNH1930.11.5.2). Arborescent, tubulo-digitate growth form. Surface shaggy, uneven, microscopically hispid. Choanosomal skeleton irregularly plumo-reticulate, with vaguely ascending multispicular primary tracts interconnected by transverse paucispicular secondary tracts cored by both smooth and partially spined choanosomal principal styles and echinated by plumose bundles of smaller acanthostyles. Spongin fibres not seen but spicules united by moderate quantities of collagen. Subectosomal skeleton plumose, with bundles of subectosomal auxiliary subtylostyles protruding through surface. No special ectosomal skeleton. Megascleres longer entirely smooth choanosomal principal styles, slightly shorter principal subtylostyles with spines on bases and distal end of shaft, small slender echinating acanthostyles with even spination, and subectosomal auxiliary subtylostyles with spined bases. Microscleres accolada toxas, some with contort centres. Chelae absent. REMARKS. The holotype of C. pauper has not yet been discovered in the Brondsted collection at UZM (0. Tendal, pers. comm.), but a fragment of it is held at the BMNH. A fragment of C. pauper (with second label stating Trotoclathria antarctica, 29.1.19?? [illegible], 340 fathoms, M Burton') is in Sydney (AMZ2239). It is possible that this material is also a fragment of the holotype, but its status and origin are not certain. Brondsted's (1927) original description conforms closely to the type material, although he appears to have overlooked the thin accolada toxa microscleres (some with asymmetrical contort central curvature). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 72zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 22. Type species of microcionid genera. A, Plocamiopsis (P. signata Topsent, MNHNDT947). B-C, Protophlitaspongia (Siphonochalina bispiculata Dendy, NMVG2319). D-E, Pseudanchinoe (Stylostichon toxiferum Topsent, MNHNDT1612). F-G, Quizciona (Microciona heterospiculata Brondsted, BMNH1901.12.26.13). H-I, Ramoses (Clathria pauper Brondsted, BMNH1930.11.5.2). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 73 Ramoses is similar to Pseudanchinoe in lacking chelae microscleres, and Koltun (1976: 155) synonymised the type species of Ramoses with P. toxiferuin Topsent, the type species of Pseudanchinoe (both species are from the Ross Sea, Antarctica). This synonymy is confirmed here from type specimens and both species are lipochelous Clathria (Clathria). Koltun (1976) suggested that C. pauper was a synonym of Microciona toxifera (Topsent), also from the Antarctic, but this synonymy is rejected. Comparison of relevant type specimens shows that the two species differ substantially in their skeletal structure, spicule size and spination on megascleres. REMARKS. The type species is from the Indowest Pacific region where most species occur. The primary and only consistent morphological feature that distinguishes Rhaphidophlus from other Clathria-like taxa is the specialised ectosomal skeleton, consisting of two differentiated categories of auxiliary subtylostyles which form brushes on the surface (either as discrete brushes (s.s.)or in a continuous palisade). This is identical to the condition described for Thalysias (which has seniority). Hallmann (1912), followed most recently by Van Soest (1984b), suggested that the two sizes of auxiliary megascleres may represent age differences in spicules, in which case the genus has a distinct localisation of adult and juvenile megascleres. This assertion, however, does not Rhaphidophlus Ehlers, 187 yet have any empirical (experimental) support. (Fig. 23A-C)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA This difference in size/age of auxiliary spicules is crucial to the definition of Rhaphidophlus, beRhapidophlus Ehlers, 1870: 19,31; Ridley, 1884a: 449cause some species of Clathria, Microciona and 453; Ridley & Dendy, 1887: 151; Topsent, 1894a: Dendrocia have a dermal skeleton of similar 19, 1925: 658, 1928a: 61, 1932: 98; Thiele, 1903a: structure but containing only one sort of auxiliary 957; Dendy, 1905:170; Hallmann, 1912: 175,1920: spicule (C. impelfecta, C. striata, C. pyramida). 769; Burton, 1932a: 320; de Laubenfels, 1936a: 112; Levi, 1960a: 56; Bergquist, 1965: 168; Similarly, some species of Antho (e.g., A. ridleyi) Simpson, 1968a: 101, 104-106; Wiedenmayer, also have relatively dense ectosomal brushes, but 1977: 140; Van Soest, 1984b: 109; Bergquist & these consist of a single undifferentiated category Fromont, 1988: 118. of subectosomal megasclere. Hallmann (1912) debated the value of ectosomal specialisation as TYPE SPECIES. Spongia crat it ia Esper, 1797: 195 (by a generic character, concluding that although the original designation) (fragment of type ZMB4577). distinction between the two genera may eventualArborescent growth form. Surface microscopi- ly breakdown, they can be consistently differencally hispid, conulose. Choanosomal skeleton ir- tiated on composition of peripheral skeleton regularly reticulate, with heavy spongin fibres rather than its development or density. These cored by multispicular tracts of choanosomal conclusions are supported here. Another character predominant in Thalysias principal subtylostyles which terminate in plumose tracts on peripheral fibres. Plumose mul- (including Rhaphidophlus) is the presence of tispicular tracts of echinating acanthostyles more than one size category of isochelae, of clumped around fibre nodes and also protruding which one or more may be contort (Fig. 5F). This from fibres at regular intervals. Subectosomal feature is not consistent among species, nor is it skeleton plumose, with well-developed multi- exclusive to the genus. Contort chelae are comspicular columns of subectosomal auxiliary sub- mon in Clathria (Thalysias) (e.g., Spongia tylostyles arising from ends of principal spicule abietina Lamarck, Spongia cactiforinis Lamarck, brushes in peripheral skeleton. Ectosome with Rhaphidophlus cervicornis Thiele, R. spiculosus bundles of erect ectosomal auxiliary subtylos- Dendy, R. topsenti Thiele, Clathria fasciculatus tyles overlaying subectosomal plumose brushes Wilson and C. spiculosus var. macilenta of spicules. Megascleres entirely smooth hastate Hentschel), Clathria (Clathria) and Clathria choanosomal principal subtylostyles, larger (Microciona) (e.g., Dictyociona adioristica de smooth subectosomal auxiliary subtylostyles, Laubenfels, Clathria mixta Hentschel, C. bulshorter thinner smooth ectosomal auxiliary sub- botoxa Van Soest, Fisherispongia ferrea de tylostyles, and echinating acanthostyles with Laubenfels, Esperiopsis obliqua George & Wilaspinose necks. Microscleres palmate isochelae son, Wilsonella conectens Hallmann, and M. of two size categories, and two forms of toxas prolifera (sensu Wilson)), Clathria (Wilsonella) (small wing-shaped and larger asymmetrical (e.g., Microciona tuberosa Bowerbank, Clathriopsanuna ^reticulata Lendenfeld), sinuous toxas). MEMOIRS OF THE QUEENSLAND MUSEUM ^ 74zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Axociella) (e.g., Esperiopsis canaliculata Whitelegge), Antho (Antho) (e.g., Holoplocamia penneyi de Laubenfels) and Holopsamma (e.g., Plectispa macropora Lendenfeld). It is also known to occur in Cornulum Carter (Iophonidae) (e.g., C. johnsoni (de Laubenfels)) and consequently it cannot be given much diagnostic value above the species level. There are several other characters used by authors to define genera present in many species of Thalysias (including Rhaphidophlus), but are not apomorphies occurring throughout related groups in all combinations (e.g., encrusting growth form and hymedesmoid architecture (e.g., Leptoclathria lambda Levi); absence of echinating megascleres (e.g., Axociella arteria de Laubenfels), absence of microscleres (e.g., Clathria fascicularis Topsent), and modified isochelae (e.g., Colloclathria ramosa Dendy)). A systematics based on these features conflicts with one based on ectosomal specialisation and dispersal of structural megascleres within the skeleton. By comparison, Dendy (1905), Wilson (1925), de Laubenfels (1936a), Hartman (1955), Wells et al. (1960), Bergquist (1965), Hooper (1990a), Hooper et al. (1990, 1992) and Hooper & Levi (1993a) amalgamated Thalysias (including Rhaphidophlus) and Clathria (including Microciona) on the basis that dermal specialisation can vary intraspecifically, especially in relation to growth form and age of an individual. These authors suggest that this feature is probably not a sound base on which to separate genera. In contrast, Simpson (1968a), Wiedenmayer (1977) and Bergquist & Fromont (1988) maintain these genera separately, although they do state that dermal specialisation may not be important at the generic level (i.e., they offer a convenient classification rather than one based on phylogeny). Simpson (1968a) showed that despite very close morphological similarities between Clathria and Rhaphidophlus, there were cytological differences between the taxa which he considered were sufficient to separate them at the generic level of classification. Thus, Simpson's (1968a) cytological data supports the conclusion above based on morphological evidence that the only morphological character of consistent diagnostic importance, and which correlates to some extent with supposed biological differences between those genera, is the possession of ectosomal specialisation. This conclusion is upheld in this study and used at the subgeneric level. There are several nomenclatural complexities that still exist for Rhaphidophlus, and these require further discussion. 1) Van Soest (1984b: 91) argued that Thalysias should be abandoned in favour of Rhaphidophlus, following Levi (1960a: 52). His argument was based on the premise that Carter (1876: 311) designated the 'representative' of Thalysias as Thalysias subtriangularis (Duchassaing, 1850), and he suggested that although Carter (1876) did not actually use the words 'type' or 'type species', there was no doubt of his intentions (T subtriangularis is a species of Xestospongia; Wiedenmayer, 1977: 255; Van Soest et al., 1983: 199). Van Soest (1984b) also suggested that the use of Rhaphidophlus over Thalysias was a pragmatic solution since the name was in current usage by the majority of contemporary authors. Conversely, Wiedenmayer (1977: 140), Hooper (1990a), Hooper et al. (1990, 1992) and Hooper & Levi (1993a) note that Carter (1876) merely compared three small specimens from the North Atlantic with Thalysias, but left them unnamed. Wiedenmayer (1977) states that although Carter did cite Duchassaing's (1850: pl. 17, fig. 1) figure of Thalysias subtriangularis, Carter's action cannot be construed as a subsequent designation of a type species under Articles 67c and 69a of the ICZN (Anonymous, 1985). Therefore, de Laubenfels' (1936a: 104) subsequent designation of Thalysias virgultosa (Lamarck, 1814) as type species of Rhaphidophlus is valid, and the genus is a junior synonym of Thalysias. 2) Simpson (1968a: 98) suggested that arguments supporting or refuting the choice of one genus name over another were irrelevant because Rhaphidophlus,Thalysias and Axocielita showed different cytological features, apparently justifying their recognition at the generic level. There are no arguments based on skeletal evidence which can contend with Simpson's hypothesis since his cytological data do not correlate with any skeletal features. However, Simpson's (1968a) conclusions are based on only three species, one in each genus, and it could be interpreted that the differences he observed may be applicable only at the species level. Simpson (1968a) found that Thalysias was distinct from both Microciona (cf. Hartman, 1955; Wells et al., 1960), Rhaphidophlus and Clathria (cf. Levi, 1960a). Thalysias was related to Rhaphidophlus, but cytologically distinct, which he stated was also apparently reflected in ectosomal cytological and morphological differences between the two genera. Rhaphidophlus REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 75 (sensu Simpson) had a specialised and extensive region containing a secondary fibre system and a continuous ectosomal skeleton, whereas Thalysias had fibre cell tracts organised into dermal columns which produce tufts of ectosomal styles. However, no similar correlations were observed in any other species examined (Van Soest, 1984b; Hooper, 1990a). Simpson's (1968a) system is intrinsically unworkable as it presently stands, given that the existing systematics is based on skeletal attributes, and essentially Thalysias and Rhaphidophlus differ only in their nomenclature. 3) Simpson (1968a) suggested that Axocielita was distinct from, but most closely related to Thalysias, but this conclusion is not supported by their morphology. Axocielita similis (Stephens) has a hymedesmoid skeleton with plumose fibre nodes, each node cored by plumose spicule tracts, echinating acanthostyles are absent and only one size class of auxiliary spicule is present, suggesting that the species is more closely related to Microciona than Thalysias (Hooper, 1990a). Simpson (1968a: 113) also agreed that echinating acanthostyles are of minor diagnostic importance. The implication of these data is that Axocielita harttnani Simpson and A. simi/is (Stephens) are either not cogeneric, which is not indicated by their morphological characters, or that morphological systematics is not corroborated by cell biology, and this is the main obstacle in using Simpson's results. 4) Simpson (1968a) also suggested that Clathria was more closely related to Rhaphidophlus and Thalysias in its cytological characteristics, than it was to the morphologically more-similar Microciona. This result conflicted with the system proposed by Levi (1960a), and developed further by Van Soest (1984b), which distinguished microcionid genera on the basis of their skeletal architecture and ectosomal characteristics, respectively. Nevertheless, it suggests that thinly encrusting microcionid sponges may have different cytological characteristics than ramose or massive forms, and this poses questions concerning environmental influences on cellular behaviour. 5) Hallmann (1920: 769) preferred Tenacia Schmidt (1870) to Rhaphidophlus Ehlers (1870) because t Tenacia had been firmly established by the redescription of T. clathrata Schmidt (Wilson, 1902: 397), whereas Rhaphidophlus was imperfectly known only from Ehlers' (1870: 18, 31) inadequate redescription of Spongia cratitia Esper. The argument is irrelevant, since S. cratitia is recognisable, and Thiele (1903a: 957) has already redescribed portions of the type material (ZMB4577, 4578), even though the whereabouts of the actual type specimen is presently unknown. Hallmann (1920) also suggested that Tenacia should be preferred to Rhaphidophlus because Schmidt's publication was abstracted in the Zoological Record for 1870, whereas Ehlers work was abstracted in 1872. There is no doubt that the genera are synonymous (Hartman, 1955: 176; Levi, 1960a: 56; Wiedenmayer, 1977: 140; Van Soest, 1984b: 91) but Rhaphidophlus has seniority over Tenacia, and Thalysias has seniority over both. It is included in Clathria (Thalysias). Seriatula Gray, 1867 Seriatula Gray, 1867: 515; de Laubenfels, 1936a: 122. (Refer to Ophlitaspongia). Sophax Gray, 1867 (Fig. 23D-E) Sophax Gray, 1867: 521; de Laubenfels, 1936a: 112. TYPE SPECIES. Microciona fallax Bowerbank, 1866: 128 (by monotypy) (lectotype BMNH1910.1.1.71; paralectotype BMNH1930.7.3.198; fragment of lectotype USNM5047). Encrusting growth form. Surface uneven, microscopically hispid. Choanosomal skeleton plumose (slightly plumoreticulate), with spongin fibres reduced to basal layer of spongin on substrate with ascending spongin fibre nodes cored by multispicular columns of choanosomal principal subtylostyles, and echinated by acanthostyles. Ascending fibres interconnected by sparse transverse spongin fibres, aspiculose or with few coring principal spicules. Ectosomal skeleton with tangential, paratangential or poorly developed brushes of subectosomal auxiliary subtylostyles. Megascleres long choanosomal principal subtylostyles with heavily microspined bases, entirely smooth, flexuous, sinuous or straight subectosomal auxiliary subtylostyles, and echinating acanthostyles of vaiiable size and evenly distributed spines. Microscleres absent. REMARKS. Gray (1867) erected Sophax for M. fallax, without further comment or comparison with other Microciona species. The type species has a Microciona-like hymedesmoid skeleton with erect spongin fibre nodes (similar to other nominal genera such as Abila, Axosuberites, Stylotellopsis, and Thalyseurypon), but unlike these it also has some transverse fibres intercon- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 76zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 23. Type species of microcionid genera. A-C, Rhaphidophlus (Spongia cratitia Esper, ZMB4577). D-E, Sophax (Microciona fallax Bowerbank, BMNH1910.1.1.71). F-G, Stylotellopsis (S. amabilis Thiele, ZMB3309). H-I, Tablis (Echinochalina anomala Hallmann, AMG10548). 77 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ necting multispicular tracts, producing a slightly plumoreticulate architecture. Sophax is referred here to Clathria (Microciona). Stylotellopsis Thiele, 1905 (Fig. 23F-G) species of Clathria (Thalysias). De Laubenfels (1936a) included the genus with Eurypon (Raspailiidae), but this is not upheld here: similarities between Stylotellopsis and Thalysias in their peripheral skeletal structure and the geometry of auxiliary spicules are closer than with the raspailiids (see Hooper, 1991). Stylotellopsis Thiele, 1905: 456; de Laubenfels, 1936a: 112. Tablis de Laubenfels, 1936 TYPE SPECIES. Stylotellopsis zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA am abilis Thiele, 1905: 456 (by monotypy) (holotype ZMB3309). (Fig. 23H-I) Thickly encrusting growth form. Surface hispid, uneven. Choanosomal skeleton hymedesmoid, with basal layer of spongin fibres lying on substrate and plumose non-anastomosing fibre nodes. Spongin fibre nodes cored by large echinating acanthostyles standing perpendicular to substrate, forming multispicular plumose tufts on basal skeleton. Subectosomal skeleton plumose, with multispicular columns of subectosomal auxiliary subtylostyles arising from ends of echinating spicule brushes, producing dendritic whispy tracts especially near periphery. Ectosomal skeleton without special spiculation but erect subectosomal auxiliary styles form nearly continuous palisade on surface. Mesohyl has heavy deposits of loose spongin. Megascleres single category of smooth subectsomal auxiliary styles with tapering bases and hastate points, and large echinating acanthostyles with swollen bases, large spines, and aspinose points. Principal spicules undifferentiated from auxiliary spicules. Microscleres absent. REMARKS. From published descriptions (Thiele, 1905; Topsent, 1928a; Koltun, 1964a) Stylotellopsis appears to be similar to Thalysias in having differentiated subectosomal and ectosomal auxiliary spicules (i.e., a specialised ectosomal skeleton), and Van Soest (1984b) synonymised the two genera on this basis. Re-examination of type material found that auxiliary spicules range greatly in their length, and principal spicules are very similar to auxiliary spicules scattered throughout the skeleton. The genus has a `microcionid' choanosomal skeletal structure (i.e., hymedesmoid with plumose fibre nodes), and an unusual subectosomal skeleton composed of dendritic or sinuous auxiliary spicule tracts. It also lacks microscleres (although several arcuate isochelae were seen in histological preparations of the holotype, but these are probably foreign). Stylotellopsis is interpreted here as being an encrusting (hymedesmoid) Tab/is de Laubenfels, 1936a: 76. TYPE SPECIES. Echinochalina anom ala Hallmann, 1912: 292 (by original designation) (holotype AMG10548). Massive, lobate-digitate growth form. Surface 'honeycomb' reticulate. Choanosomal skeleton reticulate, with heavy primary and secondary spongin fibres forming wide-meshed reticulation cored by multispicular tracts of subectosomal auxiliary subtylostyles. Coring spicules in primary fibres compressed, more dispersed in secondary fibres. Fibres heavily echinated by principal styles. Subectosomal skeleton with scattered auxiliary megascleres dispersed between fibre meshes and protruding through surface in paratangential bundles. Megascleres entirely smooth principal styles-subtylostyles, and smooth subectosomal auxiliary subtylostyles, often secondarily modified to tornotes. Microscleres absent. REMARKS. Tablis was erected for Whitelegge's (1907) specimen of Echinochalina glabra from Woolongong, NSW, a junior homonym of Ridley & Dendy's (1887) species renamed by Hallmann (1912) to E. anomala. This species has also been recorded from the Great Barrier Reef (Burton, 1934a). Three other species were referred to Tab/is by de Laubenfels (1936a): Echinochalina glabra Ridley & Dendy (a synonym of Spongia barba Lamarck), E. reticulata Whitelegge, both from SW Australia, and Spongia clavosa Lamarck. The last mentioned species is unknown: it was not mentioned in Topsent's (1933) revision of Lamarck's species nor found in the Lamarck collections (MNHN). These other 2 species differ from E. anomala in having microscleres but lacking an Holopsamma-like 'honeycombed reticulate' growth form. Tab/is is returned here to Echinochalina (Echinochalina). MEMOIRS OF THE QUEENSLAND MUSEUM 78^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Tenacia Schmidt, 1870 (Fig. 24A-C) TYPE SPECIES. Esperiopsis ^canaliculata Whitelegge, 1906: 471 (by monotypy) (lectotype AMG4325). Tenacia Schmidt, 1870: 56; Hallmann, 1920: 769; Topsent, 1920b: 17; Burton & Rao, 1932:337; Levi, 1960a: 56. Erect, branching, digitate growth form. Surface even, hispid, with oscules dispersed over lateral margins of branches. Choanosomal skeleton irregularly reticulate, nearly radial, with compressed axial and plumose extra-axial skeletons. In axis heavy spongin fibres cored by multispicular tracts of choanosomal principal styles, and in extra-axial skeleton principal styles protrude from peripheral fibres forming radial or plumose tracts, lying immediately below and supporting subectosomal skeleton. Echinating megascleres absent. Subectosomal skeleton plumose, with multispicular ascending tracts of subectosomal auxiliary styles at ends of peripheral choanosomal fibres. Ectosomal skeleton with thick continuous palisade of smaller ectosomal auxiliary styles standing perpendicular to surface. Megascleres smooth fusiform choanosomal principal styles, subectosomal auxiliary styles-subtylostyles with smooth or microspined bases, and ectosomal auxiliary styles-subtylostyles with microspined bases. Microscleres palmate isochelae of two sizes, including contorted forms, and oxhorn and wing-shaped toxas. TYPE SPECIES. Tenacia clathrata Schmidt, 1870: 56 (by monotypy) (fragment of type BMNH1870.5.3.156); = Spongia virgultosa Lamarck, 1814: 444 (fragment of type MNHNLBIMDNBE1344, 1338). Erect, arborescent, reticulate branching growth form. Surface highly conulose, uneven, microscopically hispid. Choanosomal skeleton irregularly reticulate, with well developed spongin fibres differentiated into ascending primary and transverse secondary fibres. Principal spicules confined to axis of primary fibres, but absent completely from secondary connecting fibres, and all fibres cored by paucispicular tracts of subectosomal auxiliary subtylostyles, and heavily echinated by small acanthostyles. Subectosomal skeleton plumose, with tracts of subectosomal auxiliary subtylostyles originating from deeper regions of choanosomal skeleton; subectosomal auxiliary subtylostyles also scattered abundantly throughout mesohyl between fibre meshes. Ectosome plumose, with brushes of smaller ectosomal auxiliary subtylostyles arising from ends of subdermal spicule brushes. Megascleres smooth choanosomal principal styles, smooth subectosomal auxiliary subtylostyles, smooth ectosomal auxiliary subtylostyles, and short thick echinating acanthostyles with aspinose points and necks. Microscleres palmate isochelae of two sizes, and wing-shaped, accolada and sinuous asymmetrical toxas. REMARKS. Tenacia and Rhaphidophlus are synonyms (Hallmann, 1920; Topsent, 1932; Levi, 1960a), and use of one name over another is merely a nomenclatural decision (see remarks for Rhaphidophlus). Topsent's (1932: 97) synonymy of T clathrata and Spongia juniperina Lamarck is not upheld here: the former is considered here to be a synonym of T virgultosa and restricted to Caribbean and NE. Atlantic populations, whereas T. juniperina (including T clathrata of Hallmann, 1912) is known only from the Indian Ocean. The genus is synonymised with Clathria (Thalysias). Tenaciella Hallmann, 1920 (Fig. 24D-E) Tenaciella Hallmann, 1920: 772; de Laubenfels, 1936a: 126. REMARKS. Hallmann (1920) erected Tenaciella for species like Tenacia(= Thalysias), but lacking echinating megascleres. Wells et al. (1960) also referred Esperiopsis obliqua George & Wilson to Tenaciella, but this species lacks dermal specialisation and is more closely allied with Axocielita (= Clathria). Ristau (1978) suggested that Artemisina archegona Ristau (USNM 24528) was similar to the type species of Tenaciella in having prominent subectosomal (extra-axial) spicule columns, and he suggested that the two species differed only in growth form, shape of the toxa microscleres and the appreciably more organised skeletal architecture in T canaliculata. However, differences observed in types of these species are more noticable than their similarities (e.g., ectosomal skeleton, fibre reticulation, degree of axial and extra-axial differentiation), and their supposed affinities (Ristau, 1978), are superficial. In its skeletal structure, differentiation of axial and extra-axial skeletons, fibre characteristics, the possession of similar extra-axial radial bundles of megascleres, and having a continuous ectosomal palisade of spicules the type species shows close similarities to Ceratopsion axifera (Hentschel) (Raspailiidae) from the Arafura Sea ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 79 (Hooper, 1991). In fact the resemblance between these two species is remarkable (although Esperiopsis canaliculata obviously belongs to Microcionidae, having isochelae and toxas, whereas Ceratopsion belongs to Raspailiidae in having long extra-axial spicules surrounded by bundles of ectosomal spicules and lacking chela or toxa microscleres). These structural similarities suggest a closer relationship between the two families, as proposed by Hooper (1991) in returning Raspailiidae to the Poecilosclerida, and subsequently supported by chemotaxonomic evidence (Hooper et al., 1992). In skeletal architecture E. canaliculata is identical to E. cylindrica, the type species of Axociella. Consequently, both Tenaciella and Axociella are synonymised here (the latter being the most senior available name), both having a compressed reticulate axis and plumose or plumo-reticulate extra-axial (subectosomal) skeletons, isochelae and toxa microscleres, lacking echinating spicules, and referred to Clathria (Axociella). Thalassodendron Lendenfeld, 1888 Thalassodendron Lendenfeld, 1888: 222; de Laubenfels, 1936a: 112. TYPE SPECIES. Thalassodendron typica Lendenfeld, 1888: 223 (by indication) (holotype unknown). Flabellate, cup-shaped growth form. Surface striated longitudinally. Choanosomal skeleton reticulate, with differentiated primary ascending and secondary transverse connecting spongin fibres; primary fibres cored by multispicular tracts of choanosomal principal styles, lightly echinated by acanthostyles (often secondarily incorporated into fibres), whereas secondary fibres aspiculose. Subectosomal and ectosomal skeletons unknown. Megascleres include smooth choanosomal principal styles, and short stout echinating acanthostyles. Microscleres unknown. REMARKS. This diagnosis is based on Lendenfeld's (1888) description of the type species which is virtually unrecognisable (de Laubenfels, 1936a). It is suspected that Thalassodendron typica Lendenfeld is a synonym of Echinonema typicum Carter, given Lendenfeld's propensity for describing other authors' species as his own 'new species'. However, it is not possible to associate any type materialwith the name `Thalassodendron typicum' and this synonymy remains doubtful. Echinonema typicum is also a synonym of Spongia cactifortnis Lamarck, in which case Thalassodendron would belong to Clathria (Thalysias) . Thalassodendron typica of Whitelegge (1901: 86; holotype AMZ958) is not the same as Lendenfeld's species, being a synonym of Echinodictyum mesenterinum (Lamarck) (Raspailiidae) (Hooper, 1991: 1379). Thalyseurypon de Laubenfels, 1936 (Fig. 24F-G) Thalyseurypon de Laubenfels, 1936a: 107; Wiedenmayer, 1977: 143. TYPE SPECIES. Spongia raphanus Lamarck, 1814: 444 (by original designation) (holotype MNHNLBIMDT572). Arborescent, reticulate, bushy growth form. Surface conulose, not hispid. Choanosomal skeleton more-or-less regularly reticulate, with heavy spongin fibres undifferentiated into primary or secondary components. Choanosomal fibres mostly aspiculose, or with irregularly paucispicular tracts of choanosomal principal subtylostyles, sparsely echinated by acanthostyles. Subectosomal skeleton poorly developed, consisting only of extra-fibre paratangential tracts of subectosomal auxiliary subtylostyles, becoming tangential in ectosomal region. Megascleres entirely smooth fusiform choanosomal principal subtylostyles, smooth subectosomal auxiliary subtylostyles, and small vestigially spined acanthostyles. Microscleres absent. REMARKS. Thalyseurypon was established for 8 species differing from Clathria only in lacking microscleres. Van Soest (1984b) proposed to merge the genus with Clathria, whereas Wiedenmayer (1977) synonymised it with Pandaros suggesting that the skeletal architecture and absence of microscleres in Spongia raphanus (sensu Topsent, 1932: 100) was similar to P. acanthifolium Duch. & Mich. However, re-examination of type material in both type specimens indicates that S. raphanus is most appropriately placed in Clathria (Clathria). Thalysias Duchassaing & Michelotti, 1864 (Figs 24A-C, 25A-B, 176-177) 1) Thalysias Duchassaing & Michelotti, 1864: 82; de Laubenfels, 1936a: 104; Hartman, 1955: 172; Wiedenmayer, 1977: 140. 2) Thalysias; Carter, 1876: 311; de Laubenfels, 1954: 137; Levi, 1960a: 52; Simpson, 1968a: 98. MEMOIRS OF THE QUEENSLAND MUSEUM 80zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TYPE SPECIES. 1) Interpretation by Wiedenmayer (1977), (i.e., Thalysias s.s.): Spongia virgultosa Lamarck, 1814 (by subsequent designation; de Laubenfels, 1936a: 104) (holotype MNHN missing, fragment BMNH1954.2.20.67); junior synonym of Spongia juniperina Lamarck, 1814 (lectotype MNHNLBIMDT570) (de Laubenfels, 1936a: 104). 2) Interpretation by Van Soest (1984b), i.e., Thalysias of authors: Thalysias subtriangularis Duchassaing, 1850 (by subsequent designation; Carter, 1876: 311); = Xestospongia subtriangularis (Wiedenmayer, 1977: 255). DEFINITION OF TYPE SPECIES. As for Tenacia. REMARKS. The nomenclatural complexities of Thalysias have been discussed above (see Rhaphidophlus and Tenacia). According to Van Soest (1984b), Carter's (1876) statement that T subtriangularis was 'representative' of the genus means that Thalysias sensu Carter is a haplosclerid, but this is not a valid subsequent designation. Wiedenmayer (1977) stated that de Laubenfels' (1936a) subsequent designation of T. virgultosa as the type, which makes Thalysias (established 1864) a senior synonym of Rhaphidophlus (established 1870). Moreover, Tenacia is an objective synonym of Thalysias by synonymy of their respective type species. This has been confirmed by type material (although the holotype of Spongia virgultosa is only represented by a fragment in the BMNH). Thalysias is used here as the earliest available subspecific name for Clathria with differentiated ectosomal and subectosomal spiculation (i.e., specialised ectosomal skeleton). Wetmoreus de Laubenfels, 1936 Wetmoreus de Laubenfels, 1936a: 112. TYPE SPECIES. Microciona novaezealandica Brondsted, 1924: 463 (by original designation) (holotype UZM not found). Encrusting growth form. Surface shaggy, hispid. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer of spongin on substrate with plumose non-anastomosing fibre nodes. Fibre nodes cored by erect multispicular bundles of choanosomal principal subtylostyles forming ascending plumose skeletal columns, and echinating acanthostyles at oblique angles to skeletal columns, usually forming brushes. Subectosomal skeleton paratangential or plumose, with single category of subectosomal auxiliary subtylostyles forming light dermal brushes erect on surface, or scattered individually on surface and throughout mesohyl. Megascleres choanosomal principal subtylostyles with prominent basal spines, smooth subectosomal auxiliary subtylostyles-styles, and small evenly spined echinating acanthostyles. Microscleres allegedly include both palmate and arcuate forms. Toxas absent. REMARKS. This definition is from Brondsted's (1924) description of the type. The holotype is possibly extant in Brondsted's collection at the UZM, but has not been located (0. Tendal, pers. comm.). Wetmoreus differs from Microciona (s.s.) in allegedly having both palmate and arcuate isochelae, and lacking toxas (see remarks for Paradoryx), but this cannot be verified. It is possible that both forms of chelae are merely modified (curved, thickened) palmate forms. Wetmoreus is included here in Microciona based on the possession of a hymedesmoid basal skeleton with plumose fibre nodes. Wilsonella Carter, 1885 (Fig. 24H-I) Wilsonella Carter, I885f: 366; Hallmann, 1912: 237, 1920: 768; Topsent, 1928a: 62; de Laubenfels, 1936a: 109. TYPE SPECIES. Wilsonella australiensis Carter, 1885f: 366 (by monotypy) (holotype BMNH1886.12.15.43). Erect, massive or flabelliform growth forms. Surface prominently conulose at apex of sponge, with conspicuously raised oscules. Choanosomal skeleton reticulate, with moderately light spongin fibres forming irregular meshes with distinct primary ascending and secondary transverse lines. Primary fibres cored by paucispicular tracts of robust auxiliary subtylostyles and abundant detritus, and heavily echinating by acanthostyles particularly at fibre nodes. Secondary connecting fibres with paucispicular tracts, little detritus and lightly echinated. Ectosomal skeleton lightly arenaceous, mostly with spicule fragments, lacking specialised spiculation but with light tangential or paratangential tracts of more slender subectosomal auxiliary subtylostyles. Megascleres fusiform robust auxiliary subtylostyles with smooth bases and hastate or telescoped points (inside fibres), more slender auxiliary subtylostyles with spines on both bases and points (outside fibres), and echinating acanthostyles with even spination. Microscleres palmate isochelae and wing-shaped toxas. REMARKS. Wilsonella is characterised by auxiliary megascleres coring fibres (i.e., without REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 81 true choanosomal principal spicules), incorporation of sand and foreign spicule detritus into the skeleton, and lacking any differentiation between ectosomal and subectosomal skeletons. Wilsonella australiensis is unusual amongst 'sandy' microcionid sponges in that subectosomal auxiliary styles outside fibres (e.g., in the ectosomal skeleton) are geometrically very similar to those coring fibres differing only in having spines on both their bases and points. These spicules are characteristic of the type species (although they are also known in several other microcionids, such as Clathria (C.) chelifera and Clathria (T) major). Even though spicules coring the fibres cannot be construed as being true principal megascleres, it must be concluded that this species has two distinct categories of auxiliary spicules. In this respect it is contrasted with Clathria (Dendrocia), which has a completely plumose skeletal architecture, including ectosomal and subectosomal differentiation, but only a single undifferentiated category of structural megascleres throughout the skeleton. Species referred to Paradoryx by Hallmann (1920) are similar to Wilsonella in having only auxiliary megascleres in the skeleton, but these have arcuate-like chelae instead of palmate isochelae, more than one category of structural megasclere (i.e., Clathria (Clathria)), or only 1 category of auxiliary spicules and a plumose architecture (i.e., Clathria (Dendrocia)). In contrast, the type species of Wilsonella incorporates detritus into spongin fibres, and this character is interpreted as probably being phylogenetic rather than merely ecological despite being homoplasious througout Porifera (known also in Phorospongiidae, Raspailiidae, Ircinidae, Dysideidae, etc.), because it is correlated with distribution of megascleres within skeletal regions as described above. Contemporary authors (e.g., Levi, 1967b) have included Wilsonella as a synonym of Clathria, whereas in the present work the type species of Wilsonella is synonymised with Clathriopsamnza (both with detritus in the skeleton), the former being the senior name, and used as at the subgenus level, Clathria (Wilsonella). GENERA EXCLUDED FROM MICROCIONIDAE Acarnus Gray, 1867 Acarnus Gray, 1867: 544; Hooper, 1987: 71; Hiemstra & Hooper, 1991: 434; Van Soest et al., 1991: 49; Hooper & Levi, 1993a: 1222 (for full synonymy see Hooper, 1987). TYPE SPECIES. Acarnus innominatus Gray, 1867: 544 (by monotypy) (holotype BMNH not found). Thickly encrusting, massive, flabellate or vasiform growth forms. Surface smooth or minutely hispid, uneven, often sculptured with subdermal canals in encrusting forms. Choanosomal skeleton with short plumose tracts connected by renieroid reticulation of spongin fibres, reduced to plumo-reticulate skeletal tracts, or further reduced to plumose-halichondroid skeleton in encrusting forms. Skeletal tracts composed of uni- or paucispicular tracts of smooth choanosomal principal styles or subtylostyles, sometimes with spined bases. Fibres usually heavily invested with spongin, with granular collagenous around nodes of skeletal tracts, and fibres echinated by smooth and/or spined cladotylotes of one or two size categories, with or without additional, smaller echinating acanthostyles. Ectosomal skeleton with a tangential or paratangential layer of basally spined tylotes. Microscleres palmate isochelae and up to three distinct forms of toxas: oxhom, wing-shaped and accolada toxas. REMARKS. The type species concept follows Van Soest (1984b: 61) based on material from the Caribbean given that the holotype is missing from the BMNH. The type has two categories of cladotylote megascleres, a larger smooth and smaller spined variety, whereas other species are known to have various combinations of those spicules (Van Soest et al., 1991), together with acanthostyles in some species (A. (Acanthacarnus) Levi). Although the genus has now been exhaustively discussed (Hooper, 1987; Hiemstra & Hooper, 1991; Van Soest et al., 1991), its family placement is still debatable. In its microsclere complement (palmate isochelae, diverse toxas), differentiated principal and auxiliary spicules, echinating acanthostyles in some species (as well as having a highly modified second category of echinating spicules (cladotylotes) which are unique to the genus, derived from either acanthostyles (Hooper, 1987) or ectosomal tylotes (Hiemstra & Hooper, 1991)), the genus appears to have affinities with the Microcionidae (e.g., Burton, 1959; Levi, 1960a; Vacelet et al., 1976). Other authors (de Laubenfels, 1936a; Tanita, 1963; Hechtel, 1965; Bakus, 1966; Thomas, 1970a, 1973; Hoshino, 1981; Van Soest, 1984b; Hooper, 1987; Hiemstra & Hooper, 1991; Van Soest et al., ^ MEMOIRS OF THE QUEENSLAND MUSEUM 82zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 24. Type species of microcionid genera. A-C, Tenacia (Spongia juniperina Lamarck, MNHNDT570). D-E, Tenaciella (Esperiopsis canaliculata Whitelegge, AMG4325). F-G, Thalyseurypon (Spongia raphanus Lamarck, MNHNDT572). H-I, Wilsonella (W. australiensis Carter, BMNH1886.12.15.43). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 83 1991; Hooper & Levi, 1993a) suggest that true diactinal ectosomal spicules (tylotes with terminal spines and swollen tips) and a renieroid reticulation indicate relationship to Myxillidae (in the sense of Hartman, 1982). Hajdu et al. (1994) resurrected Iophonidae for Acarnus, Megaciella, Melonchela and others with microcionid-like spiculation (terminally spined megascleres of diverse categories, palmate isochelae, toxas) as well as ectosomal tylotes (previously considered diagnostic for the Myxillidae; Van Soest, 1984b). This revised interpretation de-emphasises the primary importance placed on skeletal structure (Bergquist & Fromont, 1988) for example), and allows for the inclusion of renieroid reticulate skeletons in several families (Iophonidae, Microcionidae, Raspailiidae, Phoriospongiidae, Cornuliidae, etc.). Skeletal structure would, therefore, be a highly homoplasious character but this interpretation does allow a consistent differentiation between Microcionidae and Iophonidae based on monactinal versus diactinal-derived ectosomal spicules, whilst also acknowledging their affinities based on the possession of similar microscleres through their inclusion together in Microcionina. Amphilectus Vosmaer, 1880 Amphilectus Vosmaer, 1880: 109; Ridley & Dendy, 1887: 123; Burton, 1929a: 428; Levi, 1960a: 55. TYPE SPECIES. Isodiciya gracilis Bowerbank, 1866: 149 (by subsequent designation of Burton, 1929a: 428) (holotype BMNH1877.5.21.754). Arborescent, dichotomously branched, stipitate growth form. Surface even, hispid. Choanosomal skeleton reticulate, with multispicular ascending primary fibres and uni- or paucispicular transverse connecting fibres, both cored by small styles. Subectosomal region with tracts of spicules projecting through surface. Ectosome membraneous, without specialised spiculation. Echinating megascleres absent. Megascleres small smooth styles of a single category. Microscleres palmate isochelae. Toxas absent. REMARKS. Vosmaer (1880: 109) established Amphdectus for a heterogeneous assemblage of 42 poecilosclerid species, most of which were related to Mycale, Esperiopsis (Mycalidae), Desmacidon (Phoriospongiidae) or Myxilla (Myxillidae). Ridley & Dendy (1887) restricted the genus to taxa with smooth styles and palmate isochelae, but even so, they remarked that the taxon was undoubtedly artificial. Burton (1929a) designated Vosmaer's first-named species as genotype, and suggested that in the strict sense (i.e., the above diagnosis) the genus had affinities with Esperiopsis. Levi (1960a) decided to abandon Amphilectus because, in the sense of Vosmaer (1880), it was too vague and served only as a catch-all taxon. In the broad sense Amphilectus contains some microcionid species (e.g., Microciona annata Bowerbank) whereas in the strict sense it fits with the concept of Myxillidae (Bergquist & Fromont, 1988; Hajdu et al., 1994). Caulospongia Kent, 1871 Caulospongia Kent, 1871: 616; Burton, 1930c: 673; de Laubenfels, I 936a: 118. Plectrodendron Lendenfeld, 1888: 66; Hallmann, 1914a: 306. TYPE SPECIES. Caulospongia verticillata Kent, 1871: 616 (by subsequent designation of Hallmann, 1914a: 306) (holotype BMNH1895.7.16.1); =Spongia perfoliata Lamarck, 1814: 439 (Topsent, 1932: 85) (lectotype MNHNLBIMDT582). Distinctive foliose growth form with lamellae arranged in whorls, or in plates, around an erect stalk. Surface even, minutely hispid. Choanosomal skeleton plumoreticulate, with well developed spongin fibres cored by pauci- or multispicular tracts of choanosomal principal tylostyles; longitudinal primary fibres form dendritic branches through axis, and ascending or oblique secondary fibres produce a nearly regular secondary reticulation. Subectosomal skeleton plumose, with erect brushes of choanosomal megascleres protruding through ectosome. Ectosome with tangential or paratangential crust of smaller tylostyles. Megascleres tylostyles of two sizes but with same geometry. Microscleres absent. REMARKS. Caulospongia has been included in Suberitidae (Hadromerida) by most authors since Burton (1930c). The type species is distinctive in growth form, and although it is common in NW Australian coastal waters it has been recorded in the literature only infrequently (Lamarck, 1814; Kent, 1871; Topsent, 1932; Hooper, 1984a). De Laubenfels (1936a: 118) defined Caulospongia in Ophlitaspongiidae presumably based on similarities to ophlitaspongiids (most now included in Microcionidae) in skeletal architecture (differentiated axial and extra-axial regions), localisation of spicules to different parts of the skeleton, and growth form (erect, whereas 'typical' suberitids are massive). However, these similarities are convergent. The presence of true tylostyles in Caulospongia (which are charac- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 84zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 25. Type species of microcionid genera. A-B, Thalysias (Spongia virgultosa Lamarck, BMNH1954.2.20.67). teristic of suberitids, as opposed to subtylote styles seen in many microcionids), and the absence of microscleres (which often provide useful clues on phylogenetic affinities) suggest that the genus should be retained in Suberitidae. Plectrodendron (type species Plectrodendron elegans Lendenfeld, 1888: 66, by monotypy) was also referred to Caulospongia by Hallmann (1914a: 306). Megaciella Hallmann, 1920 (Fig. 19A-B) Megaciella Hallmann, 1920: 772. TYPE SPECIES. Amphilectus pilosus Ridley & Dendy, 1886: 350 (by original designation) (lectotype BMNH1887.5.2.125). Lobate flabellate growth form. Surface shaggy, ridged, hispid. Choanosomal skeleton reticulate, with ascending multi- or paucispicular tracts of choanosomal principal styles, interconnected by secondary uni- or bispicular tracts within light spongin, producing irregular wide meshes. Echinating spicules absent. Subectosomal skeleton radial or plumose, with erect choanosomal principal styles protruding from peripheral fibres through surface. Ectosome with tangential or paratangential layer of tylotes, often in bundles. Megascleres very large smooth choanosomal principal styles, and ectosomal tylotes, often curved or sinuous, with slightly swollen microspined bases. Microscleres minute palmate isochelae and two sizes of toxas (very large accolada and wing-shaped). REMARKS. This diagnosis is based on the lectotype which differs slightly from the original description of Ridley & Dendy (1886, 1887). Specifically, the so-called 'long thin centrally curved oxeas' are very large accolada toxas with slight central curvature and slightly reflexed hastate points, some of which exceed 100p,m long (i.e., larger than the principal styles). Similarly, these toxas are not associated with the ectosomal skeleton but are scattered throughout the mesohyl, whereas bundles of `oxeas' described by Ridley & Dendy (1887) are actually bundles of ectosomal tylotes. A second morph of toxas is 85 ^ REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR also present, being large and wing-shaped, and these too have hastate points. Hallmann (1920) suggested that a lax skeletal construction and supposedly comparable spiculation indicated affinities between Megaciella and Artemisina. He noted that Megaciella differed from that genus in lacking ornamentation on toxas, which he considered was a significant diagnostic character at the generic level. However, in regard to this latter character, it has been found that a number of microcionid species have distally spined toxas (see remarks for Clathria), and in any case, both genera have quite different ectosomal skeletons. On the basis of its ectosomal characteristics (consisting of basally spined diactinal tylote spicules), Van Soest (1984b) referred Megaciella to the Myxillidae, whereas Artemisina (with a monactinal ectosomal skeleton) was retained in Microcionidae. Under the revised scheme proposed by Hajdu et al. (1994) Megaciella is included in Iophonidae, with palmate isochelae and toxas which being the only real synapomorphy between Megaciella and the Microcionidae (i.e., Microcionina). Melonchela Koltun, 1955 (Fig. 19C-E) Melonchela Koltun, 1955a: 17, 1959: 187. TYPE SPECIES. Melonchelaclathrata Koltun, 1955a: 17 (by original designation) (paratype BMNH1963.7.29.7). Arborescent, reticulate planar, branching growth form. Surface even, microscopically hispid. Choanosomal skeleton plumose, with bundles of independent ascending fibres cored by large and small choanosomal principal styles. Ascending fibres not interconnected. Echinating acanthostyles absent. Subectosomal skeleton plumose, with principal styles projecting through surface. Ectosome with tylotes forming tangential layer or erect brushes on surface. Megascleres large and small choanosomal principal styles, with smooth or microspined bases, and diactinal ectosomal tylotes (swollen bases) and strongyles (rounded bases) with microspined bases. Microscleres abundant palmate cleistochelae, palmate isochelae, small wing-shaped toxas, and oxhorn toxas with spines, mucronate points, or telescoped points, or simply with subterminal ridge. REMARKS. This species is remarkable in several features: its erect planar reticulate growth form (superficially resembling the microcion id Clathria coppingeri and the raspaili id Echinodic- tyum cancellatum); extremely large size range of principal spicules, the largest protruding a long way through fibre bundles, reminiscent of Raspailiidae; the apparent lack of connecting fibres between the ascending plumose spicule tracts; a ridge-like subterminal ornamentation on toxas; and extremely abundant tracts of chelae microscleres throughout the mesohyl. Diactinal ectosomal spicules (tylotes, strongyles and intermediates, varying in the degree of swelling of their bases), palmate isochelae and toxas indicates that the species has affinities to Acarnus and Megaciella in the Iophonidae. Naviculina Gray, 1867 (Fig. 19H-I) Naviculina Gray, 1867:538; de Laubenfels, 1936a: 88. TYPE SPECIES. Naviculina cliftoni Gray, 1867: 538 (by monotypy); for Ilymedesmia sp. nov.' of Bowerbank, 1864: 252 (fragment of type BMNH1877.5.21.270). Growth form and surface details unknown. Ectosomal features unknown. Choanosomal skeleton evenly reticulate, wide meshed, composed of tracts of subtylostyles bound together with nodal spongin, with multispicular tracts several spicules wide interconnected by uni- or paucispicular tracts only one spicule wide, both producing even triangular meshes. Megascleres single category of subtylostyles. Microscleres cleistochelate anisochelae, dispersed throughout mesohyl particularly between fibre anastomoses. REMARKS. Naviculina is monotypic, and the type species, from Fremantle is characterised by cleistochelae ('naviculoid spiculum' of Bowerbank, 1864), but nothing else was known about the species. de Laubenfels (1936a) considered Naviculina dubious, with alleged affinities to Plocamiopsis (having cleistochelae), although several other nominal genera are also known to have cleistochelate microscleres, interpreted as modified palmate isochelae (Colloclathria, Plocamiopsis and Quizciona of the Microcionidae; and Melonchela of the Iophonidae). The holotype no longer exists. The holotype slide (Bowerbank, 1864; Gray, 1867) contains a small section of the skeleton (Fig. 19H-I), but enough detail to indicate the Mycalidae, probably Arenochalina. Paracornulum Hallmann, 1920 Paraconiulum Hallmann, 1920: 772. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 86zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA TYPE SPECIES. Cornu/um dubium Hentschel, 1912: 346 (by original designation) (holotype SMF964). Encrusting growth form. Surface smooth, even. Choanosomal skeleton hymedesmoid, with spongin fibres lying on substrate and bases of acanthostyles embedded and spicules standing perpendicular to surface. Subectosomal skeleton radial or plumose, with ascending tracts of erect tornotes. Acanthostyles also scattered throughout mesohyl. Ectosome with tangential tracts of tornotes. Megascleres radial tornotes (with fainly rugose bases) and principal acanthostyles (with some oxeote or strongylote modifications). Microscleres palmate isochelae and oxhorn toxas. REMARKS. Paracornulum does not appear to be as closely related to microcionids as inferred by Hallmann (1920), showing similarities in possession of echinating acanthostyles, hymedesmoid basal spongin fibres, palmate isochelae and toxas. Two other species referred by Hallmann to this genus (Cliona purpurea Hancock and Suberites fuliginosus Carter) are very poorly known and further comment on those taxa must await redescription of relevant type material. Based on its ectosomal characteristics and megasclere spiculation Paracornulutn was subsequently referred to Cornulidae (Levi & Levi, 1983a: 966). The type species appears close to Cornulum Carter, lophon Gray and Zyzzya de Laubenfels because the ectosomal tornotes are reduced tylotes with rugose (i.e., vestigially microspined) bases. These genera are referred to Iophonidae (Hajdu et al., 1994). Paresperia Burton, 1930 Paresperia Burton, 1930a: 501. TYPE SPECIES. Paresperia intermedia Burton, 1930a: 501 (by monotypy) (holotype BMNH1910.1.1.912). Irregularly massive, low growth form. Surface even, smooth. Choanosomal skeleton reticulate, with loose, unispicular, irregular reticulation of light spongin fibres cored by auxiliary subtylostyles, of same geometry as those in subectosomal and ectosomal skeletons. Echinating acanthostyles absent. Subectosomal and ectosomal skeletons lack specialised spiculation, but have loose tangential reticulation of subectosomal auxiliary megascleres. Megascleres only auxiliary subtylostyles, entirely acanthose or only basally spined. Microscleres palmate isochelae. Toxas absent. REMARKS. Burton (1930a) assigned Paresperia to the Microcionidae based on supposed affinities to Artemisina (sensu Burton), having an irregular, confused skeletal architecture composed of undifferentiated choanosomal and ectosomal megascleres. The genus differed from Artemisina (s.s.) in having lightly and evenly spined auxiliary megascleres and lacking toxas. Burton also noted that Paresperia had affinities with the Mycalidae, whereas Van Soest & Stone (1986) suggested that the presence of a unispicular choanosomal reticulation of acanthostyles and palmate isochelae placed the genus closer to Esperiopsis (placement still controversial in either Mycalidae or Esperiopsidae) than to other microcionids. Querciclona de Laubenfels, 1936 Querciclona de Laubenfels, 1936a: 46. TYPE SPECIES. Antherochalina quercifolia Keller, 1889: 383 (by original designation) (holotype ZMB429). Erect, arborescent, flabellate growth form. Sur- face even, hispid, regularly porous with oscules. Choanosomal skeleton reticulate, with heavy spongin fibres producing tight and irregular meshes, fibres cored by choanosomal principal styles which form primary ascending multispicular (eventually plumose) tracts, and secondary transverse uni- or paucispicular tracts, together producing a subisodictyal reticulation. Axial skeleton slightly different from extra-axial region: near axis skeletal architecture isodictyal, whereas towards periphery skeleton plumose. Echinating megascleres absent. Subectosomal skeleton plumose, with brushes of choanosomal principal styles protruding through surface. Ectosomal skeleton with brushes of smaller choanosomal styles surrounding one or few larger subectosomal styles. Megascleres small smooth choanosomal principal styles, and long smooth subectosomal auxiliary styles. Microscleres absent. REMARKS. Querciclona seems to be a case where inadequate primary taxonomy has been badly misinterpreted. The genus is restricted here to include only A. quercifolia, and its association with the Microcionidae rests solely upon the original placement of the type species in Antherochalina (whereby many of the species originally described for the genus by Lendenfeld (1888) are true microcionids). By comparison, de Laubenfels (1936a) originally intended Quer- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 87 ciclona to include Haliclona-like species (Haplosclerida), but this is only true for a second species, Arenochalina arabica Keller (which was subsequently transferred to A renosclera PulitzerFinali (1982), but is probably an arenaceous Callyspongia). Conversely, the type species belongs to the Axinellidae. Antherochalina quercifolia is structurally close to Isociella (s.s.) and Phakellia of authors. It has an almost regular isodictyal choanosomal reticulation of spongin fibres cored by primary and secondary skeletal lines, without any pronounced compression of the axial skeleton but with some differentiation of the axial and extraaxial skeletons, and it also lacks a specialised ectosomal skeleton. Details of the holotype (reexamined and redescribed above) bear little resemblance to Keller's (1889) original description, and it is concluded that Querciclona is most appropriately referred to Phakellia (Axinellidae) (Hooper & Levi, 1993b). A specimen label in M. Burton's handwriting accompanying a fragment of the holotype in the BMNH (BMNH1908.9.24.179) suggests further that A. quercifolia is a synonym of Phakellia donnani (Bowerbank), but this synonymy is so far unsubstantiated. Scopalina Schmidt, 1862 Scopalina Schmidt, 1862: 78; Gray, 1867: 535. TYPE SPECIES. Scopalina lophyropoda Schmidt, 1862: 79 (by monotypy) (holotype LMJG15117/154). Thickly encrusting growth form. Surface prominently conulose. Choanosomal skeleton hymedesmoid, with heavy spongin fibres forming basal layer of spongin on substrate, with ascending non-anastomosing fibre nodes cored by plumose brushes of auxiliary styles. Echinating megascleres absent. Ectosome lacks specialised spiculation, although brushes of auxiliary styles protrude through surface. Mesohyl matrix heavy. Megascleres only a single category of long auxiliary style. Microscleres absent. REMARKS. Gray (1867) and de Laubenfels (1936a) suggested that Scopalina was Microciona-like with plumose skeletal architecture and greatly reduced spiculation. Although skeletal architecture is hymedesmoid containing non-anastomosing ('microcionid') fibre nodes, and the possession of only a single category of auxiliary spicules could be construed as reduction of the typical Microciona condition (analogous to Dendrocia (Microcionidae) or Amphilectus s.l. (Myxillidae)), no other characters support its inclusion in the Microcionidae. Boury-Esnault (1971) and Uriz (1982) include it in the Halichondrida, and suggested close affinites with Stylinos. KEY TO GENERA 1(0). Choanosomal skeleton more-or-less undifferentiated, unstructured ^ Artemisina Choanosomal skeleton well structured, hymedesmoid to reticulate, but lacking any differentiated components 2 Choanosomal skeleton well structured, predominantly reticulate, differentiated into two distinct components ^ 7 2(1). Choanosomal fibres or skeletal tracts cored by one or more category of principal spicules . ^ 3 Choanosomal fibres or skeletal tracts cored by auxiliary spicules but partially or wholly 5 replaced by detritus ^ Choanosomal fibres or skeletal tracts cored by auxiliary spicules identical to those in ectosomal and subectosomal skeletons ^ 6 Choanosomal fibres or skeletal tracts cored by auxiliary spicules different from those in peripheral skeleton ^ Panda ros 3(2). Choanosomal skeleton without any marked axial compression or differentiated axial and extra-axial regions 4 Choanosomal skeleton with noticeably compressed axis and well differentiated axial and extra-axial (radial, plumose or plumoreticulate) regions ^ Clathria (Axociella) Choanosomal skeleton hymedesmoid or microcionid, with basal layer of spongin lying on substrate (with or without ascending, nonanastomosing fibre nodes), and bases of principal spicules standing perpendicular to substrate ^ Clathria (Microciona) Choanosomal skeleton evenly renieroid reticulate throughout with well developed spongin fibres cored by smooth principal styles Clathria (Isociella) ^ 4(3). With single category of (subectosomal) auxiliary spicule on surface forming tangential, paratangential or plumose tracts ^ Clathria (Clathria) With two categories of auxiliary spicules, smaller ectosomal spicules generally overlaying larger subectosomal spicules forming discrete bundles or continuous palisade on surface ^ Clathria (Thalysias) 5(2). Special category of (acantho)styles present echinating fibres, differentiated from principal spicules ^ Clathria (Wilsonella) MEMOIRS OF THE QUEENSLAND MUSEUM 88zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Echinating spicules styles or acanthostyles undifferentiated from principal spicules coring sponHolopsamma gin fibres 6(2). Special category of (acantho)styles present echinating fibres, differentiated from principal spicules ^ Clathria (Dendrocia) Echinating spicules styles or acanthostyles representing principal spicules, but different from those coring fibres ^ Echinochalina (Echinochalina) Echinating spicules oxeas or anisoxeas representing principal spicules, but different from those coring fibres ^ Echinochalina (Protophlitaspongia) 7(1). Primary skeleton renieroid cored by axially or basally compressed tracts of acanthostyles, secondary skeleton cored by smooth principal styles in plumose, subisodictyal or plumoreticulate tracts ^ Antho (Antho) Primary skeleton renieroid cored by axially or basally compressed tracts of acanthostrongyles, secondary skeleton cored by smooth principal styles in plumose, subisodictyal or plumoreticuAn (ho (Plocamia) late tracts Primary skeleton axially compressed spongin fibres cored by renieroid tracts of sparsely spined principal styles intermingled with plumose or plumoreticulate tracts of smooth principal styles, overlaid by secondary extraaxial plumose skeleton cored by larger smooth principal styles ^ Antho (lsopenectya) Primary renieroid reticulate skeleton cored by smooth principal styles and echinated by identical spicules, with secondary radial extra-axial skeleton on exterior edge of skeleton only cored by larger smooth principal styles Echinoclathria ^ DESCRIPTION OF AUSTRALIAN SPECIES Clathria Schmidt, 1862 Refer to subgenera for synonymy. TYPE SPECIES. Clathria compressa Schmidt, 1862: 58 (designated Schmidt, 1864: 35). DEFINITION. Monactinal auxiliary spicules in 1 or 2 categories forming ectosomal skeletons ranging from sparse, mostly membraneous (C. (Microciona)), sparse, tangential (C. (Clathria)) to relatively dense, erect (C. (Thalysias)). Choanosomal skeletal tracts usually enclosed within spongin fibres, sometimes simply with nodal spongin; fibres cored by smooth, basally spined or partially spined principal monactinal megascleres, usually geometrically different from auxiliary megascleres, sometimes secondarily lost and cored by single category of auxiliary subtylostyle (C. (Dendrocia)), or sometimes replaced partially or fully by detritus in fibres (C. (Wilsonella)). Echinating megascleres partially or entirely acanthose, occasionally smooth or vestigial spination, sometimes secondarily lost (C. (Axociella), C. (Isociella)). Choanosomal structure ranges from leptoclathrid to microcionid plumose (C. (Microciona)), renieroid (C. (Isociella)), plumoreticulate or reticulate, with (C. (Axociella)) or without compressed axis and radial extra-axial regions. Microscleres include palmate isochelae and modified forms, and toxas with smooth or spined points, occasionally absent. REMARKS. This definition is necessarily broad to encompass the 7 subgenera included in Clathria, showing a wide spectrum of character states, most of which are interpreted as secondary losses rather than unique apomorphies, and many characters show intermediate states making it virtually impossible to maintain generic boundaries recognised by earlier authors. Clathria (Clathria) Schmidt, 1862 Clathria Schmidt, 1862: 57. Allocia Hallmann, 1920: 768. Antherochalina Lendenfeld, 1887b: 741. Bipocillopsis Koltun, 1964a: 79. Diclyociona Topsent, 1913a: 579. Labacea de Laubenfels, 1936a: 125. Ligrota de Laubenfels, 1936a: 125. Litaspongia de Laubenfels, 1954: 162. Marleyia Burton, 1931a: 346. Ramoses de Laubenfels, 1936a: 109. Thalyseurypon de Laubenfels, 1936a: 107. TYPE SPECIES. Clathria compressa Schmidt, 1862: 58 (by subsequent designation of Schmidt, 1864). DEFINITION. Ectosomal skeleton composed of a single undifferentiated category of auxiliary megasclere; choanosomal skeletal structure plumoreticulate or reticulate, usually without marked difference between axial and extra-axial regions; spongin fibres cored by completely smooth, basally spined or partially spined principal megascleres, geometrically differentiated from auxiliary megascleres, but sometimes secondarily lost; echinating megascleres entirely or partially acanthose, occasionally smooth, sometimes secondarily lost. Microscleres include palmate isochelae and modified forms, and toxas with smooth or spined points. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 89 TABLE 1. Comparison between present and published records of Clathria (Clathria) a ngulifera Dendy.Measurements in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Holotype (NMO2286) (Victoria) Specimens (N=2) (SE.Queensland) Choanosomal principal styles 1444172.4)-211 x 3-(4.3)-6 Subectosomal auxiliary styles 168-(175.0)-254 x 2.5-(3.6)-4.5 x 3.5-(4.1)-4.5 128-(141.3)-150 x 3-(3.8)-5 181-(195.8)-223 x 342.3)-5 58-(61.8)-72 x 2-(3.3)-5 Chelae 5.5-(7.1)-8.5 5-(6.7)-9 Toxas 18467.4)-101 x 0.541.7)-3.3 absent Echinating acanthostyles 58-(64.2)-72 REMARKS. Of the 154 named species described in, or subsequently referred to Clathria, or one of its synonyms listed above, 112 are retained in this subgenus. There are 31 (2 new) species known from Australian waters. Clathria (Clathria) angulifera Dendy, 1896 (Figs 26-27, Plate 1A, Table 1) Clathria anguhfera Dendy, 1896: 32; Ayling et al., 1982: 100; Hooper & Wiedenmayer, 1994: 258. Thalysias angulifera; de Laubenfels, 1936a: 104. cf. Clathria conectens Hallmann, 1912: 247. MATERIAL. HOLOTYPE: NMVG2286 (fragment BMNH1902.10.18.323): Outside Port Phillip Heads, Vic, 38°09'S, 144°52'E, coll. J.B. Wilson (dredge). OTHER MATERIAL: Queensland — QMG303230, QMG303960. HABITAT DISTRIBUTION. Submerged rock reef; 10-30m depth; Bass Strait (Vic), N. Stradbroke I. and Noosa Heads (SEQ) (Fig. 260). DESCRIPTION. Shape. Thickly lamellate, ap- proximately 9 cm long, 30mm wide, 50mm high, with numerous bulbous lobate digits up to 15mm long. Colour Bright yellow-orange alive (Munsell 2.5Y 8/10), grey-brown in ethanol. Oscules. Occasional large oscules, 1.5 3mm diameter, on edges and between lobes. Texture and surface characteristics. Firm, compressible; glabrous skin-like surface. Ectosome and subectosome. Ectosomal skeleton membraneous, with loose, irregular, tangential, occasionally paratangential or erect tracts of subectosomal auxiliary subtylostyles; rarely protruding above surface. Subectosomal portion of peripheral skeleton slightly plumose, with sparse - diverging brushes of auxiliary megascleres which arise from ascending choanosomal tracts. Choanosome. Choanosomal skeleton regularly reticulate, cavernous, vaguely renieroid; fibre anastomoses produce wide oval, rectangular or sometimes slightly triangular meshes; spongin fibres thin, only lightly invested with spongin, barely differentiated from mesohyl matrix; spongin fibres cored by uni- or paucispicular tracts of choanosomal principal megascleres; echinating acanthostyles sparsely dispersed on fibres; mesohyl matrix heavy, lightly pigmented, containing few straight or sinuous subectosomal auxiliary megascleres and abundant spherical incubated larvae (275-34511m diameter) with well differentiated cellular development. Megascleres (Table 1). Choanosomal principal styles short, thin, fusiform, straight or slightly curved towards basal end, with rounded or only slightly subtylote, smooth bases. Subectosomal auxiliary subtylostyles straight, thin, relatively long, almost hastate, with smooth and only slightly subtylote bases. Acanthostyles small, thin, prominently subtylote, with rudimentary granular spination and small aspinose 'neck' near basal constriction. Microscleres (Table 1). Palmate isochelae very small, weakly silicified, unmodified. Toxas not common, relatively large, v-shaped, with sharply angular central curvature and straight arms. REMARKS. The thickly lamellate, lobate growth form, glabrous surface, relatively open, slightly renieroid, paucispicular skeletal structure (cf. Dendy, 1896), and aspects of spiculation in C. (C.) angulifera are quite distinctive and unusual amongst microcionids (cf. Hallmann, 1912). Many of these features are similar to those in C. (T)aphylla sp. nov. from the Houtman Abrolhos, WA, although the latter has a specialised ectosomal skeleton and hence is included in C. (Thalysias) rather than C. (Clathria). These species together with C. (C.) hispidula are sister species belonging to a species group termed here `angulifera' group. They are compared in detail in discussion on C. (T) aphylla. This species is also slightly reminiscent of C. (C.) conectens, differing in that megascleres coring fibres are true principal spicules rather than just undifferentiated auxiliary spicules (albiet these differences are not as well marked in C. (C.) angulifera as in many other species of Clathria). This reduction in coring spicules was interpreted by Hallmann (1912) to be charac- MEMOIRS OF THE QUEENSLAND MUSEUM 90zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 26. Clathria (Clathria) angulifera Dendy (holotype NMVG2286). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Palmate isochelae. E, V-shaped toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 91 FIG. 27. Clathria (Clathria) angulifera Dendy (A-B, G, holotype NMVG2286; C-E, QMG303230). A, Choanosomal skeleton. B, Fibre characteristics (x456). C, Ectosomal paratangential skeleton. D, Palmate isochelae. E, Echinating acanthostyles. F, Acanthcstyle spines. G, Principal styles and v-shaped toxas. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 92zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA teristic of C. (Wilsonella) (although this is shown here to be an erroneous interpretation of Wilsonella). De Laubenfels (1936a) also transferred C. (C.) angulifera to Thalysias, but this was completely unjustified. Although known only from 2 widely separated localities it is probable that it is more widely distributed within shallow coastal waters in southeastern Australia. Clathria (Clathria) arcuophora Whitelegge, 1907 (Figs 28-29) Clathria arcuophora Whitclegge, 1907: 491,500-501, p1.45, fig.29; Hallmann, 1912: 211, 229, 234- 237, 260, 263, text-fig. 49-49a; Hooper & Wiedenmayer, 1994: 258. Thalysias arcuophora; de Laubenfels, 1936a: 105. Microciona arcuophora; Vosmaer, 1935a: 611, 649, 665. MATERIAL. LECTOTYPE: AMG4346: Off Barranjoey, S. coast of NSW, 33 °35'S, 151 °20'E, 50-66m depth, coll. FIV `Thetis' (dredge). PARALECTOTYPE: AMZ1209: Off Botany Bay, NSW, 34°00'S, 151°11'E, 40-46m depth, coll. FIV `Thetis' (dredge). HABITAT DISTRIBUTION. Depth range 30-90m; substrate unknown; central and S. coast of NSW (Fig. 28G). DESCRIPTION. Shape. Thinly flabelli form, 170mm long, 120mm maximum breadth, up to 4mm thick, with long cylindrical stalk, 90mm long, 13mm diameter, and rounded or slightly lobate margins. Colour Grey-brown in dry state. Oscules. Surface covered with evenly distributed minute oscules, up to 1.5mm diameter. Texture and surface characteristics. Harsh in dry state; surface optically smooth. Ectosome and subectosome. Ectosome microscopically hispid, with regularly distributed choanosomal principal styles forming erect plumose brushes, and protruding from peripheral fibre endings; tangential layer of subectosomal auxiliary spicules lying on or just below surface, at base of principal styles. Subectosome with only slightly plumose tracts of choanosomal megascleres, virtually undifferentiated from choanosomal skeleton. Choanosome. Choanosomal skeleton more-orless regularly reticulate, sub-renieroid; axial skeleton slightly compressed, peripheral skeleton slightly plumo-reticulate; branching between moderately heavy spongin fibres produces ovoid to square meshes, and fibres differentiated into primary ascending and secondary transverse components; primary fibres with pauci- to multispicular core of choanosomal principal styles; secondary fibres with uni- or bispicular tracts of principal spicules; echinating acanthostyles sparse, slightly more abundant on peripheral fibres; mesohyl matrix light, with only few subectosomal auxiliary megascleres dispersed between fibres. Megascleres. Choanosomal principal styles thick, slightly curved, slightly fusiform, with rounded smooth bases. Length 1764258.4)444p,m, width 12-(22.2)-27p.m. Subectosomal auxiliary subtylostyles small, thin, straight, with smooth or microspined bases, bases slightly subtylote, points fusiform. Length 1924231.1)-276pm, width 3.5-(4.8)-6pm. Acanthostyles small, slightly subtylote, with relatively even granular spination. Length 68(93.2)-104pm, width 6.547.5)-10p.m. Microscleres. Palmate isochelae large, unmodified. Length 16-(21.4)-26p.m. Toxas intermediate between oxeote and oxhorn, thick, gently curved at centre, with straight arms or slightly reflexed points. Length 28(104.0)-132p.m, width 1.5-(4.8)-6pm. REMARKS. This species is distinctive by its renieroid choanosomal skeleton, plumo-reticulate subectosomal skeleton, distinctive spongin fibres which contain only few but very thick coring spicules, and an ectosomal region which is dominated by plumose brushes of principal spicules. The renieroid skeletal construction is superficially similar to Antho (lsopenectya) and Clathria(lsociella)), and several other (otherwise unrelated) groups (some Raspailiidae (e.g., Ectyoplasia), Axinellidae (e.g., Axinella aruensis (Hentschel, 1912)), Iophonidae (e.g., Acarnus) and Myxillidae (Lissodendoryx) (see also remarks for C. (C.) crassa)). Clathria (C.) arcuophora is most reminiscent of Ectoplasia frondosa (Lendenfeld) (Raspailiidae; see Hooper, 1991: figs 47-48), which was originally described in the Microcionidae, having closely comparable architecture, fibre characteristics and geometry of some spicules. Within the Microcionidae, C. (C.) arcuophora is also similar to C. (C.) biclathrata in spicule geometry and fibre characteristics, although there are substantial differences between them in skeletal construction. The species should also be compared to C. (C.) striata, which differs mainly in the morphology of its toxa. ^ REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 93 FIG. 28. Clathria (Clathria) arcuophora Whitelegge (paralectotype AMZI 209). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Palmate isochelae. E, Oxeote and oxhorn toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype AMG4346. Clathria (Clathria) biclathrata sp. nov. (Figs 30-31, Table 2) Microciona clathrata Whitelegge, 1907: 491-494, p1.46, fig. 38-38a; Vosmaer, 1935a: 608; Hooper & Wiedenmayer, 1994: 258. Clathria clathrata; Hallmann, 1912: 209. Dictyociona clathrata; de Laubenfels, 1936a: 110. Thalysias clathrata; de Laubenfels, 1953: 527. Not Tenacia clathrata Schmidt, 1870: 56,80. Not Clathria clathrata; Vosmaer, 1880: 153; Ridley & Dendy, 1887: 147; Wilson, 1902: 397-398; Alcolado, 1976: 5. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 94zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 29. Clathria (Clathria) arcuophora Whitelegge (paralectotype AMZ1209). A, Choanosomal skeleton. B, Fibre characteristics (x409). C, Echinating acanthostyle. D, Acanthostyle spination. E, Base of auxiliary subtylostyle. F, Palmate isochelae. G, Oxeote toxas. H, Oxhorn toxas. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 95 TABLE 2. Comparison between present and published records of Clathria(Clathria)biclathrata, sp.nov. All measurements are given in pm, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Choanosomal principal styles Lectotype (AMG4355) Paralectotype (AMG10530) 2534372 . 3)446 x 18(22.8)-33 226(337.5)423 x 17(22.6)-28 Specimen (N=I) 2334319.8)418 x 14(17.8)-24 Subectosomal I 32-(214.0)125-(165.4)1444197.0)auxiliary 293 x 2-(4.9)-7 243 x 3-(5.5)-7 253 x 3-(4.6)-7 styles 734158.4)-231 1144161.1)-193 1134150.2)-193 Echinating acanthostyles x 4.5-(14.7)-I9 x 11-(13.6)-18 x 8-(12.4)-I6 Chelae 7-(10.4)-I4 6-(8.6)-11 4-(7.8)-12 Toxas I 28-(93.0)-149 x 0.8-(2.4)-3.5 Toxas II 4-(10.2)-I8 x 0.841.7)-2.0 434102.8)-141 x I.8-(2.8)-3.5 6-(I8.0)-36 38-(66.6)-104 x 1-(2.7)-4 3-(12.4)-24 MATERIAL. LECTOTYPE: AMG4355: (dry) Off Woolongong, NSW, 34 °25'S, 151 ° 10'E, 110-112m depth, coll. FIV 'Thetis' (dredge). PARALECTOTYPES: AMG10530: (dry) unknown locality, NSW, coll. FIV `Thetis' (dredge). AMG10531 (presently missing): (label `Dictyociona clathrata, cotype'). OTHER MATERIAL: NEW SOUTH WALES - AMZ994. HABITAT DISTRIBUTION. Up to 112m depth, substrate unknown; S. coast of NSW (Fig. 30H). DESCRIPTION. Shape. Clavulate to subspherical, bushy, honeycombed mass, 85-105mm high, 30-55mm wide, up to 40mm maximum thickness, composed of numerous thin, cylindrical, tightly anastomosing branches; small cylindrical stalk, 8-15mm long, up to 8mm diameter. Colour Live colouration unknown, grey-brown to dark brown in dry state. Oscules. Small oscules, 1-2.5mm diameter, on edges of surface microconules. Texture and surface characteristics. Surface very hispid, minutely porous, with numerous slightly elevated microconules; texture unusually tough. Ectosome and subectosome. Ectosomal skeleton a tangential layer of subectosomal auxiliary styles in multispicular tracts, with numerous choanosomal principal spicules protruding and extending a long way through surface; subdermal skeleton, if present, totally obscured by dense mass of erect choanosomal megascleres. Choanosome. Choanosomal skeletal architecture irregularly reticulate, slightly renieroid, with heavy spongin fibres forming tight meshes, and some compression of axial fibres; spongin fibres not clearly divisible into primary or secondary components, but merely ascend and diverge towards surface; fibres mostly aspicular, sometimes with one or few choanosomal principal subtylostyles in core, heavily echinated by both acanthostyles and choanosomal principal subtylostyles (the latter `spicate' in arrangement), and some intermediate spicules with rudimentary spines on shaft; echinating megascleres most abundant on peripheral fibres; mesohyl matrix heavy, with few subectosomal auxiliary subtylostyles between meshes. Megascleres (Table 2). Choanosomal principal subtylostyles slightly curved or straight, fusiform, slightly constricted at base, heavily spined bases (smaller examples may also have scattered spines on shafts). Subectosomal auxiliary subtylostyles short, fusiform, straight, with slightly subtylote microspined bases. Acanthostyles relatively long, thick, slightly curved, with prominent subtylote bases, with evenly distributed large spines over entire spicule or with an aspinose region proximal to base. Microscleres (Table 2). Palmate isochelae, some twisted. Toxas include larger thick wing-shaped forms with large central curvature, slightly reflexed arms and microspined points, and smaller oxhorn forms, the smallest with abbreviated arms. REMARKS. This species is characterised by its compressed skeletal architecture, fibre and ectosomal features, and it is unlikely to be confused with other members of Hallmann's (1912) spicata group of microcionid species (cf. Hooper et al., 1990) which have principal spicules protruding through fibres and surface skeletons but few within fibres themselves. A feature overlooked by previous authors is the presence of spinous extremities on toxas, which are virtually identical to those of type species of Clathria, C. (C.) compressa, also occurring in C. (C.) juncea, C. (C.) lobata, and Artemisina suberitoides. The geometry of spicules in C. (C.) biclathrata is similar to those in C. (C.) arcuophora, although these species differ quite substantially in their skeletal architecture. Clathria (Clathria) caelata Hallmann, 1912 (Figs 32-33, Table 3) Clathria caelata Hallmann, 1912: 139, 177, 206, 211- 216, p1.33, fig.4, text-fig.43; Hooper & Wiedenmayer, 1994: 258. Clathria coelata; Burton & Rao, 1932: 336 [lapsus]. Pseudanchinoe caelata; de Laubenfels, 1936a: 109. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 96zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 30. Clathria (Clathria) biclathrata sp.nov. (lectotype AMG4355). A, Choanosomal principal subtylostyle and spined subtylostyle. B, Echinating acanthostyles. C, Subectosomal auxiliary subtylostyles. D, Palmate isochelae. E, Wing-shaped toxas. F, Oxhorn toxas. G, Section through peripheral skeleton. H, Australian distribution. I, Lectotype AMG4355. Not Clathria inanchorata Ridley & Dendy, 1886:475; Ridley & Dendy, 1887: 150, p1.28, fig.4, p1.29, figs 13,13a. cf. Microciona prolifera; Vosmaer, 1935a: 611, 648, 664. MATERIAL. LECTOTYPE: AMZ778: (wet) 64km W. of Kingston, SA, 36 °50'S, 139 °05'E, 60m depth, coll. FIV 'Endeavour' (dredge; label `4th consignment'). PARALECTOTYPES: AME53: (dry) same locality. AMZ952-953: unspecified locality, W. coast Tas. ('ref. G255'). OTHER MATERIAL: TASAME2273. HABITAT DISTRIBUTION. Depth 53-106m; substrate unknown; Kingston (SA), Bass Strait (Vic), Cape Barren, W coast (Tas) (Hallmann, 1912). DESCRIPTION. Shape. Planar or multiplanar, digitate fans, 110-150mm high, 70-90mm wide, with short cylindrical stalk, 8-17mm long, 5- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 97 FIG. 31. Clathria (Clathria) biclathrata sp.nov. (lectotype AMG4355). A, Choanosomal skeleton. B, Ectosomal skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spination. F, Wing-shaped toxa. G, Oxhorn toxas. H, Palmate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM 98^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 32. Clathria (Clathria) caelata Hallmann (paralectotype AME53). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Intermediate echinating and principal style. D, Echinating acanthostyle. E, Accolada toxa. F, Oxhorn toxas. G, Section through peripheral skeleton. H, Australian distribution. I, Paralectotype AMZ953. J, Paralectotype AMZ952. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 99 FIG. 33. Clathria (Clathria) caelata Hallmann (A-B, Lectotype AMZ778; C-G, E2273). A, Choanosomal skeleton. B, Fibre characteristics. C, Ectosomal skeleton. D, Accolada toxa. E, Oxhorn toxas, F, Echinating acanthostyles. G, Acanthostyle spination. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 100zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA lOmm diameter, rounded lobate, or digitate margins. Colour Llight to dark brown preserved. Oscules. Small pores on margins of lobes, up to 1.5mm diameter. Texture and surface characteristics. Surface highly rugose, with numerous close-set microconules, subdermal canals and grooves, between which extends a skin-like dermal membrane; texture firm, compressible. Ectosome and subectosome. Ectosome microscopically hispid, with plumose brushes of choanosomal principal styles protruding through surface, and with thin layer of subectosomal auxiliary styles tangential to surface; subdermal region not markedly differentiated from choansome containing bundles of diverging principal styles embedded in peripheral fibres; peripheral fibres heavily echinated on their exterior surfaces; acanthostyles may extend into ectosomal skeleton. Choanosome. Choanosomal skeletal architecture irregularly reticulate, with well developed spongin fibres forming ovoid to elongate meshes; fibre anastomoses relatively cavernous in axis, although smaller in peripheral skeleton; fibres clearly differentiated into primary ascending and secondary transverse components, although fibre diameter is consistent throughout skeleton: primary fibres contain pauci- or multispicular tracts of choanosomal principal subtylostyles, forming a radial architecture; secondary fibres uni- or aspicular; spongin fibres echinated on external surfaces only, with a variable density of echinating acanthostyles, mostly relatively light except at the periphery; choanosomal styks also echinate fibres, particularly at fibre nodes; mesohyl matrix moderately heavy, granular, pigmented; extra-fibre megascleres (subectosomal subtylostyles) usually abundant. Megascleres (Table 3). Choanosomal principal subtylostyles thick, straight or slightly curved, fusiform, with slightly subtylote smooth bases, although some examples are obvious intermediates to echinating acanthostyles, bearing rudimentary spines on the shaft. Subectosomal auxiliary subtylostyles straight or slightly curved, fusiform, relatively thin, evenly rounded or slightly subtylote bases, and smooth or microspined bases. Echinating acanthostyles variable in length, subtylote, with evenly spined shafts on smaller forms, or with aspinose necks on larger forms. Microscleres (refer to Table 3 for dimensions). Isochelae absent. TABLE 3. Comparison between present and published records of Clathria (Clathria) caelata (Hallmann). All measurements are given in m, denoted as range (and mean) of spicule length x spicule width (N=25) Paralectotype (AME53) Specimen (N=1) Choanosomal 1484215.4)-276 principal x 8-(12.2)-16 styles 1544222.2)-262 x 9-(11.8)-14 186-(254.5)-353 x 6-(9.4)-10 Subectosomal 128-(216.4)-294 auxiliary x 2-(3.2)-5 styles 45-(71.9)-122 Echinating x 245.1)-8 acanthostyles 1344234.8)-324 x 3-(4.6)-7 1664267.0)-355 x 2-(3.5)-7 64-(89.4)-131 x 4-(6.4)-8 23-(64.4)-120 x 2-(4.8)-8 SPICULE Lectotype (AMZ778) Chelae absent absent absent Toxas I 1224151.7)-190 x 0.5-(1.1)-1.5 72-(101.8)-165 x 0.8-(1.4)-2 86-(121.1)-165 x 0.5-(1.4)-2 Toxas II 2I-(55.9)-83 x 1-(2.3)-4 13-(38.8)-75 24-(48.2)-92 x 1-(2.1)-3.5^, x 1-(2.2)-4 Toxas separated into two morphs: I - long, thin accolada toxas, with straight points and slight central curvature; II - relatively thick oxhorn toxas, ranging from almost straight with only slight and angular central curvature, to widely curved at the centre with reflexed points. REMARKS. Not all of the specimens described by Hallmann (1912) belong to this species: AME771, E772 and E773 are species belonging to Axinellidae - Reniochalina (2 specimens) and Acanthella, respectively. All three specimens look very similar in external morphology to C. (C.) caelata, but differences are immediately obvious upon examination of the skeleton. Among the few known specimens of this species there is relatively high variability in choanosomal skeletal construction although this is difficult to define concisely. The development of spongin fibres, the degree to which the skeletal meshes are compressed or elongated, and the density and pigmentation of the mesohyl matrix may vary between specimens. Growth form and spicule geometry appear to be closely comparable between all specimens, but some (e.g., AME2273) have predominantly long thin toxas, whereas others (e.g., AMZ952) have mostly short and thick toxas. All specimens have both choanosomal styles and acanthostyles echinating fibres, and in this respect the species belongs to Hallmann's (1912) spicata' group, together with Clathria (C.) biclathrata, C. (C.) inanchorata, C. (Thalysias) costifera, C. (T) coppingeri, C. (T) lendenfeldi and others (Hooper et al., 1990). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 101 Clathria (Clathria) calopora Whitelegge, 1907 Clathria calopora Whitelegge, 1907: 499-500, p1.46, fig.34; Hallmann, 1912: 211; Hooper & Wiedenmayer, 1994: 259. Thalysias calopora; de Laubenfels, 1936a: 105. MATERIAL. HOLOTYPE: AM (presently missing): Shoalhaven Bight, NSW, 34 ° 49'S, 151°04'E, 30-90m depth, 1.vii.1911, coll. FIV 'Endeavour' (trawl). HABITAT DISTRIBUTION. Depth 27-32m, substrate unknown; S. coast of NSW. DESCRIPTION. Shape. Branching, 195mm high, with thin, bifurcate, cylindrical, mostly non-anastomosing digits, 3-8mm diameter, with tapering points, short cylindrical stalk. Colour Live colouration unknown, greyish-yellow preserved. Oscules. Small oscules, about lifun diameter, scattered over surface. Texture and surface characteristics. Surface uneven, minutely hispid, honeycomb-like, with prominent bulbous conulose processes scattered over surface; texture hard, incompressible in dry state. Ectosome and subectosome. Ectosome microscopically rugose, close-meshed reticulation of peripheral spongin fibres covered by a thin membrane; ectosomal skeleton with a thin tangential layer of subectosomal auxiliary subtylostyles, also forming plumose tufts on surface, through which choanosomal principal styles from peripheral fibres protrude, individually or in plumose brushes; subdermal auxiliary megascleres run parallel with peripheral fibres, but not forming organised extra-fibre tracts. Choanosome. Choanosomal skeleton irregularly reticulate, with some axial and extra-axial differentiation; spongin fibres relatively heavy, clearly divided into primary ascending and secondary transverse components; primary fibres sinuous, forming radial architecture, with a paucispicular core of choanosomal principal styles; primary fibres more compressed at axis than at periphery; secondary fibres less common, with uni- or paucispicular core of megascleres; branching of spongin fibres produces elongate meshes in axis and round or rectangular meshes in periphery; spongin fibres very heavily echinated, particularly in peripheral regions; numerous sinuous toxodragmata dispersed in mesohyl between fibres. Megascleres. Choanosomal principal styles thick, slightly curved, fusiform, with rounded smooth bases. Length 300-500p,m, width 1522m. Subectosomal auxiliary subtylostyles thin, straight or slightly curved, with smooth slightly subtylote bases. Length 150-200p,m, width 24p,m. Acanthostyles short, stout, evenly spined, spines large. Length 30-80p,m, width up to 12p.m. Microscleres. Palmate isochelae. Length 810t.m. Toxa morphology unknown, apparently long, slender, with large central curvature. Length ? 200-300p,m, width ? up to 21.1,m. REMARKS. This species is known only from Whitelegge's (1907) poor description. It is not possible to determine whether dimensions of echinating acanthostyle (cited as 0.3-0.8mm long by Whitelegge) is merely a typographical error. Whitelegge gave no indication of whether C. (C.) calopora has a special ectosomal skeleton, but described the species as having a tangential or paratangential layer of subdermal (auxiliary) megascleres, through which protrude choanosomal (principal) styles. Consequently, there was no justification for de Laubenfels (1936a) referring the species to Thalysias. Clathria (Clathria) chelifera (Hentschel, 1911) (Figs 34-35, Table 4) Spanioplon cheliferum Hentschel, 1911: 362-363, fig.42; Hentschel, 1912: 368-369. Allocia chehfera; Hallmann, 1920: 768; Bergquist & Fromont, 1988: 96, fig. 8c, pis 45e-f, 46a-c, table 73; Dawson, 1993: 44 (note). Clathria chelifera; Dendy, 1922: 70-71, p1.14, fig. 3a-e; Hooper & Wiedenmayer, 1994: 259. Not Microciona chelifera Lëvi,1960a: 70, fig. 12. MATERIAL. HOLOTYPE: ZMH (not seen) (fragment ZMB4440): precise locality unknown, Perth region, WA, 1905, coll. W. Michaelsen & R. Hartmeyer (dredge). PARATYPE: SMF 1571 (fragment MNHNDCL2327): same locality. OTHER MATERIAL: VIETNAM - PIBOC-05-216 (fragment QM G300058). HABITAT DISTRIBUTION. 10-100m depth; rock or gravel substrates; Arafura Sea (NT) and Perth region (S. WA) (Fig. 34F). Also Indian Ocean (Amirante), New Zealand (Three Kings Is) and South China Sea (Hon Trung Lon, Vinh Loi coast, S. Vietnam). DESCRIPTION. Shape. Arborescent, foliose, planar growth form, up to 50mm high made up of fused porous-reticulate lamellae 10-15mm thick. MEMOIRS OF THE QUEENSLAND MUSEUM 102^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 34. Clathria (Clathria) chelifera (Hentschel) (paratype SMF1571). A, Choanosomal principal style. B, Subectosomal auxiliary styles (quasitylotes). C, Echinating acanthostyles. D, Palmate isochelae. E, Section through peripheral skeleton (ectosome on far right). F, Australian distribution. G, QG300058. ^ 103 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU FIG. 35. Clathria (Clathria) chelifera (Hentschel) (QMG300058). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Palmate isochelae. G, Pattern of echinating spicules. H, Section of peripheral skeleton. MEMOIRS OF THE QUEENSLAND MUSEUM 104^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA TABLE 4. Comparison between present and published records of Clathria (Clathria)chelifera (Hentschel). All measurements are given in 1.1.m, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Paratype (SMF1571) (Perth, WA) Specimen Bergquist & Fromont (1988) (Three Kings Is, NZ) Specimen (N=1) (S. Vietnam) Choanosomal principal styles Subectosomal auxiliary styles 1404150.1)-161 x 4-(6.1)-7 1734193.4)-205 x 2-(3.9)-5 2704404)-550 x 15-(18)-20 280-(334)-375 x 4-(5)-6.5 150-(164.6)-174 x 4-(6.4)-10 1924204.0)-212 x 3-(4.7)-7 Echinating acanthostyles Chelae I 84-(93.2)-100 x 7-(7.8)-9 162-(207)-290 x 12.5-(14)-19 10341165)-132 x 5-(8.0)-10 13-(14.6)-1 20421)-23 15-(17.6)-20 Chelae H 7-(8.7)-10 absent 9-(10.3)-12 Colour Red alive (10R 5/8), beige or yellow brown in ethanol. Oscules. Not seen. Texture and stuface characteristics. Surface hispid, uneven, irregularly conulose, with transparent ectosomal membrane stretched between adjacent conules; texture firm, compressible. Ectosome and subectosome. Ectosomal skeleton with a tangential layer of auxiliary tylotes and numerous isochelae scattered between, supporting the membraneous ectosomal covering; subectosomal region with paratangential tracts of auxiliary tylotes supporting tangential ectosomal layer, the latter sometimes protruding through surface, with ascending primary tracts of choanosomal principal styles in turn supporting these. Choanosome. Choanosomal skeleton reticulate, with multispicular ascending primary fibres and paucispicular transverse connecting fibres; spongin fibres heavy, cored by principal styles and auxiliary tylotes (the latter also scattered throughout the mesohyl), and echinated by acanthostyles more-or-less perpendicular to fibres; mesohyl matrix light, with numerous isochelae scattered between fibres. Megascleres (Table 4). Principal styles slender, slightly curved near basal end, with abrupt hastate points, and completely smooth. Subectosomal auxiliary spicules tylotes or quasitylotes, asymmetrical (and therefore probably modified styles), usually with microspined bases and points or sometimes completely smooth at both ends. Echinating acanthostyles with spinose shaft, base and point but apinose 'neck'; spines large, recurved. Microscleres (Table 4). Palmate isochelae of two sizes, both with thickened and elongate alae. Toxas are absent. REMARKS. Dimensions of some spicules were found to differ in type material (Table 4) from those published by Hentschel (1911). Similarly, two size classes of isochelae were found in the WA population, not one as described by Hentschel (1911). In both respects this population is the same as the one described from Amirante (Dendy, 1922) and the material described above from Vietnam, whereas the specimen described by Bergquist & Fromont (1988) from New Zealand has substantially larger spicule dimensions than either of the Indian Ocean populations, including only one size class of isochela. In spicule geometry, choanosomal skeletal structure and ectosomal characteristics (including the distribution of isochelae in the ectosomal membrane), these 4 disjunct populations are relatively homogeneous and I follow Bergquist & Fromont (1988) in recognising only a single species. No intermediate populations of C. (C.) chelifera are known, and the species is relatively rare with only few known specimens in the Indowest Pacific. This species is unusual to most C. (Clathria) in possessing modified auxiliary spicules with spines on both ends, considered by some authors to be true tylotes typical of the Myxillidae (Hall mann, 1920; Bergquist & Fromont, 1988) or Iophonidae (Hajdu et al., 1994). But these spicules are clearly asymmetrical (quasitylotes), not true diactinal megascleres, and Dendy (1922) correctly assigned this species (and thus the genus Allocia) to Clathria. These modified quasidiactinal auxiliary spicules are infrequent but known in several other microcionids (e.g., C. (C.) bulbosa, C. (Thalysias) major, C. (Dendrocia) pyramida, C. (Wilsonella) australiensis, most Echinochalina, some Holopsamma, and some Echinoclathria species). Clathria (Clathria) conectens (Hallmann, 1912) (Figs 36-37, Plate 1B, Table 5) Wilsonella conectens Hallmann, 1912: 245-247, p1.32, fig.2, text-fig.50. Clathria conectens; Hallmann, 1920: 768; Hooper & Wiedenmayer, 1994: 259. MATERIAL. LECTOTYPE: AMZ220: (dry) 16km E. of Fraser I., Qld, 25°22'S, 153°07'E, 48-52m depth, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 105 coll. FIV 'Endeavour' (dredge). PARALECTOTYPE: AME1533 (dry; presently missing): same locality. OTHER MATERIAL: QUEENSLAND - QMG300455, QMGL714 (fragment NTMZ1537), QMGL2757 (fragment NTMZ1564), QMGL2770 (fragment NTMZ1581), QMG301037, QMG303190, QMG303217, QMG304980, QMG304985, QMG305135, QMG304005, QMG304016. NEW SOUTH WALES - QMG301387. HABITAT DISTRIBUTION. Acropora coral reef, fringing rock reef, boulders, algal turf, wharf pylons; inshore waters, 4-80m depth; Green I., S. Direction I., Innisfail (FNQ), Fraser I., Mudjimba I., Moreton I., N. Stradbroke I., Moreton Bay (SEQ), and Byron Bay (N. NSW) (16-28S) (Fig. 36G). TABLES. Comparison between present and published records of Clathria (Clathria) conectens (Hallmann). All measurements are given in p,m, denoted as range (and mean) of sp'cule length x spicule width (N=25). SPICULE Lectotype (AMZ220) Specimens (N=7) Choanosomal principal styles 944168.4)-268 x 2.5-(4.1)-5 1104171.4)-218 x 3-(4.3)-6 Subectosomal auxiliary styles 92-(171.3)-219 x 1.542.9)-4 174-(230.6)-295 x 1.5-(2.4)-3.5 48-(58.3)-65 x 3-(3.7)-5 36-(64.2)-78 x 2-(3.9)-5 Echinating acanthostyles Chelae Toxas 6-(7.2)-8 4.546.7)-9 22-(69.8)-111 51-(102.6)-164 x 0.5-(0.9)-1.5 x 0.5-(0.7)-1.0 DESCRIPTION. Shape. Massive, subcylindrical mass 40-65mm high, 85-120mm broad, composed of irregularly reticulate, lamellate bulbous branches, up to 15mm diameter, standing erect on substrate. Colour Live colouration bright orange-yellow to bright orange-red (Munsell 2.5YR 7/10-10R 6/10), brown in ethanol. Oscules. Small oscules, up to 1.5mm diameter, on edges and tips surface bulbs. Texture and surface characteristics. Surface uneven, porous, optically smooth. Ectosome and subectosome. Ectosome with tangential or paratangential layer of thin subectosomal auxiliary subtylostyles, on or just below a membraneous dermal layer; subectosomal auxiliary styles confined entirely to peripheral skeleton; subectosomal skeleton virtually nonexistent, with only few erect, plumose choanosomal principal styles, arising from peripheral choanosomal spongin fibres, projecting into tangential ectosomal layer. Choanosome. Choanosomal skeleton irregularly reticulate, with regular circular, oval or elongate meshes enclosing small oval choanocyte chambers; spongin fibres relatively heavy, without size differentiation of primary or secondary components, although ascending (primary) skeletal fibres cored by paucispicular or multispicular tracts of choanosomal styles, whereas connecting, transverse (secondary) fibres uni-, pauci- or entirely aspicular; echinating acanthostyles sparse, scattered evenly throughout skeleton, occasionally incorporated into fibres; mesohyl matrix very light, some choanosomal styles scattered between fibres; some specimens also incorporating detritus into mesohyl, but not into fibres. Megascleres (Table 5). Choanosomal principal styles thin, fusiform, occasionally styloid, slight- ly curved, with rounded or very slightly subtylote, smooth bases. Subectosomal auxiliary subtylostyles slightly curved, sometimes sinuous, exceedingly thin, hastate, almost vestigial, with very slightly subtylote, smooth bases. Acanthostyles subtylote, with more-or-less evenly distributed vestigial (granular) spination. Microscleres (Table 5). Palmate isochelae small, unmodified. Toxas accolada rare, thin, with sharply angular central curvature and straight arms. REMARKS. Specimen AMZ220 is designated lectotype (labelled `cotype of Wilsonella conectens, duplicate of E1533') as the latter specimen is presently missing from AM collections. Despite Hallmann's (1912) remarks to the contrary C. (C.) conectens is clearly different from C. (C.) angulifera (see above), although both species do fit into his (erroneous) concept of Wilsonella. Moreover, in C. (C.) conectens the megascleres which core fibres (choanosomal styles) are differentiated from those occurring in the dermal skeleton (subectosomal styles), and although their geometry is very similar, they have very different morphology (see Fig. 36) and thickness (see Table 5). Each category of spicule is localised in the choanosomal and ectosomal regions, respectively, and they are not intermingled as supposed by Hallmann (1912). The entire spiculation of this species is reduced, and for that reason it is easily recognisable. This species is a common member of the SolanderianPeronian biogeographical overlap zone centred around Moreton Bay, Queensland. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 106zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A FIG. 36. Clathria (Clathria) conectens (Hallmann) (lectotype AMZ220). A, Choanosomal principal styles. B, Subectosomal auxiliary style. C, Echinating acanthostyle. D, Accolada toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. Clathria (Clathria) crassa (Lendenfeld, 1887) (Figs 38-39) Antherochalina crassa Lendenfeld, 1887b: 787, p1.22, fig.41. Clathria crassa; Burton, 1934a: 558; Hooper & Wiedenmayer, 1994: 259. Microciona or Thalysias crassa; de Laubenfels, 1936a: 112. Not Reniera crassa Carter, 1876: 312. Not Aulena crassa; Lendenfeld, 1889a: 101. MATERIAL. HOLOTYPE: BMNH1886.8.27.450 (fragments AMG3460, AMZ1991): Port Jackson, NSW, 33 ° 51'S, 151 ° 16'E, other details unknown. HABITAT DISTRIBUTION. Ecology unknown; central coast NSW. DESCRIPTION. Shape. Thin fan, 230mm high, 190mm maximum width, up to 8mm thick, with short stalk, 25mm long, digitate margins. Colour Live colouration unknown, grey-brown in ethanol. Oscules. Small pores seen on both faces of fan, 1-2mm diameter, with subdermal drainage canals surrounding oscules. Texture and surface characteristics. Surface smooth, not optically hispid, with stellate sculpturing on both faces of fan (associated with aquiferous system); texture firm, flexible. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU 107 FIG. 37. Clathria (Clathria) conectens (Hallmann) (A-B, lectotype AMZ220; C-G, QMG303217). A, Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal fibres. D, Echinating acanthostyle. E, Acanthostyle spines. F, Accolada toxa. G, Palmate isochelae. 108zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 0 0 FIG. 38. Clathria (Clathria) crassa (Lendenfeld) (fragment of holotype AMZ I 991). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Oxhorn toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, holotype BMNH 1886.8.27.450. ^ REVISION OF M1CROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 109 FIG. 39. Clathria (Clathria) crassa (Lendenfeld) (holotype BMNH1886.8.27.450). A, Fibre characteristics. B, Ectosomal region. C, Choanosomal skeleton. D, Oxhorn toxas. E, Smaller toxas intermediate between oxhorn and u-shaped. F, Palmate isochelae. G, Acanthostyle spines. H-1, Echinating acanthostyles. 110zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Ectosome and subectosome. Ectosome microscopically hispid, with points of large choanosomal principal styles from peripheral fibres protruding through surface in plumose brushes; thin tangential layer of subectosomal auxiliary subtylostyles lying on or just below surface, at base of protruding choanosomal megascleres. Choanosome. Choanosomal skeleton irregularly reticulate, with slightly renieroid axis and plumoreticulate extra-axis; spongin fibres in axis flattened, very heavy, running longitudinally through lamellae; axial fibres clearly divisible into primary (longitudinal) and secondary (ascending, connecting) components; primary fibres cored by paucispicular tracts of principal choanosomal styles; secondary fibres unispicular; extra-axial skeleton diverges from axis at an oblique angle, with moderately heavy fibres, divided into primary (multispicular, ascending) and secondary (unispicular, transverse) elements; choanosomal principal styles project from primary fibres in plumose tracts; secondary fibres connect ascending primary lines, producing reneiroid reticulation, except at periphery where architecture is distinctly plumose; echinating acanthostyles moderately common, evenly distributed throughout skeleton; mesohyl matrix abundant, containing few tnicroscleres but few loose megascleres. Megascleres. Choanosomal principal styles thick, fusiform, slightly curved, with rounded or slightly subtylote, smooth bases. Length 184(292.3)-463p,m, width 9-(17.3)-22p,m. Subectosomal auxiliary subtylostyles thick, straight, fusiform, with microspined subtylote bases. Length 118-(226.7)-316p,m, width 3(4.8)-6 pun. Acanthostyles short, thick, subtylote or rounded, with evenly dispersed vestigial (granular) spination. Length 51-(66.3)-82Rm, width 5-(6.5)-9p.m. Microscleres. Palmate isochelae large, unmodified. Length 17-(l9.5)-23p,m. Toxas oxhorn, thick, with rounded central curvature, slightly reflexed points although the smaller ones may lack reflexed points and are intermediate between oxhorn and u-shaped forms. Length 28-(68.0)-112p,m, width 0.842.6)4.511M. REMARKS. Burton (1934a) designated A. crassa type species of Antherochalina, and subsequently declared that the genus was a synonym of Clathria. Lendenfeld's (1887b) brief descrip- tion of A. crassa is vague and not very useful in distinguishing it from other Clathria, but type material is still extant and recognisable. However, there is little agreement between characters in the type material and as described by Lendenfeld, C. (C.) crassa is very closely related to C. (C.) arcuophora, with similar skeletal architecture (with 2 components, renieroid and plumoreticulate), spicule geometry, spicule sizes, similar fibre characteristics and comparable growth forms. It is possible that the two species are synonyms, but their formal merger is not warranted on the basis of the existing relatively poor material. Similarly C. (C.) crassa shows some similarities with C. (Isociella), particularly to C. (I.) eccentrica. This resemblance is mostly due to the renieroid axial skeletal architecture and geometry of both principal and auxiliary styles. Clathria (Clathria) decumbens Ridley, 1884 (Figs 40-41) Cluihria decumbens Ridley, 1884a: 612, p1.53, fig.k, p1.54, fig.g-g'; Ridley & Dendy, 1887: 148; Burton, 1938a: 29, p1.3, fig.23; Hooper & Wiedenmayer, 1994: 259. Wilsonella decumbens; Hallmann, 1912: 239. MATERIAL. HOLOTYPE: BMNH1882.10.17.51: Boudouse and Etoile Is, Amirante Is Group, Indian Ocean, 6 ° S, 53°10'E, coll. HMS 'Alert' (dredge). PARATYPES: BMNH1882.10.17.71, 1882.10.17.76: same locality. OTHER MATERIAL: QUEENSLAND BMNH1887.5.2.139. HABITAT DISTRIBUTION. 6-26m depth; on sand and coral rubble substrate; Cape York, Torres Strait (FNQ) (Fig. 40F) (Ridley, 1884a). W. Indian Ocean (Ridley & Dendy, 1887), Madras (Burton, 1938a). DESCRIPTION. Shape. Small, subcylindrical, irregularly lobate, 32-50mm long, 15-40mm wide, up to 25mm thick. Colour Brown to red-brown in ethanol. Oscules. Numerous small oscules, 1-2mm diameter, dispersed between surface conules. Texture and surface characteristics. Surface rugose, irregularly conulose, with canals, grooves and ridges meandering over the surface. Ectosome and subectosome. Ectosome membraneous between ridges and surface projections, with spongin fibres from choanosome producing a dermal reticulation; dermal region lacks a mineral skeleton entirely; spongin fibres in subectosomal region closely reticulate, with relatively small mesh sizes. Choanosome. Choanosomal skeletal architecture regularly to irregularly reticulate; spongin fibres REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 111 moderately heavy, undifferentiated into primary or secondary lines, lightly cored by paucispicular tracts of both choanosomal principal styles and subectosomal auxiliary styles; heavily echinated by acanthostyles; coring spicules in peripheral fibres ascend to surface, piercing surface processes in light brushes; subectosomal auxiliary subtylostyles also sparsely scattered within mesohyl; fibre meshes in choanosomal skeleton cavernous; mesohyl matrix abundant, lightly pigmented. Megascleres. Choanosomal principal styles straight or slightly curved near base, hastate points, with evenly rounded bases, smooth or with microspines on both points and bases. Length 128-(156)-176Rm, width 4-(4.9)-6p.m. Subectosomal auxiliary subtylostyles straight, with hastate points, slightly subtylote bases, bases smooth or occasionally microspined. Length 159-(1177.8)-193Rm, width 243.7)6 Km. Acanthostyles subtylote, with strongly formed, recurved spines over apical end, shaft and base but bare neck. Length 62-(83.2)-104Rm, width Microscleres. Palmate isochelae unmodified, two discrete size classes, the smaller showing variation in fusion of alae. I: Length 14-(20.4)-28Km, II: length 5-(7.4)-10Km. Toxas absent. REMARKS. Hallmann (1912) referred this species to Wilsonella, presumably because Ridley (1884a) did not differentiate between choanosomal (principal) and subectosomal (auxiliary) megascleres. Subsequently, Burton (1938a) described additional specimens from Madras which had differentiated principal and auxiliary spicules (but were otherwise identical with Ridley's (1884) description). In type material there were consistent morphological differences between principal and auxiliary spicules, confirming that the species should be retained in Clathria (Clathria). Although C. (C.) decumbens has been described from three widely separated localities, it remains poorly known, being only poorly differentiated from other low growing, lobate Clathria species. The species is similar to C. (Thalysias) major (with spines on both the bases and points of some of its megascleres), while at the same time being similar to species in the juniperind species complex (i.e., Clathria in which the geometry of choanosomal principal and subectosomal auxiliary spicules is barely different), but the species has little else of distinction. Clathria (Clathria) echinonematissima (Carter, 1881) Wilsonella echinonematissima Carter, 1881a: 366; Carter, 1887: 210; Hallmann, 1912: 243. Clathria echinonematissima; Dendy, 1896: 33, 34; Hooper & Wiedenmayer, 1994: 259. MATERIAL. HOLOTYPE: BMNH not found (slide containing only a desilicified section is the only type material known to exist): Westernport Bay, 38 °26'S, 145°08'E, or Port Phillip, Vic, 38 °09'S, 144°52'E, coll. J.B. Wilson (dredge). HABITAT DISTRIBUTION. Ecology unknown; Victoria. DESCRIPTION. Shape. Massive. Colour. Unknown. Oscules. Unknown. Texture and surface characteristics. Unknown. Ectosome and subectosome. Ectosomal skeletal tracts heavily cored with detritus, megascleres excluded. Choanosome. Choanosomal skeleton irregularly reticulate, with relatively heavy spongin fibres; fibres of peripheral skeleton are solely arenaceous, whereas within choanosome fibres cored by subectosomal auxiliary styles; echinating acanthostyles dispersed throughout skeleton. Megascleres. Choanosomal principal megascleres apparently absent. Subectosomal auxiliary subtylostyles with smooth bases. Length 21011m, width 4t.t.m. Acanthostyles apparently divided into two size categories. Length from 145Km, width 8.311m. Microscleres. Isochelae arcuate. Length 251.1,m. Toxas absent. REMARKS. This species is barely recognisable as Clathria from Carter's (1881) description, and it is only poorly differentiated from other microcionid species. One category of auxiliary spicule and the absence of choanosomal principal spicules places it in Wilsonella (sensu Hallmann, 1912). It also was described with arcuate isochelae, similar to species grouped by Hallmann (1920) in Paradoryx, but this is uncorroborated. Clathria echinonematissima is a species inquirendum as it is only known from a slide preparation, now desilicified, allegedly made from the missing holotype. Clathria (Clathria) hispidula (Ridley, 1884) (Figs 42-43) Amphdectus hispidulus Ridley, 1884a: 429-430, p1.40, fig.c, p1.41, fig.y. 112zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 40. Clathria (Clathria) decumbens Ridley (paratype BMNH1882.10.17.71). A, Choanosomal principal styles. B, Subectosomal auxiliary styles/subtylostyles. C, Echinating acanthostyles. D, Palmate isochelae. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype BMNH1882.10.17.51. H, Paratype. Axociella hispidulus; de Laubenfels, 1936a: 114. Esperiopsis hispidula var. ramosa; Hentschel, 1911: 313. Not Hymeraphia hispidula Topsent, 1904a: 164-165, p1.14, fig.2. MATERIAL. LECTOTYPE: BMNHI881.10.21.261: Thursday I., Torres Strait, N. Old, 10 °35'S, 142 ° 13'E, 6-10m depth. 01.vi.1881, coll. HMS 'Alert' (dredge). PARALECTOTYPE: BMNH1881.10.21.319: same locality (dry). OTHER MATERIAL: WESTERN AUSTRALIA - ZMB4408. HABITAT DISTRIBUTION. Encrusting on bivalves, hydroids, and gorgonians and algae; 6-11rn depth; Torres Strait (FNQ), and Shark Bay (WA) (Fig. 42G). DESCRIPTION. Shape. Erect, irregular branch- ing reticulate mass of clathrous digits, up to 60mm long, 55mm diameter. Colour. Live colour unknown, light brown in ethanol. Oscules. Small oscules, <2mm diameter in preserved material, scattered over lateral margin. Texture and surface characteristics. Harsh, compressible, slightly elastic; surface with meandering irregular ridges and microconules scattered over branches, and tips of fibres from primary skeleton protruding. Ectosome and subectosome. Sparse paratangential skeleton of subectosomal auxiliary subtylos- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 113 FIG. 41. Clathria (Clathria) decumbens Ridley (paratype BMNH1882.10.17.71). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E-F, Terminations of principal styles. G, Principal styles. H-K, Palmate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM 114^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 42. Clathria (Clathria) hispidula (Ridley) (lectotype BMNH1881.10.21.261). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. tyles forming bundles or lying more-or-less erect on the surface, particularly on ends of surface conules; tips of conules with choanosomal principal styles also protruding only short distance through surface; choanosomal fibres immediately below surface skeleton with poorly developed subectosomal region. Choanosome. Skeleton regularly reticulate, slightly renieroid, with thin but well developed spongin fibres forming oval or rectangular, relatively wide meshes, 150-350Rm diameter, generally more cavernous in axis than in peripheral region; spongin fibres 20-701J,m diameter, imperfectly divided into primary, ascending, multispicular tracts of 4-10 spicules per tract, interconnected by uni-, pauci- or aspicular secondary transverse tracts; fibres cored by choanosomal principal styles not occupying en- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 115 Clat hria ( Clat hria) hispidula (Ridley) (lectotype BMNH1881.10.21.261). A, Choanosomal skeleton. FIG. 43. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spination. E, Terminations of auxiliary spicules. F, Palmate isochelae. G, Wing-shaped toxa. MEMOIRS OF THE QUEENSLAND MUSEUM 116zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ tire fibre diameter, and sparsely echinated by slender acanthostyles; mesohyl matrix light containing scattered microscleres and some auxiliary megascleres. Megascleres. Choanosomal principal styles slender, straight or slightly curved midway along shaft, with evenly rounded smooth bases and fusiform points. Length 884162.7)-21911m, width 4-(4.8)-611m. Subectosomal auxiliary subtylostyles similar in geometry to principal spicules, but more slender and with subtylote bases; thinner (younger) principal spicules frequently sinuous. Length 94(143.0)-175pAn, width 2-(3.4)-5p.m. Echinating acanthostyles club-shaped, slender, slightly subtylote, evenly spines or with bare 'neck' below base, fusiform or rounded points, granular spination. Length 52-(59.4)-65Rm, width 2-(4.1)-5p.m. Microscleres. Palmate isochelae very abundant, small, unmodified, with lateral alae completely fused to shaft and front ala entire; lateral and front alae of approximately equal length. Length 9(11.6)-13p,m. Toxas uncommon, small, thick, wing-shaped, with rounded central curve and slightly reflexed arms. Length 17-(56.8)-104Rm, width 1.5-(2.1)3p.m. REMARKS. This species has not yet been rediscovered from either of the known locations of early collections, and it remains relatively poorly known from museum specimens. Surprisingly, both Ridley (1884a) and Hentschel (1911) failed to describe several spicule types present in their respective material, particularly echinating acanthostyles which are uncommon but certainly present, as well as auxiliary subtylostyles and less common toxa microscleres. Furthermore, Hentschel's material essentially differs from Ridley's only in the specific dimensions of spicules and growth form (being more elongate, branching), and it is not considered to be necessary to recognise the subspecific taxon proposed by Hentschel (1911) for the WA population. Although the identity of this species has never been been clearly established from either published record, it is obviously a Clathria with relatively cavernous skeletal architecture and standard spiculation. It is similar to C. (C.) angulifera (Dendy) from Victoria and southern Queensland, and C. (T) aphylla from the Houtman Abrolhos, in having a cavernous, slightly renieroid skeletal structure ('angulifera' species group), differing in the protruding fibrous ectosomal skeleton, spicule geometries (e.g., toxas, acanthostyles), a more-or-less branching growth form (cf. lobate lamellate and foliose lamellate, respectively), and spicule sizes. Clathria (Clathria) inanchorata Ridley & Dendy, 1886 (Figs 44-45, Table 6) Clathria inanchorata Ridley & Dendy, 1886: 475; Ridley & Dendy, 1887: 150, p1.28, fig.4, p1.29, fig. 13; cf. Kieschnick, 1896: 533; cf. Thiele, 1903a: 959; Whitelegge, 1907: 492-495; Hallmann, 1912: 206, 211, 214, 215; Hooper & Wiedenmayer, 1994: 259. Pseudanchinoe inanchorata; de Laubenfels, 1936a: 109. cf. Microciona prolifera; Vosmaer, 1935a: 610, 635, 665. MATERIAL. HOLOTYPE: BMNH1887.5.2.99: Bass Strait, Tas, 36°59'S, 150°20'E, 4.iv.1874, coll. HMS 'Challenger' (trawl). OTHER MATERIAL: NSW AMG5675, AMZ131, AMZ1413, AMZ1414. HABITAT DISTRIBUTION. Depth 110-300m, substrate mud; Bass Strait (Tas) (Ridley & Dendy, 1886), S. coast (NSW) (Whitelegge, 1907). Ternate, Mollucas, Indonesia (Kieschnick, 1896). DESCRIPTION. Shape. Erect, irregularly cylindrical digits, 38-120mm high, 8-12mm diameter, encrusting on organic debris or standing free in substrate. Colour Grey- or yellow-brown preserved. Oscules. Numerous oscules, up to 2.5mm diameter, scattered over branches. Texture and surface characteristics. Surface rugose, reticulate, minutely hispid; texture firm, flexible. Ectosome and subectosome. Ectosomal skeleton prominently hispid, with choanosomal principal megascleres from peripheral fibres protruding a long way through surface, and with a sparse tangential layer of subectosomal auxiliary megascleres dispersed between erect principal spicule brushes; subectosomal skeleton plumose, undifferentiated from choanosomal fibres which are immediately subdermal. Choanosome. Choanosomal skeleton irregularly reticulate, with relatively heavy spongin fibres incompletely divided into primary ascending and secondary transverse components; secondary fibres uncored, lightly echinated by small acanthostyles; primary fibres contain sparse tracts of subectosomal auxiliary subtylostyles, identical to those occurring in the ectosomal skeleton, enclosed completely within spongin fibres, together with plumose brushes of choanosomal ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 117 FIG. 44. Clathria (Clathria) inanchorata Ridley & Dendy (holotype BMNH1887.5.2.99). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Accolada toxa. E, Oxhorn toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype. 118zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 45. Clathria (Clathria) inanchorata Ridley & Dendy (holotype BMNH1887.5.2.99). A, Choanosomal skeleton. B, Ectosomal spicule bundles. C, Ectosomal skeleton. D, Fibre characteristics. E, Echinating acanthostyle. F, Smaller echinating acanthostyle. G, Acanthostyle spination. H-I, Ozhorn toxas. J, Portion of accolada toxa. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 119 TABLE 6.Comparison between present and published records of Clathria (Clathria) inanchorata Ridley & Dendy. All measurements are given in pm, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Choanosomal principal styles Subectosomal auxiliary styles Echinating acanthostyles I Echinating acanthostyles II Chelae Toxas I Toxas II Holotype (BMNH1887.5.2.99) Specimens (N=4) 292-(417.6)-535 x 8- 273-(384.5)-540x (18.4)-28 12-(17.8)-25 2524328.6)-432 x 3- 290-(341.8)-410 x 3(8.2)-11 (8.5)-12 58-(66.4)-78 x 3(5.2)-7 58-(75.2)-88 x 4(5.4)-7 1184150.2)-175 x 4- 119-(169.3)-228 x 6(7.4)-11 (8.8)-12 absent absent 35-(73.8)-121 x 1.5- 22-(66.2)-105 x 1.5- (2.0)-3 (3.4)-5 118-(349.2)-478 x 3044408.3)-545 x 1.5-(2.1)-2.5 1.5-(2.1)-3 principal styles usually poking out of fibres; choanosomal principal styles, protruding through spongin fibres, together form multispicular ascending plumose tracts, also lightly echinated by acanthostyles; fibre anastomoses form circular to oval, cavernous meshes; mesohyl matrix very light, with few megascleres dispersed between fibres. Megascleres (Table 6). Choanosomal principal styles long, thick, fusiform, slightly curved, with rounded or slightly subtylote, smooth or minutely microspined bases. Subectosomal auxiliary subtylostyles straight, relatively thick, robust, almost hastate, with slightly subtylote microspined bases. Acanthostyles very variable in size, with slightly subtylote bases, incompletely separated into two size classes with some intermediate examples. Smaller morph usually straight, often with aspinose necks, whereas larger morphs slightly curved, with evenly distributed large spines. Microscleres. (Table 6). Isochelae absent. Toxas clearly separated into two morphs - I: most common form are oxhom toxas, small, relatively thick, with large, rounded or slightly angular curvature at centre, and reflexed points. II: Less frequent are accolada toxas, long, thin, sharply angular at centre, unreflexed arms. REMARKS. This species has a distinctive growth form, spicule geometry, and spongin fibre characteristics, but otherwise it is similar to other species included in Hallmann's (1912) spicata' group, particularly C. (T) costifera and C. (C.) caelata. There is no doubt that Whitelegge's (1907) specimens from Wollongong are conspecific although this claim was disputed by Hallmann (1912: 206). Kiescknick's (1896) record of this species from Indonesia is dubious, since his material was not described and could possibly be any one of these spicata' -like sponges. Thiele (1903a) compared Kieschnick's specimen with C. (T)coralliophila from the same region, but that comparison is misleading: both taxa have quite different spiculation and spongin fibre characteristics. Clathria (Clathria) kylista Hooper & Levi, 1993 (Figs 46-47, Plate IC, Table 7) Clathria (Clathria) kylista Hooper & Levi, 1993a: 1265-1267, figs 21-22, table 11; Hooper & Wiedenmayer, 1994: 259. MATERIAL. HOLOTYPE: QMG300035: Inner Gneering Shoals, off Mooloolaba, Qld., 26°38.5'S, 153°09.5'E, 10m depth, 10.xii.I 991, coll. J.N.A. Hooper & S.D. Cook (SCUBA). PARATYPE: QMG300690 (ORSTOM R1338; fragment NTMZ3876): N. entrance, Recif des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S, 166°44.4'E, 8m depth, 30.iv.1976, coll. G. Bargibant (SCUBA). OTHER MATERIAL: QUEENSLAND - QMG303166. HABITAT DISTRIBUTION. 8-20m depth; on rock pinnacles, in caves and coral rubble substrates; Mooloolaba and Moreton I. (SEQ) (Fig. 46F). New Caledonia (Hooper & Levi, 1993a). DESCRIPTION. (See Hooper & Levi, 1993a). DIAGNOSIS. (refer to Table 7 for spicule dimensions) Simple digitate, tubular or bulbous, erect, branching growth form; dark orange to pale orange alive; terminal osculum on single tubes and oscules scattered over the apical regions of more complex lobate digitate growth forms; oscules surrounded by membraneous lip; prominently conulose surface with large, bulbous tubercles; ectosome membraneous, with sparse, tangential layer of subectosomal auxiliary styles and erect brushes of auxiliary spicules scattered throughout both peripheral and subectosomal regions; choanosomal skeleton plumoreticulate with differentiated primary and secondary fibres; primary fibres ascending, paucispicular occupying only small proportion of fibre diameter, cored by principal spicules with fewer auxiliary megascleres interdispersed; secondary fibres transverse, uni- or aspicular; echinating styles MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVUT 120^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA B A C E 1.0 ('.1 E azyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO 0 0 FIG. 46. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hr ia ( Clat hr ia) ky list a Hooper & Levi (A-D, QMG303166; E, holotype QMG300035). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada toxas. E, Section through peripheral skeleton. F, Australian distribution. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 121 FIG. 47. Clathria (Clathria) kylista Hooper & Levi (paratype QMG300690); A, Choanosomal skeleton. B, Fibre characteristics in peripheral skeleton. C-D, Echinating acanthostyles. E, Acanthostyle vestigial spines. F, Subtylote base of auxiliary subtylostyles. 122zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 7. Comparison in spicule dimensions between Clathria (Clathria) lipochela Burton, 1932 types and specimen of zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Clat hria) kylist a (Fig. 48) Hooper & Levi. All measurements are given in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). Burton, 1932a: 319, figs 6-7, textfig.29; Burton, 1940: 109, p1.4, fig.5; Koltun, 1964a: 69; Desqueyroux, 1972: 26-27, figs 87-89, 135; Sara, 1978: 65; Hooper & Wiedenmayer, 1994: 260. liolotype Paratype Specimen Thalyseurypon lipochela; de Laubenfels, 1936a: 107. (QMG300035) (QMG300690) (QMG303166) SPICULEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA (SE. Qld) Choanosomal 64484.7)-112 principal x 2.5-(2.9)-4.0 styles (New Cal.) (SE. Qld) 61-(81.6)-97 x 1.6-(2.7)-3.6 55478.2)-104 x 2.043.1)-4.5 Subectosomal auxiliary styles 143(168.9)181 x 1.0(2.4)-4.1 138(159.4)183 x0.4(2.1)-3.5 110(142.3)166x 1.0(1.8)-3.5 Ectosomal aux -iliary styles absent absent absent 32-(35.5)-39 x 1.1-(3.1)-5.0 18-(27.8)-35 x 1.0-(2.4)-3.5 26-(32.8)-40 x Echinating acanthostyles 1.042.6)-4.0 Toxas 65-(129.6)231.4 x 0.5(1.3)-2.5 65-(135.6)266 x 0.2(1.1)-1.6 35-( 146.5)222 x 0.5(1.0)-2.0 Chelae absent absent absent moderately common on primary ascending fibres, sparse on secondary connecting fibres; choanosomal principal styles slender, straight, relatively short, with hastate points and smooth, slightly swollen, subtylote bases; subectosomal auxiliary styles long, slender, straight, with hastate points and smooth, elongated, swollen subtylote bases; echinating styles entirely smooth, short, slender, sharply pointed, with prominent basal constriction ('neck'), subtylote base and widest just below basal constriction; isochelae absent; accolada toxas abundant, moderately long, ranging from slender to raphidiform, with straight arms and prominent central curve. REMARKS. This species was assigned to Clat hria ( Clat hria) by Hooper & Levi (1993a), even though echinating spicules are smooth (cf. Echinoclat hria) , because coring (principal) and echinating megascleres have different geometries (see also C. (M.) acerat oobt usa) . Clat hria ( C.) kylist a is similar to C. ( C.) angulifera and C. (C.) noarlungae in skeletal structure, having sparsely cored ascending primary fibres and uncored secondary connecting fibres, although both these other species have acanthose echinating spicules and palmate isochelae, and C. (C.) noarlungae also has slightly curved toxas. The unusual geometry of the echinating styles in C. (C.)kylista is the strongest apomorphy for the species, not seen elsewhere in the genus. Clat hria lipochela MATERIAL. HOLOTYPE: BMNH1928.2.15.352: Eddystone Rock, Falkland Is, 105-115m depth, coll. HMS 'Discovery' (trawl). HABITAT DISTRIBUTION. 22-1I5m depth; on sand and hard substrates; King George Land (Australian Antarctic Territory), Antarctica (Koltun, 1964a) (Fig. 48F). South Georgia (Koltun, 1964a), Falkland Is (Burton, 1932a), Caleta Santa Marta, Chile (Desqueyroux, 1972), Mar del Plata, Argentina (Burton, 1940), Kerguelen Is (Koltun, 1964a), Magellan Straits (Burton, 1940), Cape Sebastiano, Punta Arenas, Rio Grande, Cape Domingo, Cape Viamonte, Tierra del Fuego (Sara, 1978). DESCRIPTION. Shape. Stalked, irregularly flabellate; digitate margins of fan. Colour. Unknown. Oscules. Not seen. Text ure and surface charact erist ics. Firm, compressible; uneven, pitted, porous surface. Ect osom e and subect osom e. Surface skeleton with sparse paratangential or erect brushes of subectosomal auxiliary styles, mainly at ends of ascending primary spicule tracts; choanosomal principal styles protrude through surface in places; detritus scattered over surface but not embedded in ectosome. Choanosom e. Skeletal architecture regularly reticulate with ascending primary multispicular fibres and transverse uni-, pauci- or occasionally aspicular fibres, interconnecting at more-or-less regular intervals producing rectangular or elongate meshes, up to 4501.1.m diameter; fibres cored by choanosomal principal styles and moderately heavily echinated by acanthostyles evenly scattered over fibres in both axial and peripheral regions of skeleton; spongin fibres well developed; few spicules scattered between fibres; mesohyl matrix light, smooth, virtually unpigmented. Megascleres. Choanosomal principal styles short, relatively slender, fusiform, rounded and smooth bases, straight or only slightly curved near basal end. Length 153-(206.7)-254Rm, width 7-(11.0)-14p.m. Subectosomal auxiliary styles long, slender, slightly hastate pointed, slightly subtylote or REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 123 FIG. 48. Clathria (Clathria) lipochela Burton (holotype BMNH1929.2.15.352). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle and style. C, Echinating acanthostyle. D, Palmate isochelae. E, Section through peripheral skeleton. F, Antarctic distribution. G, Choanosomal skeletal structure. H, Fibre characteristics. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 124zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA rounded bases, microspined or smooth bases. Length 164-(179.4)-198p,m, width 3-(3.9)-5p.m. Echinating acanthostyles relatively long, slender, straight, with subtylote bases, fusiform points, spines concentrated in basal and apical regions and bare 'necks'. Length 79493.1)111p.m, width 6-(8.7)-1211m. Microscleres. Palmate isochelae small, unmodified, relatively abundant. Length 7-(8.5)11p,m. Toxas absent. TABLE 8. Comparison between present and published records of Clathria (Clathria)multipes Hallmann. All measurements are given in p,m, denoted as range (and mean) of sp'cule length x spicule width (N=25). Lectotype Paralectotype Paralectotype (BMNH1887.4. SPICULEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP (AMG9038) (AMG9162) 27.9) Citoaposomal 142-(136.6)-174 principal styles Subectosomal auxiliary styles I45-(157.6)-178 112-(147.8)-262 x 4-(5.9)-7 x 648.2)-10 x 7-(9.3)- 12 162-(194.6)-239 x I.5-(2.8)-4 1324167.0)-222 x 2-(3.1)-4 143-(189.5)-262 x 2-(3.2)-4.5 REMARKS. Burton (1932a) named this species 52462.8)-91 x 71-(85.8)-98 x 64-(78.5)-93 x Echinating for the apparent absence of chelae, but these were acanthostyles 4-(6.6)-9 4-(5.6)-8 5-(6.4)-8 found to be common in sections of the holotype. 5-(4.8)-8 4-(6.1)-8 3-(5.7)-8 Chelae Burton's (1932a) comparison with Raspaxilla 16-(112.2)-147 19-(101.4)-141 1304124.4)-178 phakellina Topsent (Hooper, 1991: 1199), is misToxas x 1-(4.3)-6 x 2-(4.5)-6 x I-(3.9)-5 leading as they do not resemble each other in skeletal structure or spicule geometry. Clathria (C.) lipochela resembles C. (T) vulpina Colour. Live colouration brick red, grey-brown (Lamarck) in geometry of principal and echinat- dry. ing spicules and skeletal architecture to some Oscules. Numerous large oscules, up to 3mm extent, although they differ in many other fea- diameter, distributed over all surfaces. Texture and surface characteristics. Surface irtures.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA regularly rugose, with small elevated conules scattered mainly on lateral sides of branches. Clathria (Clathria) multipes Hallmann, 1912 Ectosome and subectosome. Ectosome, rarely in(Figs 49-50, Table 8) tact in dry type material, consists of three dimenClathria (Plectispa) arborea, in part, Whitelegge, sional fibre reticulation, with fibre endings 1901: 88, p1.11, fig.15. forming small surface conules, and choanosomal Plectispa macropora,in part, Lendenfeld, 1888: 225-6. principal styles protruding through peripheral Not Plectispa arborea Lendenfeld, 1888: 226. fibres in light brushes or singly; subectosomal Clathria (Plectispa) multipes Hallmann, 1912: 204, auxiliary subtylostyles dispersed in a tangential 211. layer around projecting dermal fibres. Clathria multipes; Hooper & Wiedenmayer, 1994: Choanosome. Choanosomal skeletal architecture 260. irregularly reticulate, consisting of relatively MATERIAL. LECTOTYPE: AMG9038(dry): Tug- heavy spongin fibres forming incompletely difgerah Beach, Illawarra region, NSW, 34 ° 32'S, ferentiated primary (vaguely ascending, multi150°50'E (beach debris, label 'Clathria arborea Len- spicular) and secondary fibres (transverse uni-, denfeld; ms name = Clathria plicatella'). PARALECpauci- or aspicular), and producing relatively TOTYPES: AMG9162 (dry): Maroubra Bay, NSW, 33°455, 151°20'E (label Tlectispa arborea Lend. = tight oval to elongate meshes; fibres echinated by Clathria arborea'). BMNH1887.4.27.9 (fragment small, sparsely distributed acanthostyles; AMG3590): Port Jackson, NSW, 33 ° 51'S, 151°16'E mesohyl matrix light, with moderate quantities of (label `Thalassodendron reticulata RvL, MS'). Other subectosomal subtylostyles and microscleres distype fragments ZMB2264, 6894. SYNTYPE of P. persed. macropora: BMNH1925.11.1.555: Manly Beach, Megascleres (Table 8). Choanosomal principal NSW, 33°50'S, 151°17'E, other details unknown. styles hastate or stepped, relatively thick, slightly HABITAT DISTRIBUTION. Ecology unknown; curved, with tapering or slightly subtylote and smooth bases. central and S coast (NSW) (Fig. 49F). Subectosomal auxiliary subtylostyles long, DESCRIPTION. Shape. Branching, reticulate thin, fusiform, straight, curved or sinuous, with branches, planar, 85-140mm long, 62-73mm slightly subtylote, smooth bases. wide, with compressed, cylindrical and regularly Acanthostyles subtylote, with vestigial spinaanastomosing branches,4-8mm diameter, and tion and an aspinose neck. several small basal stalks (multiple points of at- Microscleres (refer to Table 8 for dimensions). tachment). Palmate isochelae minute. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 125 FIG. 49. Clathria (Clathria) multipes Hallmann (lectotype AMG9038). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Palmate isochelae. E, U-shaped toxas. F, Australian distribution. G, Section through peripheral skeleton. H, Paralectotype AMG9162. I, Lectotype. Toxas u-shaped, relatively thick, oxeote, with hastate points, typically curved at right angles at the centre, with straight and unreflexed points. REMARKS. All known specimens are in poor condition, and it is not possible to accurately determine ectosomal characteristics. The species is most closely related to Clathria (Clathria) rather than Clathria (Thalysias). Whitelegge (1901) remarked that the species was frequently washed up onto coastal beaches of S NSW after storms, inferring that it was a relatively prominent component of the benthos, and therefore it is surprising that it has not been collected since that time despite intensive trawling. The growth form of C. (C.) multipes (reminiscent of MEMOIRS OF THE QUEENSLAND MUSEUM 126^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 50. Clathria (Clathria) multipes Hallmann (lectotype AMG9038). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F,U-shaped toxas. G, Palmate isochelae. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 127 EI tO FIG. 51. Clathria (Clathria) murphyi sp.nov. (holotype QMG300656). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype in situ. C. (T) coppingeri and Echinodictyum cancellatum (Raspailiidae)), the small size of isochelae, the peculiar angular shapes of toxas, and the vestigial acanthostyles differentiates this species from other Clathria. Clathria (Clathria) murphyi sp. nov. (Figs 51-52, Plate 1D) MATERIAL. HOLOTYPE: QMG300656 (NCIQ66C2904-N, fragment NTMZ3754): Old jetty, E. end of Princess Royal Drive, Albany, WA, 35°02.3'S, 117°54.2'E, 9m depth, 27.ii.1989, coll. NCI (SCUBA). HABITAT DISTRIBUTION. Wood jetty piles; 9m depth; SW WA (Fig. 51G). DESCRIPTION. Shape. Thickly encrusting, bulbous digitate lumps, up to 80mm diameter, resembling the tropical Higginsia massalis (Desmoxyidae). Colour. Orange-red alive (Munsell lOR 6/10), pale brown preserved. MEMOIRS OF THE QUEENSLAND MUSEUM 128zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 52. Clathria (Clathria) murphyi sp.nov. (holotype QMG300656). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Base of acanthostyle. E, Acanthostyle spines. F, Palmate isochelae. G, Accolada toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 129 Oscules. Large oscules, more than 5mm diameter, on ends of bulbs. Texture and surface characteristics. Surface porous, microvillose, uneven, with epi- and zoophytes; texture soft, compressible, easily torn off wooden piles. Ectosome and subectosome. Ectosome membraneous, with heavy, brown collagen and brushes of choanosomal principal styles poking through surface in plumose bundles, mainly on ends of microconules; subectosomal auxiliary styles in irregular, paratangential bundles below surface, at base of principal style brushes, not protruding through surface. Choanosome. Choanosomal skeleton renieroid reticulate, cavernous, without visible spongin fibres, with differentiated primary and secondary spicule tracts; primary skeletal tracts ascending, multispicular, 20-40p,m diameter, composed of plumose brushes of choanosomal principal styles, 3-10 spicules abreast; secondary tracts uni-, bi- or paucispicular, transverse, 8-20p,m diameter, connecting primary tracts; echinating acanthostyles relatively sparsely dispersed throughout choanosome; mesohyl matrix heavy, granular, darkly pigmented, surrounding large oval, paired choanocyte chambers, up to 320Rm diameter, with abundant toxa and isochelae microscleres dispersed throughout. Megascleres. Choanosomal principal subtylostyles short, stout, straight, fusiform points, tapering smooth bases, terminally subtylote, with slightly swollen subterminal region. Length 87(116.5)-149pin 1, width 5-(7.1)-911m. Subectosomal auxiliary subtylostyles short, slender, fusiform, slightly subtylote, smooth bases. Length 114-(138.4)-165p..m, width 1.5- Echinating acanthostyles short, slender, slightly subtylote bases, spines recurved, evenly spined except for bare neck. Length 42455.7)-6811m, width 3.5-(4.2)-5.5vm. Microscleres. Palmate isochelae, small, poorly silicified, about 10% with twisted shafts. Length 5-(8.4)-14p.m. Toxas accolada form, long, very slender hairlike, with straight, unreflexed arms and angular central curvature. Length 72-( 115.6)-16411m, width 0.5-(0.7)-0.4,m. ETYMOLOGY. For Dr Peter Murphy, Australian Institute of Marine Science, Townsville. REMARKS. This species is separated from other Clathria (Clathria) by its distinctive spicule geometries (principal styles with marked basal constrictions and swollen 'necks'), relatively small, poorly silicified spicules of all categories, renieroid choanosomal skeletal structure (with multispicular ascending tracts and uni- or paucispicular plumose transverse connecting spicule tracts, both sparsely echinated by acanthostyles), bulbous-digitate growth form, orangered colouration, microvillose surface with plumose brushes of choanosomal styles protruding through the ectosome especially on the tips of microconules. Clathria (C.)inurphyi has a skeletal architecture reminscent of C. (C.) arcuophora and C. (C.) crassa (both of which have much larger spicules of different geometry), and in this respect the species is included in the `striata' group (see remarks for C. (C.) striata). Clathria (Clathria) nexus (Koltun, 1964) (Figs 53-54)zyxwvutsrqponmlkjihgfedcb Bipocillopsis nexus Koltun, 1964a: 79-80. Clat hria nexus; Hooper & Wiedenmayer, 1994: 260 MATERIAL. HOLOTYPE: ZIL 10644 (not seen): Clarie Coast or Wilhelm Land, Australian Antarctic Territory, 65 °48' S, 89 °49'E, 310-400m depth (dredge). PARATYPES: BMNH1963.7.29.56, ZIL 11525: same locality. HABITAT DISTRIBUTION. 310-400m depth; substrate unknown; Australian Antarctica Territory (Fig. 53F). DESCRIPTION. Shape. Erect arborescent growth form, 88-100mm high, 32-55mm wide, with tightly anastomosing cylindrical branches, up to 4mm diameter; tips of branches bifurcate, relatively sharply pointed. Colour. Grey-brown preserved. Oscules. Numerous small oscules, 1-2mm diameter, scattered over lateral margins of all branches. Texture and surface characteristics. Surface hispid, raised into irregularly distributed, sharply pointed microconules. Ectosome and subectosome. Ectosomal skeleton with choanosomal principal styles erect on surface, and bundles of subectosomal auxiliary spicules surrounding protruding principal spicules, or lying paratangential to surface. Choanosome. Choanosomal skeleton subrenieroid-reticulate, without clearly defined spongin fibres, but with heavy collagen enclosing principal subtylostyles and sparse acanthostyles echinating; mesohyl matrix heavy, with numerous isochelae, few auxiliary styles and some detritus dispersed throughout. MEMOIRS OF THE QUEENSLAND MUSEUM 130^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA oEl FIG. 53. Clathria (Clathria) nexus Koltun (paratype BMNH1963.7.29.56). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Unguiferous palmate isochclae. E, Section through peripheral skeleton. F, Antarctic distribution. G, Paratype. Megascleres. Choanosomal principal styles long, Microscleres. Isochelae, strongly curved sig- robust, straight or slightly curved towards the distal end, tapering to sharp points, with slightly swollen subtyloted bases, liberally microspined; basal spines may partially extend up shaft near basal end. Length 5184567.3)-62011m, width 18(22.4)-2511m. Subectosomal auxiliary styles long, straight, abruptly pointed,with only slight basal swelling and pointed-hastate bases. Length 366-(394)415m, width 3-(5.6)-7[Lm. Echinating acanthostyles relatively long, robust, subtylote, sharply pointed, with heavy recurved spines and heaviest concentrations of spines on basal end. Length 214-(241.3)-278Rm, width 9-(13.8)-17Rm. moid, unguiferous with vestigial teeth, of anchorate or arcuate modification. Length 14(16.6)-19iLm. Toxas absent. REMARKS. This species is similar to C. (T) michaelseni in having bidentate sigmoid isochelae, but differing in skeletal architecture and in most other features. These unguiferous, sigmoid-like chelae with vestigial, pointed alae, are not unique to these austral species, also known in W. Indian Ocean C. (C.) spongodes Dendy (including its synonym C. madrepora Dendy). In C. (C.) spongodes these reduced chelae were initially thought to be sigmas (Dendy, 1922; Burton, 1959a; Vacelet et al., ^ 131 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT FIG. 54. Clathria (Clathria) nexus Koltun (paratype BMNH1963.7.29.56). A, Choanosomal skeleton. B, Peripheral skeleton C, Ectosomal structure. D, Fibre characteristics. E, Principal subtylostyles. F, Bases of principal subtylostyles. G, Base of auxiliary style. H, Echinating acanthostyle. I, Acanthostyle spines. J, Unguiferous palmate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM 132zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 1976), but the type material showed that these are in fact reduced unguiferous isochelae with vestigial teeth. Clathria (C.) nexus is also unusual in Microcionidae in having auxiliary styles obviously associated with (surrounding) protruding principal spicules, reminiscent (or analogous to) Raspailiidae. Unlike raspailiids, however, auxiliary spicules are also dispersed within the peripheral skeleton, paratangential to the surface. The other alternative, that the species belongs to Raspailiidae, is rejected due to the chelae microscleres. The bidentate-derived chelae is unusual to the Microcionidae and grounds to exclude the species from it under the phylogeny of Poecilosclerida hypothesised by Hajdu et al. (1994), but in all other respects the species fits in this group supporting the present classification. Clathria (Clathria) noarlungae sp.nov. (Figs 55-56, Plate 1E, Table 9) MATERIAL. HOLOTYPE: SAMTS4047 (fragment NTMZ1632): Port Noarlunga, SA, 35 °09'S, 13 29'E, 1973, coll. SA Fisheries (trawl). OTHER MATERIAL: S AUST - QMG300247 (NCIQ66C-2468-X, fragment NTMZ3566). HABITAT DISTRIBUTION. 5-30m depth; from rock reef substrate; Port Noarlunga and Kingston, SE S.Aust. DESCRIPTION. Shape. Branching, 58-190mm long, cylindrical digitate, bulbous-lobate branches, 15-33mm diameter, with bulbous lobes on tips of digits, single or bifurcate tips. Colour. Orange alive (Munsell 1OR 6/10), greybrown preserved (7.5YR 5/4). Oscules. Large oscules, 3-5mm diameter, in-line on lateral sides of branches and on apex of terminal bulbs. Texture and suiface characteristics. Texture rubbery, compressible, fibrous, difficult to tear; surface optically smooth, with few low rounded bulbous projections; detachable skin-like dermis, which is microscopically evenly porous, without projecting spicules. Ectosome and subectosonie. Ectosomal skeleton consisting of relatively even, light series of plumose brushes of ectosomal auxiliary subtylostyles, standing perpendicular or paratangential to surface, usually surrounding inhalant pores and forming a spiculo-fibrous reticulation on surface; ectosomal skeleton thin, 75-149pm wide, perched on ends of peripheral, ascending choanosomal fibres; subectosomal skeleton not differentiated from dermal skeleton, and ectosomal spicule brushes composed of a single category of auxiliary megascleres only; megascleres coring peripheral fibres sometimes project into, but not through ectosomal skeleton, but this is exceptional. Choanosome. Choanosomal skeletal architecture arborescent, cavernous, consisting of well differentiated primary ascending and secondary transverse components; primary spongin fibres multispicular, 35-98Rm diameter, relatively heavily invested with spongin, forming dendritic, arborescent, radial, relatively even structure; coring spicules in primary fibres occupy only a small proportion of sponge diameter in axial skeleton, becoming more heavily cored and increasingly plumose towards peripheral skeleton; spongin fibres cored by larger subectosomal auxiliary styles, and echinating acanthostyles occur only sparsely throughout entire skeleton; secondary spongin fibres entirely free of coring megascleres, 12-57p.m diameter, forming a plumo-reticulate structure; meshes formed by primary and secondary spongin fibre branching are ovoid-elongate to rectangular in shape, relatively even, and markedly cavernous (155560p,m maximum diameter); choanocyte chambers paired, 90-311 p.m maximum diameter, becoming larger towards periphery; mesohyl matrix slightly granular, with toxas dispersed singly or in dragmata; thin, rhaphidiform, vestigial auxiliary megascleres also dispersed between fibres, difficult to distinguish from toxas. Megascleres (refer to Table 9 for dimensions). Choanosomal principal megasclere absent, or completely undifferentiated from subectosomal auxiliary spicules. Subectosomal auxiliary styles, coring fibres, thin, mostly straight, varying from hastate to almost fusiform, with smooth tapering, rounded or very slightly subtylote bases. Ectosomal auxiliary styles geometrically similar to larger auxiliary styles, with slightly more pronounced, smooth, subtylote basal constrictions. Acanthostyles vestigial, with subtylote bases, fusiform points, rudimentary spination, aspinose points and necks. Vestigial auxiliary megascleres dispersed within mesohyl are styloid or quasidiactinal, often sinuous, with tapering or rounded bases, long fusiform points. Microscleres (refer to Table 9 for dimensions). Palmate isochelae minute, abundant, frequently twisted (80% of samples), two size classes observed. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 133 Toxas accolada, rhaphidiform, common, occurring individually or in dragmata, exceedingly thin, rhaphidiform with very slight rounded central curvature and straight points, or less often with more angular central curvature and slightly reflexed arms. ETYMOLOGY. Named for type locality. REMARKS. There are some minor differences in spicule dimensions between the two known specimens of C. (C.) noarlungae (Table 9), although there is no doubt that they are conspecific. In having sparsely cored ascending primary fibres and uncored secondary connecting fibres this species is similar to C. (C.)kylista and C. (C.) angulifera although spicule geometry and dimensions differs between all three. This species is also similar to C. (T) cactiformis (Lamarck) in growth form and skeletal structure but they differ in geometry and size of spicules, number of spicule categories and structure of ectosomal and subectosomal skeletons. This species is enigmatic in its higher systematic placement. On the one hand C. (C.) noarlungae has a classical, albiet thin ectosomal skeleton typical of Thalysias species, consisting of erect plumose brushes of smaller ectosomal auxiliary spicules. This feature is structurally discrete from the primary dendritic, and secondary plumo-reticulate choanosomal skeleton. Conversely, there is only a single category of auxiliary style which forms these ectosomal brushes, whereas the larger (subectosomal) auxiliary spicules are confined to inside the choanosomal fibres. Consequently this species technically belongs to C. (Clathria) (in having an undifferentiated ectosomal-subectosomal skeletal composition). Clathria (C.) noarlungae may be confused with C. (Dendrocia) curvichela and C. (D.) elegantula due to superficial similarities in spiculation and fibre characteristics, but it differs from these (and other Dendrocia) in having two different auxiliary spicule geometries, with palmate isochelae instead of arcuate-like isochelae, and different skeletal construction. This species differs from all other known Clathria (and other microcionids) in several features: orange colouration; bulbous-lobate digitate growth form; rounded bulbous surface projections; absence of choanosomal principal styles, where dendritic multispicular primary spongin fibres are cored by subectosomal auxiliary styles, and the secondary fibre system is aspicular, plumo-reticulate, and sparsely echinated by yes- tigial acanthostyles; and specific spicule geometries. Clathria (Clathria) oxyphila (Hallmann, 1912) (Figs 57-58, Table 10) Wilsonella oxyphila Hallmann, 1912: 249-253, p1.34, fig.3, text-fig.52; Guiler, 1950: 9. Paradoryx oxyphda; Hallmann, 1920: 768. cf. Clathria elegantula; Hallmann, 1912: 253. cf. Clathria pintformis; Carter, 1885f: 354; Hallmann, 1912: 253. Clathria oxyphila; Hooper & Wiedenmayer, 1994: 260. MATERIAL. HOLOTYPE: AMZ51 (dry; fragment AME8 I 7): Off Marsden Point, Kangaroo I., SA, 35°30'S, 137°45'E, 34m depth, 19.viii.1909, coll. FIV `Endeaour (dredge; label Wilsonella oxyphila (curvichela) Hallmann, type'). PARATYPE: AME495 (dry): Oyster Bay, Tas, 42°40'S, 148°03'E, 60-80m depth, coll. FIV 'Endeavour', dredge; (specimen label `Wilsonella oxyphila, Type'; AM register MS name Taraclathria oxyphila sp. nov.'). HABITAT DISTRIBUTION. 34-80m depth; substrate unknown; E. coast (Tas.) and Kanagaroo I. (S. Aust.) (Fig. 57G). DESCRIPTION. Shape. Fan, 152mm high, 155mm wide, planar or multiplanar, with a small compressed, cylindrical basal stalk, 22mm long, 18mm diameter, thickly lobate, rounded, even margins. Colour. Grey-brown preserved. Oscules. Oscules dispersed on margins of branches. Texture and surface characteristics. Surface relatively even, with radiating longitudinal subdermal grooves and pitted and porous ectosome. Ectosome and subectosome. Ectosome membraneous with relatively heavy mesohyl matrix below surface, sparsely arenaceous, with very few choanosomal principal megascleres protruding through surface, and with sparse ectosomal skeleton of subectosomal auxiliary megascleres lying paratangential to surface, sometimes in light brushes around fibre endings. Choanosome. Choanosomal fibres immediately subdermal, becoming slightly plumose near periphery; choanosomal skeleton sometimes dendritic in places, usually plumo-reticulate. Spongin fibres well developed, clearly divided into primary (ascending, sinuous, multispicular) and connecting (transverse, aspicular) elements; coring megascleres vestigial principal styles, barely differentiated from subectosomal auxiliary styles; spongin fibres heavily echinated MEMOIRS OF THE QUEENSLAND MUSEUM 134zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ All FIG. 55. Clathria (Clathria) noarlungae sp.nov. (holotype SAT54047). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle and modified style. D, Raphidiform-accolada toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, fragment of holotype. I, Specimen NTMZ3566. by acanthostyles, evenly dispersed over fibres; mesohyl matrix heavy, with few scattered subectosomal megascleres. Megascleres (Table 10). Choanosomal principal styles slightly shorter and thicker, but otherwise with similar geometry to those occurring in peripheral skeleton; straight, hastate, with tapering or quasidiactinal, smooth bases. Subectosomal auxiliary styles long, thin, slightly curved or sinuous, often with blackened axial canals; geometrically similar to principal megascleres. Acanthostyles slightly subtylote, evenly spined, including oxeote modifications. Microscleres (Table 10). Palmate isochelae sig- moid with reduced alae or vestigial teeth resem- bling true sigmas; isochelae predominant on membraneous ectosome. Toxas intermediate between wing-shaped and u-shaped, uncommon, with low, rounded central curves, slightly reflexed points. Larvae. Incubated parenchymella larvae ovalelongate, 120 x 210vt,m, with moderately light mesohyl. REMARKS. This species was included in Wilsonella by Hallmann (1912, 1920) and other authors on the basis that principal and auxiliary styles were allegedly undifferentiated, but re-examination of type material found them to be similar but not identical in geometry (both relatively vestigial and modified quasidiactinal form). In this respect the species is similar to C. ^ 135 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT FIG. 56. Clathria (Clathria) noarlungae sp.nov. (QMG300247). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating spicules in situ. D, Echinating acanthostyle. E, Acanthostyle spines. F, Raphidiform-accolada toxas. G, Palmate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM 136^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 57. Clathria (Clathria) oxyphila (Hal!mann) (holotype AMZ51). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Intermediate wing shaped - u-shaped toxa. E, Sigmoid palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Paratype AMFA95. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 137 FIG. 58. Clathria (Clathria) oxyphila (Hallmann) (fragment of holotype AME817). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spines. F, Sigmoid palmate isochelae. G, Intermediate wing-shaped - u-shaped toxa. MEMOIRS OF THE QUEENSLAND MUSEUM 138zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 9. Comparison between present and published records of Clathria (Clathria)noarlungae sp.nov. All measurements are given in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). TABLE 10. Comparison between present and published records of Clathria (Clathria) oxyphila (Hallmann). All measurements are given in p,m, denoted as range (and mean) of spicule length x spicule width (N=25). Holotype (SAMTS4047) Specimen (NC1Q66C2468X) Holotype (AMZ51) Paratype (AME495) Choanosomal principal styles absent absent Choanosomal principal styles 104-(133.0)-152 x 1.8-(2.5)-3 119-(135.4)-154 x 1.5-(2.2)-3 Subectosomal auxiliary styles 167-(204.4)-236 x 2.5-(4.8)-7.8 232-(250.2)-278 x 3(4.3)-5 Subectosomal auxiliary styles 142-(171.1)-199 x 0.8-(1.4)-2 143-(164.0)-186 x 1.0-(1.6)-2.0 Ectosomal auxiliary styles 87.5-(112.8)-156 x 1.8-(3.4)-4.6 117-(146.0)-175 x 2(2.7)-3.5 Echinating acanthostyles 49-(60.3)-72 x 2.5(4.1)-4.5 54-(63.6)-71 x 3.5(4.2)-5.5 Echinating 36-(46.6)-54 x 3.8(5.2)-6.5 48-(55.6)-61 x 4.5(5.2)-6 Chelae 17-(21.8)-25 16-(19.6)-23 acanthostyles Vestigial auxiliary styles 133-(186.2)-202 x 1.5-(2.1)-2.6 118-(170.1)-186 x 1.0-(1.6)-2.0 Toxas 35@20)-64x 1.5(1.7)-2.0 I9-(50.6)-84x 1.0(1.4)-2.5 SPICULE Chelae I 3-(6.3)-8.5 5-(7.3)-I0 Chelae II 12-(13.5)-16.5 13-(15.7)-I9 36-(159.5)-216 x 0.4-(0.8)-1.1 19-(145.3)-265 x 0.5-(0.9)- 1.5 Toxas (C.) piniformis, C. (C.) raphanus and C. (Dendrocia) elegantula, all referred here to an artificial species-group termed the `oxyphila' group, loosely corresponding to Hallmann's (1912, 1920) concept of Wilsonella (which is also possibly artificial). The present species differs from these by its sigmoid microscleres (virtually intermediate between palmate and arcuate-like geometry). Hallmann (1912) suggested that these chelae were arcuate, and thus he considered that the species had affinities with C. (Dendrocia) curvichela, but they actually appear to be no more than slightly modified, vestigial palmate forms. Similarly, Hallmann (1912) did not record toxas in his description, which are definitely present in type material. The modification of echinating acanthostyles to acanthoxeote megascleres in this species is reminscent of Crellidae but this modification is superficial. Clathria (C.) oxyphila was transferred to Paradoryx on the basis of its alleged arcuate chelae, but in all other respects the species is clearly a Clathria (Clathria). Clathria (Clathria) partita Hallmann, 1912 (Figs 59-60) Clathria partita Hallmann, 1912: 223, p1.32, fig.3, text-fig.46; Hooper & Wiedenmayer, 1994: 260. Clathria (Clathria) cf. partita; Rudman & Avern, 1989: 335. Pseudanchinoe partita; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: AME1024(dry): Unknown locality, South Australian coast, 60m depth, SPICULE 12.vii.1909, coll. FIV `Endeavour' (dredge); (label `Pseudanchinoe partita, type'). HABITAT DISTRIBUTION. 60m depth; substrate unknown; SA (Hallmann, 1912), and possibly S WA (Rudman & Avern, 1989) (Fig. 59F). DESCRIPTION. Shape. Arborescent branching, 340mm long, 160mm wide, with a long cylindrical stalk, 55mm long, 14mm diameter, planar, bifurcate and occasionally anastomosing flattened branches, 12-25mm wide, approximately 8mm thick. Colour. Orange alive (Rudman & Avern, 1989), brown, with a slightly whitish ectosomal crust. Oscules. Small oscules, up to 2mm diameter, on lateral sides of branches. Texture and surface characteristics. Surface rugose, laterally striated, resembling Ectyoplasia tabula (Raspailiidae; Hooper, 1991: Fig. 50), and margins of branches slightly wider and more spatuliferous than basal portions. Ectosome and subectosome. Ectosomal skeleton almost halichondroid, well differentiated from choanosomal region, consisting of paratangential multispicular bundles of subectosomal auxiliary styles, forming criss-cross tracts on surface. Choanosome. Choanosomal architecture basically isodictyal, consisting of differentiated axial and extra-axial sections of skeleton; axial region composed of very heavy and sinuous spongin fibres, forming relatively tight, oval or elongated triangular meshes; axial fibres divided into larger primary and smaller secondary elements, both containing paucispicular core of choanosomal principal styles; extra-axial skeletal architecture an irregular isodictyal reticulation of sinuous spongin fibres of approximately same diameter as axial secondary elements, relatively lightly invested with spongin, containing uni-, bi-, or more REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 139 rarely paucispicular tracts of choanosomal styles; extra-axial fibres incompletely divided into primary (ascending) and secondary (vaguely transverse) components, differentiated mainly by number of coring spicules; echinating acanthostyles sparse, evenly distributed throughout skeleton, rarely incorporated into fibres (cf. Hallmann, 1912); some choanosomal principal styles may protrude through (echinate) fibres, particularly in extra-axial skeleton; mesohyl matrix very light containing few subectosomal auxiliary megascleres. Megascleres. Choanosomal principal styles short, slightly curved or straight, almost hastate, with smooth rounded or very slightly subtylote bases. Length 1124172.6)-23911m, width 5(8.7)-1211m. Subectosomal auxiliary styles long, slightly curved or straight, fusiform, with rounded or slightly subtylote bases. Length 2224267.7)315p.m, width 3.5-(5.1)-7p.m. Acanthostyles slightly subtylote, with vestigial spines and an aspinous basal region. Length 56(60.4)-8811m, width 3-(5.0)-61.m. Microscleres. Isochelae absent. Toxas sinuous rhaphidiform, usually symmetrical with slight angular central curvature and straight points, but asymmetrical and reflexed examples also occur. Length 724115.6)-2121m, width 0.5-(0.7)-1.2Rm. REMARKS. This species was referred to Pseudanchinoe by de Laubenfels (1936a) because it lacked chelae microscleres, but that feature has since been shown to be of little systematic value (e.g. Hooper, 1991: 288). As noted by Hallmann (1912), C. (C.) partita is known only from a single dry specimen, and it is possible that isochelae may be eventually discovered in other better preserved specimens should additional material become available. Clathria (C.) partita has an unusual skeletal construction, combining features of myxillid-like microcionids (e.g. a basically isodictyal extraaxial skeleton closely resembling C. (Isociella) eccentrica) and raspailiid-like microcionids (e.g. compressed axis of C. (Axociella) canaliculata), and features of the compressed axial skeleton are also reminiscent of the skeletal plan seen in certain Axinellidae such as Cymbastela. On the basis of these characteristics C. (C.) partita is similar to C. (C.) rubens (Lendenfeld). The record of C. (C.)partita from WA (Rudman & Avern, 1989), is based on a photograph without accompanying sample. The photograph was taken of a nudibranch (Rostanga calumnus Rudman & Avern) feeding on an orange sponge from Esperance Bay (33°51'S, 121°57'E). Clathria (Clathria) paucispicula (Burton, 1932) (Fig. 61) Rhaphidophlus paucispiculus Burton, 1932a: 320, p1.56, fig.1, text-fig.30; Burton, 1940: 111; Desqueyroux, 1975: 68; Koltun, 1964a: 75; Desqueyroux-Faundez & Moyano, 1987: 49. Clathria paucispicula; Hooper & Wiedenmayer, 1994: 260 MATERIAL. HOLOTYPE: BMNH1928.2.15.243a: Near Shag Rocks, South Georgia, 53°43.4'S, 40°57.0'W, 177m depth, coll. R.R.S. 'Discovery' (dredge). HABITAT DISTRIBUTION 74-198m depth; on mud, sand and rock substrates; Australian Antarctic Territory: Queen Mary Land (Koltun, 1964a) (Fig. 61D). Also Falkland Is, South Georgia and S. Shetland Is (Burton, 1932a), Mar del Plata, Argentina (Burton, 1940), Low Is, Chilean Antarctic Territory (Desqueyroux, 1975), Tierra del Fuego (DesqueyrouxFaundez & Moyano, 1987). DESCRIPTION. Shape. Massive, flabellatedigitate with irregular ridges and convoluted branches. Colour Live colouration unknown, light brown preserved. Oscules. Large oscules 3-5mm diameter sparsely scattered on apex of ridges. Texture and surface characteristics. Texture firm, tough, compressible; surface uneven, minutely conulose. Ectosome and subectosome. Surface skeleton tangential confused crust of smaller ectosomal auxiliary styles, choanosomal principal styles and foreign spicule fragments together forming a dense crust 0.5-1.0mm thick; immediately below tangential ectosomal skeleton are erect bundles of both auxiliary styles (producing an irregular radial palisade of spicules), and principal styles (more sparsely dispersed). Choanosome. Skeletal architecture reticulate, slightly plumo-reticulate near periphery, less organised towards axis; spongin fibres moderately well developed cored by pauci- or multispicular ascending tracts and interconnected by uni- or paucispicular tracts of choanosomal principal styles, producing rectangular or triangular meshes up to 250Rm diameter; echinating megascleres absent; mesohyl matrix light, smooth, unpigmented. MEMOIRS OF THE QUEENSLAND MUSEUM 140zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 59. Clathria (Clathria) partita Hallmann (holotype AME1024). A, Choanosomal principal styles. B, Subectosomal auxiliary style. C, Echinating acanthostyle. D, Sinuous-raphidiform toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 141 FIG. 60. Clathria (Clathria) part ita Hallmann (holotype AME1024). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Vestigial acanthostyle spines. F, Fragments of sinuous and raphidiform toxas. MEMOIRS OF THE QUEENSLAND MUSEUM 142^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 61. Clathria(Clathria)paucispicula (Burton) (holotype BMNH1928.2.15.243a). A, Choanosomal principal styles. B, Subectosomal auxiliary styles. C, Section through peripheral skeleton. D, Antarctic distribution. E, Choanosomal skeletal structure. Megascleres. Choanosomal principal styles long, thick, slightly hastate pointed or occasionally slightly telescoped pointed, curved at centre, with smooth rounded or slightly subtylote bases. Length 535-(663.4)-754Rm, width 18423.3)31pm. Subectosomal auxiliary styles relatively long, thick, straight or slightly curved near basal end with evenly rounded smooth bases and tapering fusiform points. Length 255-(318.6)-402Rm, width 6410.4)-1411m. Echinating spicules absent. Microscleres. Absent. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 143 REMARKS. This species resembles a Suberitidae (Hadromerida) in skeletal structure, with the main clue as to its affinities with Microcionidae being the possession of megascleres that are not truly tylote in geometry, two distinct categories of principal and auxiliary megascleres, and slight compression of the skeleton in the axial region not generally seen in suberitids. Certain allocation of this species is difficult given that spicule diversity and skeletal structure are the main diagnostic characters for microcionids, and the species is aptly named for the reduction in these characters. It is well known and possibly widely distributed in Antarctic waters. Clathria (Clathria) pauper Brondsted, 1926 (Figs 62-63) Clathria pauper Brondsted, 1926: 3, text-fig.3; Burton, 1929a: 398; Burton, 1940: 109; Koltun, 1964a: 69, p1.12, figs 4-6; Sara, 1978: 66-67, text-fig.40; Desqueyroux-Faundez & Moyano, 1987: 50. Ramoses pauper; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: NRHM (fragments BMNH 1930.11.5.2, AMZ2239): N. of Discovery Inlet, Victoria Land, Antarctica, 640m depth, 10.ii.I924, coll. Sten Warren (dredge). HABITAT DISTRIBUTION. 10-640m depth; hard and soft substrates; Australian Antarctic Territory: Victoria Land, Banzare Coast, Wilkes Land (Brondsted, 1926; Koltun, 1964a) (Fig. 62G). Also SW. Maldonado, Mar del Plata, Argentina, S. Brazil (Burton, 1940), Cape Sebastiano, Cape Domingo, Rio Grande, Tierra del Fuego (Sara, 1978). DESCRIPTION. Shape. Erect branching digitate sponge, up to 80mm high. Colour. Grey-brown preserved. Oscules. Unknown. Texture and surface characteristics. Texture firm, compressible, elastic; surface conulose with irregularly digitate processes, up to 20mm long and 4mm thick, and with convoluted distinct ectosomal membrane visible between conules. Ectosome and subectosome. With irregular plumose brushes of subectosomal auxiliary styles protruding from end of choanosomal primary skeletal columns. Choanosome. Skeletal architecture plumosereticulate, with differentiated primary and secondary fibre systems; primary fibres ascending, multispicular, diverging near surface, cored by erect choanosomal principal styles with points of spicules directed upwards and outwards; skeletal columns dominated by spicules with poor spon- gin; secondary fibres uni- or paucispicular, interconnecting adjacent primary tracts, forming irregular or triangular meshes up to 350Rm diameter; echinating acanthostyles protruding at acute angles from fibres, or clumped in plumose brushes, scattered over the whole length of primary tracts although more sparse near surface; mesohyl moderately heavy, lightly pigmented, granular. Megascleres. Choanosomal principal styles long, thick, straight or slightly curved at midsection, tapering to long fusiform points, most commonly with rounded smooth bases or occasionally with microspined subtylote bases. Length 372(606.1)-810p,m, width 11-(15.8)-21Rm. Subectosomal auxiliary styles shorter, more slender than principals, straight, with fusiforrn points and microspined subtylote bases. Length 352-(480.8)-590Rm, width 3-(7.6)-10p.m. Echinating acanthostyles in two size categories. Larger form intermediate between the smaller and the principal styles, slightly curved at midsection, fusiform pointed, subtylote, with heavily spined bases, lightly spined shafts and often with bare base. Length 2194293.0)384iim, width 10-(12.3)-15Rm. Smaller acanthostyles straight, slender, subtylote, fusiform, more-or-less evenly spined except usually for an aspinose point. Length 92-(148.4)-183p.m, width Microscleres. Chelae absent. Toxas of two forms: Accolada toxas, long, thin, with long straight arms, sharply angular central curve or spirally twisted centre, and non-reflexed points. Length 934139.5)-18511m, width 0.8(0.9)-1.511m. Wing-shaped toxas, short, thick, gradually curved at centre, curved arms, slightly reflexed points. Length 31-(45.5)-52Rm, width REMARKS. Koltun (1976) suggested that C. pauper is synonymous with Stylostichon toxiferum Topsent and Microciona basispinosa Burton, but respective types indicate that they differ substantially in a number of significant features, considered here to warrant recognition of C. pauper as a distinct species (see remarks for C. (Microciona) antarctica below). Clathria pauper differs from other species by a combination of characters (plumose-reticulate skeletal structure with differentiated primary and secondary fibre systems; principal spicules protruding through fibres in `spicate' arrangement; two sizes of acanthostyles echinating fibres; two toxa morphologies; and absence of chelae). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 144zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A FIG. 62. Clathria (Clathria) pauper Brondsted (fragment of holotype BMNH1930.11.5.2). A, Choanosomal principal style. B, Subectosomal auxiliary style. C, Young auxiliary style. D, Larger category of echinating acanthostyles. E, Smaller form of echinating acanthostyles. F, Short wing-shaped toxas and larger accolada toxas. G, Known Antarctic distribution. H, Section through peripheral skeleton. I, Fragment of holotype. ^ 145 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR FIG. 63. Clathria (Clathria) pauper Brondsted (fragment of holotype BMNH1930.11.5.2). A, Choanosomal skeleton. B, Ectosomal skeleton. C, Peripheral skeletal structure. D, Fibre characteristics. E, Choanosomal principal subtylostyle. F, Base of principal subtylostyle. G, 2 sizes of echinating acanthostyles. H, Spined base of acanthostyle. I, Acanthostyle spine morphology. J, Accolada toxa. K, Wing-shaped toxa. MEMOIRS OF THE QUEENSLAND MUSEUM 146zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ A FIG. 64. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hr ia ( Clat hr ia) pet for at a (Lendenfeld) (lectotype AMMG9010). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary style. C, Echinating acanthostyle. D, Palmate isochelae. E, Section through peripheral skeleton. F, Distribution. G, Skeletal structure. H. Surviving portion of type specimen. Clathria (Clathria) perforata (Lendenfeld, 1887) (Fig. 64) in part, Lendenfeld, 1887b: 788; in part, Lendenfeld, 1888: 89-90. (Not Lendenfeld, 1887b: p1.22, fig.44.) Ant herochalina perforat a, Whitelegge, 1902b: 275, 279, 287; Hooper & Wiedenmayer, 1994: 260. Clat hria perforat a; MATERIAL. LECTOTYPE: AMG9010: Port Phillip, Vic, 38°09'S, 144°52'E, other details unknown (specimen label reads `Antherochalina perforata REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 147 Lend.', whereas AM register reads `Suberapsamma philippi Lend., Port Phillip'). (Not SYNTYPE BMNH1886.8.27.459: Broughton I., Port Stephens, NSW, 32°36'S, 152°19'E). forata could also be assigned to Pandaros or Echinoclathria, whereas the few vestigial acanthostyles seen in the lectotype suggest that Clathria (Clathria) is more appropriate. HABITAT DISTRIBUTION. Ecology unknown; SE. coast of Australia (Vic, NSW) (Fig. 64F). Clathria (Clathria) piniformis (Carter, 1885) DESCRIPTION. Shape. Irregularly flabello- digitate, thin, with a small stalk. Colour. Live colouration unknown, dark brown preserved. Oscules. Unknown. Texture and surface characteristics. Surface arenaceaous, porous; texture harsh.. Ectosome and subectosome. Ectosome fibrous, arenacious, little remaining of original dermal skeleton. Choanosome. Choanosomal skeleton irregularly isodictyal reticulate, with relatively heavy spon- gin fibres forming oval or hexagonal meshes, cored by one or few vestigial choanosomal principal styles; echinating acanthostyles very scarse, scattered evenly throughout skeleton; mesohyl matrix contains heavy deposits of siliceous detritus, especially large sand grains, with heavily pigmented and granular mesohyl matrix. Megascleres. Choanosomal principal subtylostyles thin, fusiform, straight, with slightly subtylote, smooth bases. Length 96-(105.0)-11811m, width 4-(4.8)-6p.m. Subectosomal auxiliary styles barely differentiated from principal megascleres, usually much thinner, only seen scattered between fibres, few at surface. Length 854103.3)-14611m, width I - (2.7) 411m. - Acanthostyles small, slightly subtylote, with vestigial or granular spination. Length 46457.7)72p.m, width 2.5-(3.1)-4.5p.m. Microscleres. Palmate isochelae rare. Length 8(9.7)-11[Lm. Toxas absent. REMARKS. Lendenfeld (1887) originally nominated two `syntypes', but one (BMNH) (Lendenfeld, 1887b: p1.22, fig.44) is Antho (see A. (Isopenectya)chartacea), whereas the description was based on the other AM specimen (Lendenfeld, 1887b: 788, 1888: 89). This latter specimen is designated lectotype, but is in poor condition and the species is poorly known: ectosomal features cannot be clearly discerned; most megascleres within fibres are broken; and microscleres are extremely difficult to pick out between the abundant detritus within the mesohyl. In having flattened fibres which form irregular (isodictyal) anastomoses, C. (C.) per- (Figs 65-66, Table 11) Dictyocylindrus piniformis Carter, 1885f: 354. Clathria piniformis; Dendy, 1896: 34; Hooper & Wiedenmayer, 1994: 261. Wilsonella piniformis; Hallmann, 1912: 241. Paradoryx piniformis; Hallmann, 1920: 768. MATERIAL. HOLOTYPE: BMNH1886.12.15.62 (fragments MNHNDCL60, AMG2803): Port Phillip Heads, Vic, 38°17'S, 144°39'E, 38-40m depth, coll. J.B. Wilson (dredge). OTHER MATERIAL: VIC NMVRN412-508. HABITAT DISTRIBUTION. 38-40m depth; substrate unknown; Port Phillip Bay(Fig. 65G). DESCRIPTION. Shape. Massive, subspherical lobate or lobo-digitate growth form, 110-130mm long, 55-65mm wide, with thick cylindrical stalk, 45mm long, 18-22mm diameter. Colour Bright orange-brown alive, olive-brown preserved. Oscules. Large oscules, 2-4mm diameter dispersed over both faces of digitate lobes. Texture and surface characteristics. Surface rugose, with irregularly distributed conules, and a skin-like dermis. Ectosoine and subectosome. Ectosome membraneous, with tangential and paratangential bundles of subectosomal auxiliary styles, sometimes forming quite thick, sinuous tracts. Choanosome. Choanosomal skeleton reticulate; spongin fibres well developed, thin, forming sinuous tracts, branching and splitting, and producing oval or elongate meshes; spongin fibres without clear differentiation between primary or secondary elements, although thicker fibres, usually ascending and cored by principal styles, whereas connecting fibres usually aspiculose; spongin fibres cored with paucispicular tracts of vestigial choanosomal principal styles, and echinated by small acanthostyles in heavy concentrations and relatively evenly dispersed; mesohyl matrix heavy, with numerous extra-fibre spicules, especially acanthostyles and auxiliary styles in the peripheral skeleton. Megascleres (Table 11). Choanosomal principal subtylostyles very thin, vestigial, with blackened axial canals, fusiform, usually straight, some- MEMOIRS OF THE QUEENSLAND MUSEUM 148^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA LO C FIG. 65. Clathria (Clathria) piniformis (Carter) (holotype BMNH1886.12.15.62). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary style. C, Echinating acanthostyle and modified form. D, Raphidiform toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. G, Holotype. I, Specimen NMVRN412. J, Choanosomal skeleton. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 149 FIG. 66. Clathria (Clathria) piniformis (Carter) (holotype BMNH1886.12.15.62). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Modified acanthostyle. G, Modified acanthostyle spination. H, Base of principal style. 1, Principal style. MEMOIRS OF THE QUEENSLAND MUSEUM 150zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ times slightly curved, with rounded, tapering or more frequently quasidiactinal, smooth bases. Subectosomal auxiliary styles almost identical to principal styles, longer, thin, slightly curved or sinuous, fusiform, with basal endings similar to principal megascleres. Acanthostyles slightly subtylote with vestigial, evenly dispersed spines, include quasidiactinal modifications to base. Microscleres (Table 11). Palmate isochelae rare, unmodified. Toxas rhaphidiform, uncommon, with slight central curvature and reflexed points, sometimes sinuous. REMARKS. In vestigial spicule geometry and generally reduced skeletal structures C. (C.) pintformis is most similar to C. (C.) oxyphila, both of which have most of their megascleres modified to quasidiactinal forms (see remarks for C. oxyphila), but the present species has quite different microsclere geometry, skeletal construction and growth form. Clathria (Clathria) raphanus (Lamarck, 1813) (Figs 67-68) Spongia raphanus Lamarck, 1813: 444; 1814: 373. Clathria raphanus; Topsent, 1932: 100, p1.4, fig.9; Hooper & Wiedenmayer, 1994: 261. Clathria raphana; Van Soest, 1984b: 108. Thalyseurypon raphanus; de Laubenfels, 1936a: 107. Panda ros raphanus; Wiedenmayer, 1977: 143-4. MATERIAL. HOLOTYPE: MNHNDT572(dry): Australian seas, Peron & Lesueur collection, no other details known. HABITAT DISTRIBUTION. Ecology unknown; Australia. DESCRIPTION. Shape. Arborescent fan, 90mm high, 95mm wide, with flattened multiplanar branches 48-80mm long, 5-10mm thick, which anastomose tightly to form several lobes. Colour Grey-brown dry. Oscules. Large oscules, up to 3.5rnm diameter, scattered over both faces of flattened branches. Texture and surface characteristics. Surface microconulose with tapering processes; texture harsh in dry state. Ectosome and subectosome. Ectosomal skeleton a sparse tangential or paratangential layer of subectosomal auxiliary styles. Choanosome. Choanosomal skeleton almost regularly reticulate, with heavy spongin fibres forming oval or elongate meshes, without differentiation of primary or secondary elements; TABLE 11. Comparison between present and published records of Clathria (Clathria) piniformis (Carter). Measurements in pin, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Holotype (BWINH- 1886.12.15.62) Specimen (N=1) Choanosomal principal styles 124-(137.6)-156 x 1.5-(2.2.)-3 Subectosomal auxiliary styles 147-(164.4)-182 x 2.5-(3.1)-4 128-(144.4)-154 x 2(2.6)4 149-(178.3)-197 x 2(3.9)-6 Echinating acanthostyles 51-(57.1)-64 x 2.5(3.2)-4 49-(54.2)-58 x 2(3.3)-6 Chelae 9-(10.2)-12 10-(10.8)-12 Toxas 92-(117.2)-143 x 0.5-(0.7)-1 1104122.0)-135 x 0.540.7)-0.8 spongin fibres almost totally uncored, and where present, choanosomal principal styles form unior paucispicular tracts within axis of fibres; spongin fibres sparsely echinated by acanthostyles; mesohyl light (dried material). Megascleres. Choanosomal principal subtylostyles fusiform, slightly curved, with rounded or slightly subtylote, smooth bases. Length 108(148.6)-230p.m, width 3.5-(5.6)-9p.m. Subectosomal auxiliary subtylostyles straight or slightly curved, thin, fusiform, with subtylote, smooth bases. Length 168-(225.7)-315p,m, width 1.5-(2.6)-4tim. Acanthostyles curved, slightly subtylote, with vestigial, granular, evenly dispersed spines. Length 47-(65.2)-82p,m, width 2.5-(4.1)-51.tm. Microscleres. Absent. REMARKS. This species is known only from an antiquated dried specimen collected from 'somewhere in the vicinity of Australia' (Topsent, 1932). It was made the type species of Thalyseurypon de Laubenfels and has since been the subject of lengthy discussions by de Laubenfels (1936a), Hechtel (1965), Wiedenmayer (1977), Van Soest (1984b) and Fromont & Bergquist (1990). These discussions have centred mostly on the value of the loss of meniscoid (chelae) microscleres used as a valid taxonomic character. These arguments have been addressed in the synopsis of Thalyseurypon (see Included Genera, above). Clathria raphanus bears little resemblance to Pandaros as supposed by Wiedenmayer (1977) which has flattened fibres, or Echinoclathria (as interpreted by Wiedenmayer, 1989) which has smooth echinating megascleres. This species is most appropriately placed in Clathria (Clathria) based on the distribution and geometry of its megascleres. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 151 Clathria (C.) raphanus is depauperate in morphological features, with a reduced spicule diversity and vestigial spicule development, and a relatively heavy evenly reticulate fibre skeleton, and in this respect the species is associated with a species-group (the oxyphila' group; also containing C. (C.) pinifonnis, C. (C.) oxyphila and C. (Dendrocia) elegantula). Clathria (Clathria) rubens (Lendenfeld, 1888) (Figs 69-70, Table 12) Thalassodendron rubens Lendenfeld, 1888: 223. Not Thalassodendron rubens var. dura, in part, Lendenfeld, 1888: 223-224. Not Thalassodendron rubens var. lamella, in part, Lendenfeld, 1888: 224, p1.7. Clathria rubens, in part; Whitelegge, 1901: 85-86, p1.11, fig.13. Clathria rubens; Hallmann, 1912: 218-223, p1.32, fig.1, text-fig.45; Burton, 1934a: 558; Burton, 1938b: 20; Shaw, 1927: 425-426; Guiler, 1950: 7; Hooper & Wiedenmayer, 1994: 261. Clathria tenuifibra Whitelegge, 1901: 82-83, p1.11, fig.10; Whitelegge, 1902b: 274, 279, 287; Hallmann, 1912: 211. Thalysias tenuifibra; de Laubenfels, 1936a: 105. Placochalina pedunculata var. monis, in part; Lendenfeld, 1888: 91-92; Whitelegge, 1902b: 274. cf. Microciona prolifera; Vosmaer, 1935a: 611, 644, 668. Not Microciona rubens Bergquist, 1961a: 38. MATERIAL. LECTOTYPE: AMG9119 (dry): Port Jackson, NSW, 33°51'S, 151°16'E, no other details known (label `Thalassodendron rubens RvL, type'). PARALECTOTYPES: AMZ455 (slide AMG3585: same locality (dry, label 'Clathria rubens, type or cotype, Lendenfeld's duplicate A50'). AMZ634: same locality; (dry, label `Thalassodendron rubens RvL, type or cotype, Lendenfeld's no.240'). BMNH1887.1.24.35 (fragment AMG3586): same locality, Ramsay collection (listed as 'var. intermedia MS name'). HOLOTYPE of C. tenuifibra: AMG3045(dry): Lake Illawarra region, NSW, 34°32'S, 150°50'E (label `Thalysias tenuifibra Whitelegge; type'). OTHER MATERIAL: NSW QMG300452. HABITAT DISTRIBUTION. Shallower than 56m depth; on shell, gravel and Halimeda substrates; Port Jackson (Lendenfeld, 1888), Broken Bay, Lake IIlawarra, and Woolongong (NSW) (Whitelegge, 1901; Hallmann, 1912); Direction and Eagle Is, Great Barrier Reef (FNQ) (Burton, 1934a); Maria I. (Tas) (Shaw, 1927; Guiler, 1950); St Vincent Gulf (SA) (Hallmann, 1912) (Fig. 69G). DESCRIPTION. Shape. Arboresecent digitate sponge,100-190mm high, 60-110mm wide, with numerous anastomosing, repeatedly bifurcate, stoloniferous branches, 30-75mm long, up to 19mm diameter, without definite basal stalk; branches subcylindrical, slightly flattened at ends. Colour Grey-brown in ethanol. Oscules. Large oscules, 2-4mm diameter, scattered over surface of branches, often raised on a small lip. Texture and surface characteristics. Surface even, optically smooth, but with prominent subdermal grooves and stellate drainage canals radiating from a central osculum (not seen in dry material). Ectosome and subectosome. Ectosomal region membraneous, with a sparse tangential layer of subectosomal auxiliary styles, through which protrude sparse brushes of choanosomal principal styles; subectosomal skeleton has auxiliary styles dispersed paratangentially, and peripheral choansomal fibres diverging slightly just below surface. Choanosome. Choanosomal skeletal architecture subisodictyal, with triangular or irregular meshes and thin spongin fibres, cored by uni-, bi- or paucispicular tracts of choanosomal principal styles; spongin fibres imperfectly divided into axial and extra-axial components, differing only in fibre diameter; axial fibres comparatively thick, paucispicular, forming cavernous and irregular ovoid meshes; extra-axial skeletal meshes more obviously triangular, subisodictyal, with thin, paucispicular ascending primary spongin fibres interconnected by uni- or bispicular secondary fibres; echinating acanthostyles lightly and evenly dispersed over fibres; mesohyl matrix heavier in peripheral skeleton than at core, containing few principal and auxiliary styles and microscleres. Megascleres (Table 12). Choanosomal principal styles thick, mostly straight, hastate-fusiform, with rounded, tapering or subtylote bases, usually smooth, occasionally microspined. Subectosomal auxiliary subtylostyles longer than principal styles, thin, straight, fusiform, with slightly subtylote, microspined bases. Acanthostyles subtylote, with mostly granular, evenly dispersed spines, sometimes with aspinose necks, and some clearly intermediate forms of choanosomal styles. Microscleres (refer to Table 12 for dimensions). Palmate isochelae rare, small, unmodified. Toxas accolada, thin, long, usually with very large, angular, sometimes coiled, central curves, slightly reflexed points, or they may be nearly straight with coiled central flexion. MEMOIRS OF THE QUEENSLAND MUSEUM 152zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ (C.) partita), geometry of choanosomal styles and toxas (Figs 59, 69), and having similar growth forms (although branches are cylindrical in C. (C.) rubens but flattened in C. (C.) part ita). Nevertheless, spicule geometries, fibre characteristics and spicule dimensions indicate that they are probably separate sibling species. There are 3 other AM specimens which have been associated with C. (C.) rubens. One, AMZ4810 from Port Jackson (with label `Thalassodendron rubens, donated by A. Dendy'), is probably a fragment of the BMNH paralectotype, but this has not been verified. AMZ2241 from Tasmania (apparently identified by M. Burton, with an old label Rhaphidophlus typicus, dried up in 1917-18'), and AMZ2246 (locality unknown, also identified by M. Burton, with an old label which reads `Crella incrustans'), were both obtained from the Antarctic Publications Committee (donated by Prof. Haswell). All three are in too poor condition to identify reliably, but they appear related to C. (Thalysias) cactifonnis. The origin of the QM specimen, obviously of great antiquity, is not known (L. FIG. 67. Clathria (Clathria) raphanus (Lamarck) (holotype MNHNDT572). Cannon, pers. comm.), but it A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary sub- may have been an exchange tylostyle. C, Echinating acanthostyle. D, Section through peripheral specimen from the AM during skeleton. E, Holotype. Whitelegge's era. That specimen certainly belongs to REMARKS. Clathria (C.) rubens is very similar C. (C.) rubens. to C. (C.)partita in skeletal architecture and fibre The holotype of C. tenuifibra is also in poor characteristics. Both species have a more-or-less condition, and irrespective of skeletal reconstitudifferentiated axial and extra-axial choanosomal skeleton. Whereas the present species has an ir- tion following rehydration during sectioning only regularly subisodictyal reticulate skeleton with few features were discernible. What little can be ovoid meshes and heavy fibres, C. (C.)partita has seen of the skeletal structure is identical to C. (C.) a nearly regularlsociella-like isodictyal skeleton. These two species are also similar in the extent to rubens, particularly the fibre characteristics and which axial and extra-axial skeletons are spiculation, and on this basis the two species are separated (with more marked separation in C. merged here. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 153 FIG. 68. Clathria (Clathria) raphanus (Lamarck) (holotype MNHNDT572). A, Choanosomal skeleton. B, Fibre characteristics. C, Membraneous ectosomal region. D, Echinating acanthostyles. E, Vestigial acanthostyle spines. Clathria (Clathria) squalorum Wiedenmayer in Hooper & Wiedenmayer, 1994 (Figs 71-72) Clathria squalorum Wiedenmayer, in Hooper & Wiedenmayer, 1994: 261. Clathria dura var. moths Hentschel, 1911: 370-372, text-fig.45; Hallmann, 1912: 242. Not Clathria dura Whitelegge, 1901: 83. Not Clathria mollis Kirkpatrick, 1903: 249. MATERIAL. LECTOTYPE: ZMB4444: Freycinet Reach, Shark Bay region, WA, 26°05'S, 113°30'E, 3.5-11m depth, 5.ix.1905, coll. W. Michaelsen & R. Hartmeyer (dredge). PA RA LECTOTYPES: HM: same locality (9 dry specimens). HM: Geographe Bay, Bunbury region, WA, 33°35'S, 115°26'E (5 dry specimens). HABITAT DISTRIBUTION. 3.5-11m depth; on sand, stone, mud and algal bed substrates; Bunbury-Shark Bay region (WA) (Fig. 71G). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 154zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 69. Clathria (Clathria) rubens (Lendenfeld) (lectotype AMZ9119). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Palmate isochela. E, Accolada toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 155 Clat hria ( Clat hria) rubens (Lendenfeld) (A-B, NTMZ1527; C-G, QMG300452). A, Choanosomal FIG. 70. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA skeleton. B, Fibre characteristics (x313). C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of auxiliary subtylostyle. F, Palmate isochelae. G, Accolada toxas. MEMOIRS OF THE QUEENSLAND MUSEUM 156zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 12. Comparison between present and published records of bases. Length 854148.5)Clathria (Clathria) rubens (Lendenfeld). All measurements are given in 184p.m, width 242.8)-411,m. p.m, denoted as range (and mean) of spicule length x spicule width Acanthostyles slightly sub(N=25). tylote, with evenly dispersed Lectotype (AMG9119) Paralectotypes (N=3) relatively large spines, slightly Holotype of C.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Specimen (N=I) less spinose below basal region. t enuij ibra Length 48-(62.3)-7411m, width (AMG3045) Choanosomal principal styles 96-(135.3)-162 x 5-(9.0)-I I 128-(135.6)-147 x 5.5-(7.1)-9 1124139.2)-159 x 8-(9.4)-11 1184140.6)-155 x 6-(7.8)-10 Subectosomal auxiliary styles 106-(157.2)-212 x 1.5-(2.9)-4 1324146.2)-168 x 2.5-(2.9)-4 112-(153.7)-185 x 2.543.7)-4.5 105-(139.2)-165 x 2.5-(3.3)-4.5 Echinating acanthostyles 55-(71.8)-111 x 3-(5.6)-7 58-(77.3)-112 x 4.5-(5.6)-6.5 51-(68.6)-118 x 5-(6.4)-8 48-(69.4)-I08 x 4-(5.8)-7.5 SPICULE Chelae 6-(9.6)-I3 7-(9.8)-13 8-(9.7)-11 6-(9.0)-12 Toxas 101-(153.4)-215 x 0.8-(1.3)-1.5 115-(161.6)-208 x I-(1.4)-2 94-(126.3)-153 x I-(1.2)-1.5 114-(138.6)-194 x 1-(1.4)-2 DESCRIPTION. Shape. Variable growth form, ranging from thickly encrusting with small stoloniferous digits, to digitate planar arborescent with anastomosing branches. Colour Live colouration unknown, beige preserved. Oscules. Large oscules on apex and lateral margins of stoloniferous branches Texture and surface characteristics. Surface microconulose. Ectosome and subectosome. Ectosomal skeleton hispid, with points of choanosomal principal styles protruding through surface individually or in small bundles, piercing a sparse paratangential layer of subectosomal auxiliary subtylostyles. Choanosome. Choanosomal skeletal architecture wide-meshed nearly renieroid reticulate, more regular in peripheral skeleton than in axis, with differentiated primary (ascending, multispicular) and secondary (transverse, pauci- or unispicular) spongin fibres; fibre diameter generally small (primaries 30-501..im; secondaries 10-22p,m), fibre anastomoses rectangular, and fibres cored by choanosomal principal styles; coring spicules in axial region heavier, more plumose (protruding through fibres) than in peripheral skeleton; echinating acanthostyles relatively sparse, evenly distributed throughout skeleton; mesohyl matrix light, usually containing choanosomal and/or subectosomal megascleres dispersed between fibres. Megascleres. Choanosomal principal subtylostyles slightly curved, fusiform, entirely smooth, very slightly subtylote. Length 124-(152.4)165Rm, width 4-(8.8)-12Rm. Subectosomal auxiliary subtylostyles straight, thin, fusiform, with smooth slightly subtylote 2-(4.2)-61.m. Microscleres. Palmate isochelae unmodified. Length 448.6)12Rm long. Accolada toxas very thin, straight arms, sharply angular central curvature, occasionally raphidiform. Length 105220v,m, width 1-(2.2)-311,m. REMARKS. Hallmann (1912) noted that C. dura var. mollis Hentschel was distinctly different from C. dura Whitelegge and he considered that Hentschel's (1911) species required a new name. The subspecific name 'moths' is preoccupied by C. monis Kirkpatrick, 1903, and so a new name C. squalorwn was proposed by Wiedenmayer (in Hooper & Wiedenmayer, 1994). Some of the characters and measurements of the lectotype differ from those published by Hentschel (1911). His description was presumably based on a series of syntypes, but these have not been located or re-examined. Comparison between the type material of C. (C.) squalorum and C. (Dendrocia) dura show that both species are different in several significant respects: (1) differences in the geometry of structural spicules where C. (C.) squalorum has differentiated principal and auxiliary spicules (and hence is referrable to C. (Clathria)), and C. (D.) dura has completely undifferentiated structural megascleres (and hence is referrable to C. (Dendrocia)); (2) palmate isochelae versus arcuate-like isochelae; (3) long thick accolada toxas versus no toxas; (4) nearly renieroid reticulation of thin fibres versus a regularly reticulate skeleton with heavy fibres, respectively. In having a slightly renieroid skeletal architecture C. (C.) squalorum is reminiscent of C. (lsociella) eccentrica, although spicule geometry and spicule dimensions are otherwise different. Clathria (Clathria) striata Whitelegge, 1907 (Figs 73-74, Plate 1F, Table 13) Clathria striata Whitelegge, 1907: 495-496, p1.45, fig.27; Hooper, & Wiedenmayer, 1994: 261. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 157 Thalysias striata; de Laubenfels, 1936a: 105. Rhaphidophlus tenebratus Whitelegge, 1907: 501- 503, p1.45, fig.I9. Clathria tenebrata; Hallmann, 1912: 211. cf. Spongia sartaginula Lamarck, 1813: 383; 1814: 362. MATERIAL. LECTOTYPE: AMG4344 (in part): Near Coogee, Sydney, NSW, 34°05'S, 151°10'E, coll. Hy 'Thetis' (dredge; label 'Clathria striata; type'). PARALECTOTYPES: AMG4344 (in part), Z823 (in part): Off Botany Bay, NSW, 34°00'S, 151°11'E, 4046m depth, coll. FIV 'Thetis' (dredge; dry, two specimens, label `Thalysias striata; cotypes'). HOLOTYPE of R. tenebratus: AMG4336: Off Woolongong, NSW, 34°30'S, 150°50'E, coll. FIV 'Thetis' (dredge; dry, label 'Clathria tenebrata Whitelegge; type'). OTHER MATERIAL: NSW QMG303755. HABITAT DISTRIBUTION. 40-50m depth; on rock substrate; central and S. coasts (NSW) (Fig. 73G). DESCRIPTION. Shape. Flabellate, digitate or a combination of both, 170-285mm long, 30110mm wide, with short, tapering, cylindrical stalk, 35-55mm long, 6-13mm diameter, cylindrical or flattened branches and even, slightly undulating, ragged margins. Colour. Live colouration unknown, dark greybrown preserved. Oscules. Not observed on flabellate specimens, but small oscules, up to 2mm diameter, seen on lateral margins of branches for digitate specimen. Texture and surface characteristics. Surface optically smooth, with prominent longitudinally radiating subdermal grooves and low ridges. Ectosome and subectosome. Ectosome strongly hispid, with discrete, plumose, or paratangential tufts of subectosomal auxiliary spicules surrounding protruding choanosomal principal styles; principal styles in peripheral skeleton usually larger than principal spicules within fibres; ectosomal skeleton relatively dense in places, merely paratangential or sometimes tangential to the surface in other places; thick tracts of palmate isochelae mostly confined to dermal and subdermal regions; subectosomal region plumose, with diverging tracts of choanosomal principal styles intermingled with extra-fibre tracts of auxilairy styles, together producing relatively dense peripheral skeleton. Choanosome. Choanosomal skeleton predominantly renieroid but with differentiated plumose and renieroid-reticulate components, and clearly differentiated axial and extra-axial regions but no well-marked separation of primary or secondary fibre elements (cf. Whitelegge, 1907); axial skeleton with very heavy, thick spongin fibres forming relatively tight ovoid meshes, with paucispicular core of choanosomal styles forming vaguely ascending skeletal tracts; extra-axial region with much heavier, rectangular, almost renieroid spicule skeleton, with a criss-cross of longitudinal and transversely orientated spicules, and spongin fibres lighter than in axis; longitudinal spongin fibres in periphery cored by paucispicular tracts of principal styles; transverse/ascending tracts with multispicular tracts of spicules, many protruding through fibres and forming plumose brushes at right angles to surface; tendency for some larger principal styles to form ascending tracts, and small styles to occur mainly in longitudinal tracts; ultimate choanosomal spicule tracts diverge into subectosomal region; echinating acanthostyles relatively sparse in axis, only marginally more abundant in peripheral skeleton; mesohyl matrix heavy, granular, with few megascleres between fibres. Megascleres (Table 13). Choanosomal principal styles thick, curved, relatively variable in length, fusiform, mostly sharply-pointed, less frequently with rounded points, with smooth rounded, or occasionally very slightly subtylote bases. Subectosomal auxiliary subtylostyles thin, fusiform, straight or slightly curved, occasionally sinuous, usually with smooth, sometimes microspined, slightly subtylote bases, occasionally polytylote bases. Acanthostyles subtylote, with small and evenly distributed spines. Microscleres (Table 13). Palmate isochelae abundant, unmodified. Toxas accolada, long, moderately thick, with sharply angular or slightly angular central curvature, straight unreflexed points. REMARKS. Rhaphidophlus tenebratus is a synonym of Clathria striata, both species having identical skeletal architecture, fibre characteristics and spicule geometry, although there is some variation in spicule dimensions between these two nominal species (Table 13). They differ only in growth form (C. striata being flabellate, R. tenebratus being digitate, with flattened and anastomosing branches; Fig. 731). In his original description Whitelegge (1907) omitted to mention toxas, abundant in the preserved specimens but relatively rare in the dry holotype, and also. that echinating acanthostyles were 0.6-0.8mm long (typographical error for 0.06-0.08mm). 158zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 71. Clathria (Clathria) squalorum Wiedenmayer (holotype ZMB4444). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. H, Skeletal structure. I, Subrenieroid reticulate fibres. Clathria (C.) striata is similar to C. (C.) sartaginula (Lamarck) in shape, slightly renieroid skeletal architecture, and to some extent spiculation. However, fibre characteristics differ markedly between the two species, particularly the density of coring and echinating spicules on fibres, the degree to which peripheral fibres radiate from the skeletal axis, and the very small but thick, stumpy choanosomal styles in C. (C.) sartaginula. This species is also compared with C. (C.) arcuophora and C. (C.) biclathrata in spicule geometry, spicule dimensions, and to ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 159 FIG. 72. Clathria (Clathria) squalorum Wiedenmayer (holotype ZMB4444). A, Choanosomal skeleton. B, Ectosomal skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Base of auxiliary subtylostyle. G, Accolada toxa. H, Palmate isochelae. some extent fibre characteristics. Clathria (C.) striata also has a similar skeletal architecture as C. (C.) arcuophora, differing substantially only in toxa geometry. All these species (C. (C.) striata, C. (C.) arruophora, C. (C.) sartaginula, C. (C.) crassa and C. (C.) biclathrata), are characterised by their slightly renieroid or subrenieroid choanosomal skeletons and spongin fibres cored by few, thick principal spicules. They are grouped together here in the striata' species 160zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 73. Clathria (Clathria) striata Whitelegge (lectotype AMZ4344). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Holotype of R. tenebratus AMG4336. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 161 FIG. 74. Clathria (Clathria) striata Whitelegge (paralectotype AMZ823). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Palmate isochelae. G, Accolada toxas. H, Base of auxiliary subtylostyle. 162zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 13. Comparison between present and published records of Clathria bundles superficially resem(Clathria) striata Whitelegge. All measurements are given in pm, denoted bling a Pseudaxinella (Axinelas range (and mean) of spicule length x spicule width (N=25). lidae). Colour. Pale yellow-brown Lectotype (AMG4344) Paralectotypes (N=2) Subectosomal auxiliary styles 1684268.3)-523 x 12-(17.7)-28 142-(245.6)-363 x 3-(4. 1)-6 184-(285.2)-496 x 14-(19)-24 153-(229.2)-294 x 3-(4.2)-6 Echinating acanthostyles 71-(81.8)-93 x 6-(7.3)-9 75-(86)-94 x 7-(7.6)-9 SPICULE Choanosomal principal styles Holotype of R.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB preserved. QMG303755 t enebrat us Oscules. Not seen. (AMG4336) Texture and surface charac193-(282.2)-546 2094299.7)-462 Texture harsh, comteristics. x 14-(16.5)-22 x 15-(19.1)-23 pressible; surface prominently 175-(274.4)-387 132-(181.4)-273 conulose, hispid. x 2-(2.9)-5 x 3.5-(5.6)-8 Ectosome and subectosome. Surface skeleton with protruding single principal styles em13-(15.1)-17 14-(16.4)-19 I4-(16.4)-20 14-(16.5)-21 Chelae bedded in peripheral skeleton 165-(224.3)-265 110-(211.0)-255 I 08-(202.4)-305 163-(221.8)-325 Toxas and forming a plumose hispid x 0.841.7)-3.0 x 0.8-(1.4)-2 x 1-(1.3)-2 x 0.8-(1.4)-2 surface; thick bundles of both subectosomal auxiliary styles group. Clathria (C.) striata differs from other and toxa microscleres surrounding protruding striata' species in growth form, spicule principal spicules, and also scattered paratangengeometry and spicule dimensions. De Laubenfels (1936a) referred C. striata to tially across surface. Thalysias but his decision is not supported. Al- Choanosonze. Skeletal architecture plumo-reticuthough there are relatively dense plumose ec- late, with slightly differentiated axial and extratosomal brushes composed of auxiliary axial regions; in peripheral region skeleton more megascleres (structurally similar to Clathria plumose than reticulate, with skeletal columns (Thalysias)) there is only one undifferentiated diverging but without connecting elements; size category (142-38711m long) (composition of skeletal columns composed of erect multispicular bundles of light spongin fibres fully cored by Clathria (Clathria)). choanosomal principal styles; spicules mostly contained within fibres but also slightly protrudClathria (Clathria) toxipraedita Topsent, 1913 ing through fibres producing the plumose (Figs 75-76) ('spicate) arrangement; towards base skeleton more disorganised, predominantly reticulate, and Clathria toxipraedita Topsent, 1913a: 620-621, p1.5, spicules more-or-less completely contained fig.4, p1.6, fig.12; Burton, 1932a: 319; Burton, 1934b: 32, p1.4, figs 2-3, text-fig.3; Koltun, 1964a: within fibres; echinating acanthostyles moderate68-69, p1.12, figs 15-24; Koltun, 1976: 187; Hooper ly abundant; mesohyl matrix heavy, smooth, containing abundant microscleres. & Wiedenmayer, 1994: 262. Thalysias toxipraedita; de Laubenfels, 1936a: 105. Megascleres. Choanosomal principal styles long, Rhaphidophlus toxipraedita; Van Soest, 1984b: 115. robust, straight or slightly curved near base, with cf. Clathria toxipraedita; Sim & Byeon, 1989: 38 fusiform points and rounded or slightly subtylote (Korea; possible misidentification). smooth bases. Length 5184616.2)-894 width 19-(27.7)-33Rm. MATERIAL. HOLOTYPE: RSME1921.143.1400: BurSubectosomal auxiliary subtylostyles short, wood Bank, off Tierra del Fuego, S. Atlantic, 54°25'S, 57°32'W, 112m depth, 1.xii.1903, coll. R.R.V. very slender, straight, hastate pointed, subtylote, with pointed (mucronate) or minutely spined 'Scotia' (dredge). bases. Length 278-(310.2)-343p.m, width 4HABITAT DISTRIBUTION. 93-540m depth; on mud, (4.8)-61.1,m. sand and hard substrates; Australian Antarctic TerEchinating acanthostyles small, straight, long, ritory: MacRobertson Land (Koltun, 1976) (Fig. 7511). tapering fusiform pointed, subtylote or tylote Also Tierra del Fuego (Topsent, 1913a), South Georgia, Shag Rock (Burton, 1932a, 1934b), South bases, with small spines and aspinose bases and Sandwich Is, Falkland Is (Koltun, 1964a), Palmer Ar- points. Length 102499.2)-1541m, width 4-(7.8)11 vm. chipelago, Antarctica (Burton, 1932a). Microscleres. Palmate isochelae in two size DESCRIPTION. Shape. Massive encrusting, categories, both modified. Smaller chelae with subspherical, 70mm long, 60mtn wide, up to relatively small alae and a small plate/ridge on 25mm thick, composed of fused irregular fibre central inner margin of shaft protruding between 84-(97.8)-112 x 6-(9.2)-11 58-(78.0)-92 x 4-(8.1)-11 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 163 the alae. Length 12-(15.7)-18Rm. Larger chelae often cleistochelate, with teeth partially or completely fused and a large central plate/ridge on inner margin of shaft between alae. Length 20(22.3)-24Rm. Two categories of toxas. Longer toxas accolada, very long, thick, with sharply angular but only slight central curvature, straight arms and smooth fusiform points. Length 63541022.6)147011m, width 4-(5.2)-6Rm. Smaller toxas wing-shaped, more widely curved at centre, occasionally looped/twisted at centre, with slightly reflexed arms and points. Length 56485.3)132p,m, width 0.5-(1.1)-2.0pLm. REMARKS. Clathria (C.) toxipraedita has peculiar, large palmate cleistochelae resembling sigmancistras (in some Mycalidae), with alae varying from partially to completely fused, and with the addition of a well formed central platelike ridge protruding between the alae (Fig. 76G). Smaller chelae have smaller diffuse alae and a smaller central plate-like ridge, indicating that they are ontogenetic stages of the larger forms. Cleistochelae are known in a number of other poecilosclerids including a several microcionids (e.g. C. (Microciona)cleistochela Topsent, C. (T) ramosa (Dendy),Antho(Plocamia)signata (Topsent)), whereas the central plate-like ridge appears to be unique to the family and only previously recorded from Mycalidae (E. Hajdu, pers.comm.). Clathria toxipraedita is unusual amongst the antiboreal species in having a great diversity of spicule geometries; generally microcionids from Antarctic waters appear to have reduced spicule diversity in comparison to the tropical fauna. Clathria (Clathria) transiens Hallmann, 1912 (Figs 77-78, Plate 2A, Table 14) Clathriatransiens Hallmann, 1912: 205, 226-234, 253, 254, p1.33, figs 1-3, p1.34, fig.2, text-figs 47-48; Shaw, 1927:426; Burton, 1934a: 599; Guiler, 1950: 7; Burton, 1959a: 244; Wiedenmayer, 1989: 57, p1.5, fig.8, p1.23, fig.6, text-fig.38; Hooper & Wiedenmayer, 1994: 262. Thalysias transiens; de Laubenfels, 1936a: 105. cf. Microciona prolifera, tropus stylota and tropus senta, Vosmaer, 1935a: 611, 649-650, 666. MATERIAL. LECTOTYPE: AME302: Off Devonport, N. coast Tas, 41°11'S, 146°21'E, coll. FIV 'Endeavour' (dredge) (label 'Clathria transiens; typical form a'). PARALECTOTYPES: AMZ49 + E819: 64km W. of Kingston, SA, 36°50'S, 139°05'E, 60m depth, coll. FIV 'Endeavour' (dredge) (specimen cut in half, form b). AME779: same locality; (form c). AMZ744: Port Phillip Bay, Vic, 38°09'S, 144°52'E, coll. J.B. Wilson (dredge) (label 'Clathria transita Hallmann', AM register ref. 342/85, JBW sp.3, cotype, form d). (uncertain type status - AMZ743: Port Phillip Bay, Vic., 38°09'S, 144°52'E, coll. J.B. Wilson; (dredge; label 'Clathria transita Hallmann', dry, `type')). OTHER MATERIAL: VIC. - NCIQ66C3231-C (fragment NTMZ3694). TAS. - QMG300268 (NCIQ66C-3638-U, fragment NTMZ3802). HABITAT DISTRIBUTION. Shallow subtida1-60m depth; on rock substrates; Port Phillip (Vic) (Hallmann, 1912; present study), Furneaux Is and Devonport (Tas) (Shaw, 1927; Guiler, 1950; Wiedenmayer, 1989; present study), Kingston (SA) (Hallmann, 1912), Qld. (Burton, 1934a) (Fig. 77G). Burton (1959a) recorded the species from the Red Sea but his material was not described, and his record is questionable. DESCRIPTION. Shape. Usually arborescent with short, bulbous branches, 75-185mm long, 40-160mm wide; stipitate with short cylindrical stalk, 25-55mm long, 8-22mm diameter; branches simply bifurcate, or repeatedly bifurcate and arborescent, flabello-digitate, or they may anastomose to form thickly clathrous, bushy lobes. Colour. Live colouration vermilion red (Munsell 5R 3/10), beige-brown or dark brown preserved. Oscules. Small oscules, up to 3mm diameter, in ridges and grooves of branches. Texture and surface characteristics. Surface markedly conulose, with conules actually being rounded vestigial branches; with or without irregular ridges striating surface; surface covered by distinct membraneous skin-like ectosome. Ectosome and subectosome. Ectosome minutely hispid, with the points of larger choanosomal principal styles protruding through membraneous surface, with a paucispicular, usually tangential layer of subectosomal auxiliary subtylostyles, lying on or just below surface; occasionally auxiliary styles form plumose brushes, surrounding 1 or more principal style; subectosomal skeletal structure vaguely plumose, formed by diverging peripheral spongin fibres and spicule tracts of smaller choanosomal principal styles. Choanosome. Choanosomal skeletal architecture almost regular, radial-renieroid, with poorly differentiated axial and extra-axial skeletons; axis formed by evenly anastomosing spongin fibres, forming relatively cavernous, rectangular or circular meshes; spongin fibres cored by pauci- or multispicular tracts of smaller choanosomal principal styles; primary and secondary skeletal tracts in axis vaguely differentiated, but not pronounced, and spongin fibre elements only dif- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 164zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A EzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 0 0 CV FIG. 75. Clathria(Clathria)toxaepraedita Topsent (holotype RSME1921.143.1400). A, Choanosomal principal styles and modifications. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada toxas. E, Wing-shaped toxas. F, Modified palmate isochela. G, Section through peripheral skeleton. H, Antarctica distribution. I, Holotype. J, Choanosomal structure. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 165 FIG. 76. Clathria(Clathria)toxaepraedita Topsent (holotype RSME1921.143.1400). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Base of auxiliary subtylostyle. E, Echinating acanthostyle. F, Acanthostyle spines. G, Transitional series from cleistochelae to modified palmate isochelae. H, Accolada and wing-shaped toxas. I, Accolada toxas. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 166zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A Clat hr ia ( Clat hr ia) t r ansiens Hallmann (lectotype AME302). A, Choanosomal principal styles. B, FIG. 77. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Oxhorn toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, lectotype. ferentiated by density of coring spicules; extraaxial skeletal architecture regularly renieroid, with ascending plumo-reticulate primary and secondary fibre elements which diverge increasingly towards surface; fibre anastomoses much closer toward periphery than at core; primary ascending fibres heavily multispicular, secondary transverse fibres more irregular, pauci- or multispicular; all fibres relatively heavily invested with spongin; echinating acanthostyles abundant on all fibres, only slightly more dense at fibre nodes; mesohyl matrix light, containing numerous subectosomal styles and microscleres scattered between fibres. Megascleres (Table 14). Choanosomal principal styles divided into 2 (overlapping) size classes: larger found in brushes protruding from surface; smaller coring choanosomal spongin fibres; both similar in geometry, fusiform, straight or slightly curved, with smooth, rounded or very slightly subtylote bases. Subectosomal auxiliary subtylostyles straight, thin, fusiform, with smooth subtylote bases, variable length but only 1 size class. Acanthostyles slightly subtylote, very sharply pointed, with granular, vestigial, evenly dispersed spines, sometimes completely smooth. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 167 FIG. 78. Clathria (Clathria) transiens Hallmann (paralectotype AMZ743). A, Choanosomal skeleton. B, Fibre characteristics (x306). C, Echinating acanthostyles. D, Vestigial acanthostyle spines. E, Echinating spicules in situ. F, Accolada toxa. G, Palmate isochelae. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 168zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Microscleres (Table 14). Palmate isochelae large, unmodified, rare in some material. Toxas oxhorn, relatively thick, with rounded, large or small central curvature, slightly reflexed or straight arms. Larvae. Larvae viviparous, ovoid or elongate parenchymella, up to 210p,m diameter, with larval megascleres dispersed throughout axis. REMARKS. The status of specimen AMZ743 (from Port Phillip Bay) is not completely certain. The label states that it is a 'dry, type', but the four type specimens described by Hallmann (1912) are accounted for by other AM register numbers (cited above). It is possible that the specimen is a fragment of the paralectotype (AMZ744), as both come from the same locality. Clathria (C.) transiens is relatively well known from SE. Australian waters (e.g. Wiedenmayer, 1989). The species appears to be the temperate water equivalent of the widely distributed tropical species C. (Thalysias) lendenfeldi (see Hooper et al., 1990). Both species have a large range of growth forms and comparable surface features, although this resemblance is superficial, in the field, and the two species differ markedly in most other features. The ectosomal features of C. (C.) transiens (with protruding choanosomal styles), and fibre characteristics (almost a subrenieroid peripheral skeleton), are also reminiscent of C. (T) vulpina, although growth form and spicule diversity differ significantly between these two species. Clathria (T) darwinensis sp.nov. from northwest Australia is similar to C. (T) transiens in ectosomal features, vaguely subrenieroid skeletal architecture, and toxa morphology, but differs in acanthostyle geometry, possession of 2 size classes of auxiliary styles, gross morphology and spicule dimensions. Despite a relatively variable growth form the skeletal architecture of this species is consistent. There are only minor differences between specimens in the degree to which the axial skelton is compressed and differentiated from extra-axial structures (e.g. compare Hallmann's (1912) 'form a' and 'form c'). Similarly, in some places on the ectosomal skeleton are distinct plumose brushes of auxiliary spicules, radiating from bases of protruding choanosomal principal styles, whereas in other places ectosomal structure is simply tangential. This plumose structure is generally associated with the ectosomal skeleton characteristic of Clathria (Thalysias) species, but only a single size class of auxiliary styles is TABLE 14. Comparison between present and published records of Clathria (Clathria) transiens Hallmann. All measurements are given in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Lectotype (AM E302) Paralectotypes (N=4) Specimens (N=2) Large choanosomal principal styles 273-(339.4)409 x 12(16.8)-19 264-(359.7)528 x 13(16.1)-19 2694408.2)632 x 9-(13.8)19 Small choanosomal principal styles 133-(l 88.4)24 x 3 8(13.2)-18 113-(170.3)247 x 6-(9.5)13 1 32-(191.8)275 x 8411.2)13 Subectosomal auxiliary styles 162-(272.4)357 x 2.5(4.0)-5.5 96-(249.2)416x 1.5(3.6)-6 142(277.0)402x 1.5(4.2)-6 42-(71.1)-98 x 2-(4.9)-7.5) 63-(76.2)-92 x 3.5-(5.1)-7 15-(17.8)-20 14-(18.4)-22 I4-(19.2)-22 17-(84.6)-131 x 0.8-(1.5)-3 18-(109.1)192 x 0.8(1.8)-3.5 7I-(108.4)134 x 0.8(1.1)-1.8 58-(70.7)-88 x Echinating acanthostyles 3-(4.9)-6.5 Chelae Toxas present in C. (C.) transiens and it technically belongs to Clathria (Clathria). Clathria (Clathria) wilsoni Wiedenmayer, 1989 (Figs 79-80) Clathria wilsoni Wiedenmayer, 1989: 57-58, p1.5, fig.9, p1.24, fig.1, text-fig.39; Hooper & Wiedenmayer, 1994: 262. Rhaphidophlus wilsoni; Carpay, 1986: 27. MATERIAL. HOLOTYPE - NMVF51967: Garden Cove, N. side of Deal I., Kent Is Group, Bass Strait, Tas, 39°29'S, 147°20'E, 10m depth, 25.ii.1981, coll. F. Wiedenmayer et al. (SCUBA). HABITAT DISTRIBUTION. 10m depth; on algal covered boulders; Bass Strait and E. coast (Tas) (Fig. 79H). DESCRIPTION. Shape. Small sponge, thickly encrusting with irregular low-formed branches, 5mm high, up to 20mm diameter. Colour. Live colouration dull vermilion, beige preserved. Oscules. Oscules small, up to 2mm diameter, scattered on base and sides of branches. Texture and surface characteristics. Surface uneven, microconulose, hispid, particularly on points of branches. Texture easily compressible, spongy. Ectosome and subectosome. Ectosome skeleton a layer of subectosomal auxiliary styles, in small ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 169 D FIG. 79. Clathria (Clathria) wilsoni Wiedenmayer (holotype NMVF51967). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Accolada toxa. E, Oxhorn toxa. F, Palmate isochela. G, Section through peripheral skeleton. H, Australian distribution. I, Choanosomal skeletal structure. J, Holotype. bundles, erect on surface, with acanthostyles styles, embedded in peripheral fibres, form echinating peripheral fibres also contributing to plumose brushes, which approach but usually do ectosomal skeleton; choanosomal principal not pierce surface. MEMOIRS OF THE QUEENSLAND MUSEUM 170^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 80. Clathria (Clathria) wilsoni Wiedenmayer (holotype NMVF51967). A, Choanosomal skeleton. B,D, Echinating acanthostyles. C,E, Acanthostyle spines. F, Base of principal style., G, Base of auxiliary styles. H, Palmate isochelae. I, Larger accolada toxa. J, Smaller oxhorn toxas. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU 171 Choanosome. Choanosomal skeleton irregularly reticulate, consisting of moderately light, slender spongin fibres, up to 35pm diameter, forming irregular rectangular or polygonal meshes; spongin fibres cored by multispicular tracts of choanosomal styles, in rows of 2-10 spicules abreast, echinated by regularly dispersed acanthostyles; mesohyl matrix light, many vestigial and fully formed subectosomal styles, and numerous rhaphidiform toxa microscleres. Megascleres. Choanosomal principal styles slightly curved, with evenly rounded or slightly subtylote, smooth or rarely microspined bases. Length 164-(191.8)-290p,m, width 2.445.1)Subectosomal auxiliary subtylostyles straight, occasionally slightly curved, with tapering or rounded, smooth bases. Length 834153.6)282p,m, width 1-(2.3)-3.5p,m. Acanthostyles short, thick, subtylote, with relatively evenly dispersed spines. Length 31.6(62.6)-71 p.m, width 2.6-(6.6)-10Rm. Microscleres. Palmate isochelae very rare, small, poorly silicified. Length 4-(5.5)-9p,m. Toxas accolada, straight, very thin rhaphidiform, occasionally with roughened points; some smaller examples slightly curved, with reflexed arms. Length 71-(121.6)-168p,m, width 0.4-(0.7)-2p.m. REMARKS. This species is transitional between the ectosomal structures of Clathria and Thalysias, and it is not surprising that Carpay (1987) referred it to the latter. However, although there is a large size range of auxiliary styles, ectosomal spicule brushes are composed only of a single size of spicule, and the species is technically correctly included in Clathria (Clathria). Nevertheless, this example further illustrates the inappropriateness of recognising a strict (phylogenetic) boundary between these two genera. Clathria (C.) wilsoni is peculiar in having almost straight raphide-like toxas, which are known to occur in only a few microcionids (e.g. C. (Thalysias) costifera and C. (Wilsonella) reticulata, both from Australia, C. (T.) juniperina from the Caribbean, C. (C.) microxea from Madagascar, and Artemisina archegona from the Pacific coast of America). Wiedenmayer (1989) notes some similarities between C. (C.) wilsoni and other species of Hallmann's (1912) spicata' group, but this is not upheld here. OTHER SPECIES OF CLATHRIA (CLATHRIA) Clathria (Clathria)acanthostyli (Hoshino, 1981)zyxwvutsrqpo Thalysias acant host yli Hoshino, 1981: 156-157, text-fig.68, p1.7, fig.2 [Uchinoura, Japan]. MATERIAL. HOLO'TYPE: MMBSSAT-020. Japan. Clathria (Clathria) acanthotoxa (Stephens, 1916) Eurypon acant hot oxa Stephens, 1916: 238-239 [SW coast, Ireland]. Microciona acant hot oxa; Stephens, 1921: 50, p1.5, fig.!: Alander, 1942:62 [Sweden]; Burton, 1959b: 43 [Iceland]; Levi, 1960a: 75 [Isle of Man, W. Ireland]. Not Microciona acant hot oxa; Lilly et al., 1953: 97. Dict yociona acant hot oxa; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953: 527. Clat hr ia acant hot oxa; Van Soest, 1984b: 7 [generic synonymy for Dict yociona) . Not Microciona acant hot oxa Levi & Levi, 1989: 81, fig. 49 [see C. claudei, nom.nov.]. MATERIAL. HOLOTYPE: INM5R253.8.1916. NE. Atlantic, North Sea. Clathria (Clathria) anchorata (Carter, 1874) Dict yocylindrus anchorat us Carter, 1874: 251 [Gulf of Manaar, Ceylon]. Clat hria anchorat a; Vosmaer, 1880: 153 [Atlantic]; [?] Stephens, 1916:242 [W coast, Ireland]; Burton, 1959b: 42 [Iceland]; Levi, 1960a: 63-64 [English Channel, Atlantic]; [7] Vacelet, 1969: 206, text-fig.45 [Mediterranean]. Microciona anchorat a; Alander, 1942: 62 [Sweden]. Echinonem a anchorat um ; Dendy, 1889a: 44. Wilsonella anchorat um ; Hallmann, 1912: 152, 185, 189, 210, 211, 243, 296, 298, 299. Cionanchora anchorat a; de Laubenfels, 1936a: 108 [note]. Not Echinonem a anchorat um Carter, 1881a: 362, 379, 380 [S. coast of Australia]. Not Echinonem a anchorat um , varr.; Lendenfeld, 1888: 219220 [Port Jackson, New South Wales]. Not Echinonem a anchorat um var. ram osa; Whitelegge, 1901: 81. Clat hria t ypica, in part; Dendy, 1896: 32; Kirkpatrick, 1903: 248-249. Clat hria longichela Topsent, 1928a: 300, p1.10, fig.9. cf. Microciona prolifera; Vosmaer, 1935a: 610, 632, 668. MATERIAL. HOLOTYPE: LFM destroyed, fragmentsBMNH1953.11.11.63-69. NE. Atlantic, Mediterranean, Gulf of Manaar. Clathria (Clathria) antyaja (Burton & Rao, 1932) Dendrocia ant yaj a Burton & Rao, 1932: 348-350, p1.28, fig.12, text-figs 13-14 [Indian Ocean] MATERIAL. HOLOTYPE: IMP791/1. Indian Ocean. Clathria (Clathria) anthoides Levi, 1994 Clat hria ant hoides Levi, 1994: 36-37, fig. 21A, p1.7, fig.5 [Norfolk Rise, 610m depth]. MATERIAL. HOLOTYPE: MNHNDCL3637. SW. Pacific. Clathria (Clathria) arbuscula (Row, 1911) Row, 1911:347-349, p1.39, fig.22, p1.40, fig.25, text-fig.22 [Red Seat Ophlit aspongia arbuscula Ophlit aspongia horrida Row, 1911: 349-351, p1.40, fig.26, text-fig.23 [Red Seat 172zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Litaspongia arbuscula; de Laubenfels, 1954: 162 [note]. Kerasemna arbuscula; Pulitzer-Finali, 1982: 105. Kerasemna horrida; Pulitzer-Finali, 1982: 105. MATERIAL. HOLOTYPE: BMNH1912.2.1.63 (0. arbuscula); BMNH1912.2.1.65 (0. horrida). Red Sea. Clathria (Clathria) asodes (de Laubenfels, 1930) Eutypon asodes de Laubenfels, 1930: 27; de Laubenfels, 1932: 92-93, text-fig.54 [Carmel,California]; Lee & Gilchrist, 1985: 24-32 [biochemistry]. Dictyociona asodes; de Laubenfels, 1936a: 110 [note]. Clathria asodes; Van Soest, 1984b: 7 [generic synonymy]. Leptoclathria asodes; Sim & Bakus, 1986: 10 [California]; Bakus & Green, 1987: 72 [S. California]. MATERIAL. HOLOTYPE: USNM21442. NE. Pacific. Clathria (Clathria) atoxa (Bergquist & Fromont, 1988) Dictyociona atoxa Bergquist & Fromont, 1988: 105-106, p1.49, figs a-c [Tokatu]; Dawson, 1993: 38 [note]. MATERIAL. HOLOTYPE: NMNZPOR116. New Zealand. Clathria (Clathria) axociona Levi, 1963 Clathria axociona Levi, 1963: 49, text-fig.56, p1.8A [Cape of Good Hope, South Africa]; Uriz, 1988a: 83-84, pls 20-21a, text-fig. 59 [Namibia]. MATERIAL. HOLOTYPE: MNHNDCL632. SW. and S. Africa. Clathria (Clathria) barleei (Bowerbank, 1866) Isodiciya barleei Bowerbank, 1866: 333; Bowerbank, 1874: p1.57. Tragosia barleei; Topsent, 1894a: 25. Axinella barleei; Arndt, 1935: 88, fig.186. Clathria barleei; Van Soest & Stone, 1986: 45-46, fig.4 [Norway]; Ackers, Moss & Picton, 1992: 138 [Ireland]. Halichondria foliata Bowerbank, 1874: 198, p1.73, figs 1-5; Carter, 1876: 310, p1.12, fig.10, p1.29, fig.29; Bowerbank, 1882: 106. Esperia foliata; Fristedt, 1885: 41. Homeodictya foliata; Topsent, 1894a: 12. Echinoclathria foliata; Topsent, 19 13b: 38; Stephens, 1916: 234 [Ireland]; Stephens, 1921: 57; Hentschel, 1929: 894, 971 [Arctic]. Artemisina foliata; Burton, 1930a: 501, 529, p1.2, figs 1-2 [Norway]; Burton, 1959b: 42-43 [Iceland]. Amphilectus foliatus; Vosmaer, 1880: 118. Antho foliata; Alander, 1942: 63 [Sweden; defined in Esperiopsidae]. ? Halichondria mutula Bowerbank, 1874: 209, p1.74, figs 4-8; Bowerbank, 1882: 96 [as H. mutulus]. ? Amphilectus mutulus; Vosmaer, 1880: 118. Isodiciya laciniosa Bowerbank & Norman, 1869: 333. Amphilectus laciniosus; Vosmaer, 1880: 116. Clathria laciniosa; Arndt, 1935: 81, fig.167; Burton, 1959b: 42 [Iceland]. Art emisina laciniosa; Burton, 1950: 891 [revision]. MATERIAL. HOLOTYPE: BMNI-11930.7.3.338. Arctic, NE. Atlantic. Clathria (Clathria) basilana Levi, 1961 Clathria basilana Levi, 1961b: 520, fig.11 [Zamboanga, Philippines]; Van Soest, 1989: 1-2, fig.6 [Lesser Sumba Is, Indonesia]. MATERIAL. HOLOTYPE: MNHNDCL722. S. Philippines - E. Indonesia. Clathria (Clathria) borealis sp.nov. Clathria robusta Koltun, 1959: 186, p1.25, fig.5, text-fig. 147 [Arctic, USSR]; Van Soest & Stone, 1986: 47 [note]. Not Microciona strepsitoxa var. robusta Dendy, 1922: 60. Not Tenacia robusta; Burton & Rao, 1932: 339-340. MATERIAL. HOLOTYPE: ZIL, no fragment in BMNH. Note: C. (C.) robusta (Dendy, 1922) has priority. Arctic. Clathria (Clathria) bulbosa Hooper & Levi, 1993 Clathria (Clathria) bulbosa Hooper & Levi, 1993a: 12681270, figs 23-24 [Chesterfield Is, Coral Sea]. MATERIAL. HOLOTYPE: QMG300021. SW. Pacific. Clathria (Clathria) burtoni sp. nov Clathria proltfera Burton, 1940: 109, p1.4, figs 3-4, text-fig.2 [Argentina]. Not Spongia prolifera Ellis & Solander, 1786: 189. MATERIAL. HOLOTYPE: MA BA 15582, fragmentsBMNH1934.1.17.13, 116, 117. Note: C. prolifera (Ellis & Solander, 1786) has seniority. Province: SW. Atlantic. Clathria (Clathria) caespes (Ehlers, 1797), unrecognizable. Spongia tubulosa, in part, Esper, 1797: 196, p1.44 [Cape of Good Hope, South Africa]. Scopalina caespes; Ehlers, 1870: 19-20, 31. MATERIAL. HOLOTYPE: unknown. S. Africa. Clathria (Clathria) calypso Boury-Esnault, 1973 Clathria calypso Boury-Esnault, 1973: 286, text-fig.47 [Brazilian Basin] MATERIAL. HOLOTYPE: MNHNDNBE1035. tropical SW. Atlantic. Clathria (Clathria) cantabrica (Orueta, 1901) Rhaphidophlus filifer var. cantabrica Orueta, 1901: 331-335, pls 3,4 [Bay of Biscay]. Tenacia filifer var. cantabrica; Hallmann, 1920: 771. Clathria cantabrica; Levi, 1960a: 55-56, 63. MATERIAL. HOLOTYPE: Madrid. NE. Atlantic. Clathria (Clathria) compressa Schmidt, 1862 Clathria compressa Schmidt, 1862: 8, 10, 58-59, 86, p1.6, fig.1; Schmidt, 1864: 35, p1.4, fig.3; Crivelli, 1863: 299; Gray, 1867: 513; Claus, 1868: 23; Carter, 1875: 195; Vosmaer, 1880: 150 [Triest]; Graeff, 1882: 318; Vosmaer, 1884: 119; Ridley, 1884a: 443-449, 612-615; Schmidt, 1880: 34-35, 45, p1.4, fig.3; Ridley & Dendy, 1887: 147; Dendy, 1889c: 8; Hope, 1889: 337; Norman, 1892: 13; Topsent, 1892a: 17; Topsent, 1894a: 18; Heider, 1895: 280; Topsent, 1896: 123; Thiele, 1899: 13; Topsent, 1911: 10, 13; Dendy, 1922:64; Dendy, 1924a: 352-354; Wilson, 1925: 439; Topsent, 1925: 647-650, fig.8 [Banyuls, Gulf of Naples]; Topsent, 1928a: 62,299; Burton & Rao, 1932: 334-337; Burton, 1932a: 319; Burton, 1934a: 558; Topsent & Olivier, 1943: 1 [Monaco]; Koltun, 1959: 184; Levi, 1960a: 50, 52, 61, 62 [Mediterranean, Atlantic]; Levi, 1960b: 761, fig. 14 [Kayar, W. Africa]; Melone, 1963: 1-8; Sara & Melone, 1963: 362; Sara, 1964: 229; Simpson, 1968a: 102, 104-106, p1.17, table 49; Boury-Esnault, 1971: REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 173 323 [Banyuls]; Pulitzer-Finali, 1983: 610; DesqueyrouxFaundez & Stone, 1992: 9, 35, 103 [list]. Thalysias compressa; de Laubenfels, 1936a: 105-106. ? Spongia clathrus Esper, 1797; Vosmaer, 1880: 150. Not Citalina compressa; Carter, 1882b: 112-113. Not Halichondria compressa Carter, 1886g: 450 [Westemport Bay, Victoria]. Not Pseudoclathria compressa; Dendy, 1897: 259. MATERIAL. HOLOTYPE: LMJG15509, fragments BMNH1867.7.26.78, BMNH1910.1.1.2362, 2363. Mediterranean, NE. Atlantic, W. Africa. Clathria (Clathria) conica Levi, 1963 Clathria conica Levi, 1963: 50-51, text-fig.57, p1.8E [Cape of Good Hope]. MATERIAL. HOLOTYPE: MNHNDCL617. S. Africa. Clathria (Clathria) contorta (Bergquist & Fromont, 1988) Dictyociona contorta Bergquist & Fromont, 1988: 105, p1.48, figs d-f [Manukau]; Dawson, 1993: 38 [note]. MATERIAL. HOLOTYPE: NMNZPOR115. New Zealand. Clathria (Clathria) coralloides (Olivi, 1792) Spongia coralloides Olivi, 1792: 264; Bertoloni, 1819: 228; Blainville, 1819: 125; Lamouroux, 1824: 369; Martens, 1824: 535. Grantia coralloides; Nardo, 1833: 522. Halichondria corona Lieberkiihn, 1859: 521, 529, p1.11, fig.3. Clathria coralloides , Schmidt, 1862:7, 10, 11, 58, 85, p1.5, figs 10-11; Kiilliker, 1864: 52, 71; Schmidt, 1864: 34; Schmidt, 1868: 9,41; Schmidt, 1870: 56, 60; Gray, 1867: 513, 533, 552; Dybowsky, 1880: 5, 70, p1.4, figs 7,9; Krukenberg, 1880: 70, 72; Vosmaer, 1880: 149-150 [Mediterranean]; Ridley, 1881: 485; Vosmaer, 1881: 4; Vosmaer, 1882-6: 119, 121-122; Vosmaer, 1884b: 492; Vosmaer, 1885a: 186, 356; Carter, I882a: 281; Graeffe, 1882: 318; Carter, 1884a: 204; Ridley & Dendy, 1887: 147; Lendenfeld, 1889a: 410-415, 498-505, 586, 592, 594, 598, 602, 604, 608, 612, 614, 618, 622, 624, 628, 634, 640, 644, 646, 650, 669, p1.34, figs 193-205; Lendenfeld, 1890a: 72; Maas, 1892:427-428; Norman, 1892: 13; Maas, 1893: 331, 334-335, 414, 441, 444, p1.20, fig.19, p1.21, fig.32; Topsent, 1893d: 445; Topsent, 1894a: 18,24; Topsent, 1894b: 35; Heider, 1895: 280; Dendy, 1896: 36; Loisel, 1898: 38; Zimmermann, 1907: 308; Korschelt & Heider, 1910: 321; Walther, 1910: 21; Babic, 1921: 84 [Adriatic]; Babic, 1922: 244; Topsent, 1925: 646-647, fig.7 [Gulf of Naples]; Topsent & Olivier, 1943: 1 [Banyuls and Naples]; de Laubenfels, 1954: 139; Levi, 1960a: 61-62 [Mediterranean, Adriatic,Naples, Banyuls]; Laubier, 1966 [Banyuls]; Vidal, 1967 [Mediterranean]; Boury-Esnault, 1971: 323 [Banyuls]; Pulitzer-Finali, 1983: 568-569, 610 [Mediterranean]; Boury-Esnault & Lopes, 1985: 194-195, fig.43 [Azores]; Pansini, 1987 [Adriatic Sea]; Desqueyroux-Faundez & Stone, 1992: 9, 35, 103 [list]. Ophlitaspongia coralloides; Bowerbank, 1874: 10. Ophlitaspongia corona; Levi, 1960a: 61 MATERIAL. HOLOTYPE: unknown; fragments LMJG 15356, BMNH1867.7.26.74, 1910.1.1.2364, 2365. Mediterranean, NE. Atlantic. - Clathria (Clathria) curvispiculifera (Carter, 1880) Microciona curvispiculifera Carter, 1880a: 43,44,151,153, p1.4, fig.6a-d [Gulf of Manaar]; Vosmaer, 1935a: 608. ? Rhabdoploca curvispiculifera; Topsent, 1904a: 157-159. MATERIAL. HOLOTYPE: LFM missing, no fragment in BMNH. Gulf of Manaar. Note: virtually unrecognisable. Clathria (Clathria) dayi Levi, 1963 Clathria dayi Levi, 1963:51, p1.8B, text-fig.58 [Cape of Good Hope]; Sim & Byeon, 1989: 39, p1.4, figs 1-2 [Korea; possible misidentification]. MATERIAL. HOLOTYPE: MNHNDCL611. S. Africa. Clathria (Clathria) depressa Sara & Melone, 1966 Clathria depressa Sara & Melone, 1966: 2-4, text-figs 1-2, p1.1 [Olivetta, Portofino, Levante coast, Italy]; PulitzerFinali, 1983: 610. MATERIAL. HOLOTYPE: IMZUB. E. Mediterranean. Clathria (Clathria) discreta (Thiele, 1905) Microciona discreta Thiele, 1905: 447-449, 494, p1.3 I , fig. 65a-e [Calbuco, Chile]. Diciyociona discreta; Topsent, 1913a: 580, 583, 614, 618620, 642, p1.3, fig.5 [Gough I., S. Atlantic Ocean]; Burton, 1932: 324, p1.56, figs 3-4 [Falklands]; Burton, 1940: 112, p1.4, figs 1-2, p1.6, fig.2 [Argentina and Chile]; de Laubenfels, 1953a: 527; Desqueyroux, 1972: 29-30, figs 95-102, 136-137 [Chile]; Desqueyroux & Moyana, 1987: 49 [Chile, Tierra del Fuego, Argentina, Falkland Is, Kerguelen Is]. cf. Microciona prolifera; Vosmaer, 1935a: 608,646. MATERIAL. HOLOTYPE: ZMB3302, fragment: BMNH1908.9.24.159. SW. Atlantic, SE. Pacific, Subantarctic. Clathria (Clathria) elastica Levi, 1963 Clathria elastica Levi, 1963: 52, text-fig.59, p1.8C [Cape of Good Hope]. Not Clathria elastica Sara, 1978: 70 [see C. sarai, nom.novi. MATERIAL. HOLOTYPE: MNHNDCL604. S. Africa. Clathria (Clathria) elegans Vosmaer, 1880 Clathria elegans Vosmaer, 1880: 152 [North America]; Vosmaer, 1884b: 492. ? Anaata elegans; de Laubenfels, 1936a: 109. Not Plectispa elegans Lendenfeld, 1888: 226. Not Antherochalina elegans Lendenfeld, 18876: 787. cf. Microciona prolifera; Vosmaer, 1935a: 630. MATERIAL. HOLOTYPE: unknown. NE. Pacific. Note: virtually unrecognisable. Clathria (Clathria) foliacea Topsent, 1889 Clathria foliacea Topsent, 1889: 39-40, text-fig.5 [Banc de Campeche]; Van Soest, 1984b: 107, table 4; Van Soest & Stentoft, 1988: 126 [table]. Thalyseurypon foliacea; de Laubenfels, 1936a: 107-108 [Florida]. Pandaros foliaceum; Wiedenmayer, 1977: 144 [note]. Clathria carteri Topsent, 1889: 38-39, text-fig.4B [Banc de Campeche]; Van Soest, 1984b: 108. Thalyseurypon carteri; de Laubenfels, 1936a: 107. Pandaros carteri; Wiedenmayer, 1977: 144 [note]. MATERIAL. HOLOTYPE: MNHN missing, reference specimen USNM22516. NE. Atlantic, Caribbean. Clathria (Clathria) foliascens Vacelet & Vasseur, 1971 MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVUT 174^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA fig.49, p1.4,figs 5-6 [Tulear Madagascar]; Vacelet & Vasseur, 1977: 114; Vacelet et al., 1976: 71 [Tulear, Madagascar] MATERIAL. HOLOTYPE: MNHNDJV30. W. Indian Ocean. MATERIAL. HOLOTYPE: MNHN missing (fragment BMNH1954.2.20.85, unconfirmed); 'representative specimen': BMNH1926.2.19.2. Note: Burton's species is probably the same as Lamarck's but this cannot be verified and the species is taken in the sense of Burton (1931a). South Africa. Clathria (Clathria) frondiculata (Schmidt, 1864) Clathria (Clathria) labyrinthica (Schmidt, 1864) Clat hria foliascens Vacelet & Vasseur, 1971: 95-96, text- Reniera ( ?) frondiculat a Schmidt, 1864: 39, 40, 45, p1.4, fig. 10 [Adriatic]. Pit alia frondiculat a; Gray, 1867: 524. Clat hriafrondiculat a; Vosmaer, 1880:153 [Triest]; Vosmaer, 1885:236; Norman, 1892: 13; Topsent, 1892b: 23; Heider, 1895: 282; Burton, 1930a: 514. cf. Clat hria com pressa and C. coralloides; Vosmaer, 1935a: 625 [intermediate form]. MATERIAL. HOLOTYPE: LMJG, fragment: BMNH1910.1.1.542. E. Mediterranean. Clathria (Clathria) gorgonoides (Dendy, 1916) Echinodict yum gorgonoides Dendy, 1916a: 129 [Kattiawar, Indian Ocean]; Hooper, 1991: 1348. MATERIAL. HOLOTYPE: BMNH1920.12.9.38. W. India. Clathria (Clathria) hexagonopora Levi, 1963 Clat hria hexagonopora Levi, 1963: 53-54, text-fig.60, p1.8D [Cape of Good Hope]. MATERIAL. HOLOTYPE: MNHNDCL620. S. Africa. Clathria (Clathria) indica Dendy, 1889 Clat hria indica Dendy, 1889b: 73, 84, p1.4, fig. 10 [Gulf of Manaar, Ceylon]; Burton 8c Rao, 1932: 336-337, p1.18, figs 8-9 [Tuticorin, India]; Thomas, 1979a: 58, p1.3, fig. 10 [Mozambique]. Wilsonella indica; Dendy, 1905: 171 [Ceylon]; Hallmann, 1912: 242; Burton, 1931a: 346 [Natal Coast]. MATERIAL. HOLOTYPE: BMNH1887.8.4.19, fragment: MNHNDCL2528. E. Africa, SE. India, Gulf of Manaar. Clathria (Clathria) inhacensis Thomas, 1979 Clat hria inhacensis Thomas, 1979: 27-28, p1.2, fig.2 [Inhaca I., Mozambique]. MATERIAL. HOLOTYPE: MRAC511. E. Africa. Clathria (Clathria) intermedia Kirk, 1911 Clat hria int erm edia Kirk, 1911: 579, text-fig.5 [Kermadec Is]; Fell, 1950: 11, text-fig.2; Berquist & Fromont, 1988: 109-110; Dawson, 1993:37 [note]. Thalysias int erm edia; de Laubenfels, 1936a: 105. MATERIAL. HOLOTYPE: NMNZ unregistered. N. New Zealand. Clathria (Clathria) irregularis (Burton, 1931) Mar leyia ir r egular is Burton, 1931a: 346, p1.23, fig.6, text- fig.5 [Durban, Natal]; de Laubenfels, 1936a: 109 [note]; Van Soest, 1984b: 130 [note]. MATERIAL. HOLOTYPE: NM1279; paratype: BMNH1934.10.1.17. S. Africa. Clathria (Clathria) juncea Burton, 1931 Clat hria j uncea Burton, 1931a: 343, p1.23, fig.5, text-fig.3 [Natal coast]; Levi, 1960a: 85; Levi, 1963: 66 [note]. Labacea j uncea; de Laubenfels, 1936a: 66 [note]. cf. Alcyonium j unceum Lamarck, 1816: 77. cf. Anom oclat hria opunt ioides var; Topsent, 1933: 26 [note]. Reniera labyrint hica Schmidt, 1864:39, p1.4, fig.9 [Adriatic]. Clat hr ia labyr int hica; Burton, 1930a: 514 [imperfectly known]. MATERIAL. HOLOTYPE: LMJG. Mediterranean. Clathria (Clathria) laevigata Lambe, 1893 Clat hria laevigat a Lambe, 1893:31-32, p1.2, figs 9-9f [Pacific coast, Canada]; Lambe, 1900: 160; Koltun, 1959: 185, text-fig. 146 [Kurile Is]. Thalysias laevigat a; de Laubenfels, 1936a: 105. cf. Microciona prolifera, tropus sent a; Vosmaer, 1935a: 641. MATERIAL. HOLOTYPE: NMCIC1900-2892. NW. and NE. Pacific. Clathria (Clathria) lissosclera Bergquist & Fromont, 1988 Clat hria lissosclera Bergquist & Fromont, 1988: 106-107, p1.49, figs d-f [Mayor I.]; Dawson, 1993: 37 [note]. MATERIAL. HOLOTYPE: NMNZPOR112. New Zealand. Clathria (Clathria) lobata Vosmaer, 1880 Clat hria lobat a Vosmaer, 1880: 151 [Cape of Good Hope]; Vosmaer, 1882: 45; Carter & Hope, 1889: 101, 105; Stephens, 1915: 444-445 [Cape of Good Hope]; Levi, 1963: 54, text-fig.61, p1.8F [Cape of Good Hope]. Rhaphidophlus lobat us, var. horrida; Ridley & Dendy, 1887: 153-155, p1.28, fig.1, p1.29, fig.4a-c [Simon's Bay, Cape of Good Hope]. Ligrot a lobat a; de Laubenfels, 1936a: 125. A rt em isina lobat a; Van Soest, 1984b: 130 [generic synonymy for Ligrot a] . cf. Microciona prolifera; Vosmaer, 1935a: 631-632. MATERIAL. HOLOTYPE: unknown, fragments RMNH276-277, BMNH1887.5.2.100. S Africa. Clathria (Clathria) macroisochela Levi, 1994 Clat hria ( Clat hria) m acroisochela Levi, 1994: 37, fig.21b, p1.7, figs 7-9 [Norfolk Rise, 680m depth]. MATERIAL. HOLOTYPE: MNHNDCL3638. SW Pacific. Clathria (Clathria) maeandrina Ridley, 1884 Clat hria m aeandrina Ridley, 1884a: 588, 614, p1.53, fig.1, p1.54, fig.H [Amirante Is]; Burton, 1930c: 668 [Gulf of Manaar]; Burton, 1959: 244 [Red Sea]. MATERIAL. HOLOTYPE: BMNH1882.10.17.55. Red Sea, W Indian Ocean, Gulf of Manaar. Clathria (Clathria) marissuperi PulitzerFinali, 1983 Clat hria m arissuperi Pulitzer-Finali, 1983: 569-571, text- fig.67 [Italy]. MATERIAL. HOLOTYPE: IZUG(MSNG 47179). Mediterranean. Clathria (Clathria) menoui Hooper & Levi, 1993 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 175 Clathria (Clathria) menoui Hooper & Levi, 1993a: 12701273, figs 25-26 [New Caledonia]. MATERIAL. HOLOTYPE: QMG301267. SW. Pacific. Clathria (Clathria) meyeri (Bowerbank, 1877) Ophlitaspongia meyeri Bowerbank, 1877: 456 [Selat Japen, Irian Jaya, E. Indonesia]. Clathria meyeri; Vosmaer, 1880: 154. ? Anaata meyeri; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: Dresden Musuem destroyed (fragments BMNH1877.5.21.1306-1309). Indonesia. Clathria (Clathria) microchela (Stephens, 1916) Eurypon microchela Stephens, 1916: 240-241 [SW coast, Ireland]; Lilly et al., 1953 [Lough me, Ireland]; Burton, 1959b: 44 [Iceland]; Van Soest & Weinberg, 1980: 10 [Lough Inc. Ireland]. Dictyociona microchela; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953: 528. Clathria microchela; Van Soest, 1984b: 7 [generic synonymy]. Not Microciona microchela Hechtel, 1965: 41. MATERIAL. HOLOTYPE: INMSR353.10.1916. NE. Atlantic. Clathria (Clathria) microxa Desqueyroux, 1972 Clathria microxa Desqueyroux, 1972: 27-28, text-figs 7680,134 [Gulf of Corcovado, Chile]. MATERIAL. HOLOTYPE: ICBUC. SE Pacific. Clathria (Clathria) mortensenii Brondsted, 1923 Clathria mortensenii Brondsted, 1923: 143-144, text-fig.22 [Campbell Is]; Burton, 1940: 111 [Argentina]; Bergquist & Fromont, 1988: 107-108, p1.50, figs a-c [N. New Zealand]; Dawson, 1993: 37 [note]. Microciona mortensenii; de Laubenfels,I936a: 111; Burton, 1940: 111 [Argentina]. Microcionaheterospiculata; Bergquist, 1961a: 39 [N of New Zealand]. Not Microciona heterospiculata Brondsted, 1924: 465, textfig.20. MATERIAL. HOLOTYPE: UZM, fragment: BMNH1930.8.11.10. SW. Atlantic, Subantarctic, New Zealand. Clathria (Clathria) mosulpia Sim & Bakus, 1989 Clathria mosulpia Sim & Byeon, 1989: 38-39, p1.3, figs 1-5. [South Korea]. MATERIAL. HOLOTYPE: HNUKP0r9. S. China Sea. Clathria (Clathria) obliqua (George & Wilson, 1919) Esperiopsis obliqua George & Wilson, 1919: 148-150, p1.60, figs 20-23, p1.66, fig.58 [North Carolina]. Microciona obliqua; de Laubenfels, 1936a: 111. cf. Microciona prolifera; de Laubenfels, 1947; de Laubenfels, 1949a. Haliclona oculata; Pearse & Williams, 1951 [North Carolina]. Chalina arbuscula; Coues & Yarrow, 1879: 312; Verrill & Smith, 1873: 743 [North Carolina]. Tenaciella obliqua; Wells et al., 1960: 218-219, text-figs 16,25 [North Carolina]. Not Tenaciella obliqua; Alcolado, 1976: 5. Clathriaobliqua; Van Soest, 1984b: 104, 108, table 4 [affinity with Clathria proliferat Not Dictyociona adioristica de Laubenfels, 1953a: 526; Van Soest, I984b: 108 [possible synonym]. MATERIAL. HOLOTYPE: USNM23612, paratypeUSNM23613. NW. Atlantic-Caribbean. Clathria (Clathria) oculata Burton, 1933 Clathria oculata Burton, 1933: 250-251, text-fig.2 [Natal]; Levi, 1963: 67 [Natal]. Thalysias oculata; de Laubenfels, 1936a: 105. MATERIAL. HOLOTYPE: BMNH1933.7.4.65. S. Africa. Clathria (Clathria) oxneri (Topsent, 1928) Hymedesmia oxneri Topsent, 1928a: 256 [Azores]. Dictyociona oxneri; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953a: 528. Clathria oxneri; Van Soest, 1984b: 7 [generic synonymy for Dictyociona]. MATERIAL. HOLOTYPE: MOM (fragment MNHNDT963). NE. Atlantic. Clathria (Clathria) pachystyla Levi, 1963 Clathria pachystyla Levi, 1963: 56, text-fig.63, p1.8G [Cape of Good Hope]. MATERIAL. HOLOTYPE: MNHNDCL609. S. Africa. Clathria (Clathria) papillosa Thiele, 1905 Clathria papillosa Thiele, 1905: 449-450, text-fig.66 [Calbuco, Chile]; Burton, 1932a: 319 [Falkland Is]; Desqueyroux-Faundez & Moyana, 1987: 49 [Chile, Argentina, Falkland Is, Kerguelen Is]. Pseudanchinoepapillosa; de Laubenfels, 1936a: 109; Burton, 1940: 115, p1.5, figs 1-7 [Argentina]. MATERIAL. HOLOTYPE: ZMB not located, fragment: BMNH1908.9.24.162. SE. Pacific, SW. Atlantic. Clathria (Clathria) pellicula Whitelegge, 1897 Clathria pellicula Whitelegge, 1897: 327-328 [Funafuti (Ellice Is)]. Hymeraphia pellicula; Hallmann, 1912: 208 209. cf. Microciona prolifera tropus tegens; Vosmaer, 1935a: 641-2. - MATERIAL. HOLOTYPE: AMG1660. SW central Pacific. Clathria (Clathria) plurityla Pulitzer-Finali, 1983 Clathria plurityla Pulitzer-Finali, 1983: 571-572, 610, textfig.68 [Italy]. MATERIAL. HOLOTYPE: IZUG(MSNG47180). Mediterranean. Clathria (Clathria) productitoxa (Hoshino, 1981) Thalysias productitoxa Hoshino, 1981: 157-159, text-fig.69, p1.7, fig.3 [Uchinoura]. MATERIAL. HOLOTYPE: MMBSSAT-018. Japan. Clathria (Clathria) prolifera (Ellis & Solander, 1786) Spongia prolifera Ellis & Solander, 1786: 189-190, p1.58, fig.5; Linnaeus, 1791: 3822; Esper, 1794: 178,281; Bose, 1802:143; Lamarck, 1814: 372; Lamouroux, 1816: 81-82; MEMOIRS OF THE QUEENSLAND MUSEUM 176^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Montagu, 1818: 86; Blainville, 1819: 106; Lamouroux, 1821: 31, 109, p1.58, fig.5; Lamouroux, 1824: 368; Templeton, 1836: 472; Johnston, 1842: 170-171; Gray, 1848: 19, 23. Not Spongia prolifera; Grant, 1826: 115-116, 123, 135, 138, 347. Microciona prolifera; Verrill, 1873: 741-742, pls 1-5 [USA]; Verrill, 1880: 232; Coues & Yarrow, 1879: 312 [North Carolina]; Hyatt, 1885: 131; Norman, 1892: 14; Wilson, 1900: 350 [Beaufort, N.Carolina]; Wilson, 1902:396-397; Wilson, 1907: 246; Wilson, 1910: 1269; Wilson, 1911: 3-11, 14, 29-30, p1.1, figs 1-6, p1.2, figs 7-12, p1.3, figs 13-20, p1.4, figs 21-25, p1.5, figs 26-32; George & Wilson, 1919: 157-158, p1.62, figs 31, 33, p1.63, figs 35-36, p1.66, fig.57 [North Carolina]; Allee, 1923: 175; Galtsoff & Pertzoff, 1926: 239-254 [physiology]; Burton & Rao, 1932: 344 [Arabian Sea; probable misidentification]; Proctor, 1933: 104; de Laubenfels, 1936a: 111; McDougall, 1943: 331-332; Bergmann, Schedl & Low, 1945: 580; de Laubenfels, 1947:35; Hopkins, 1956a: 44; Hopkins, 1962: 124; Hartmann, 1958a: 36-41, text-fig.10, table 11 [New England, USA]; Levi, 1960a: 52; Wells eta!., 1960: 213216, text-figs 18,22 [North Carolina]; Little, 1963: 49 [Honda]; Wells et al., 1964: 757-758 [North Carolina]; Bagby, 1966: 167-181, pls 3-5 [cytology]; Simpson, 1968a: 18, pls 1-5 [Connecticut, USA]; Simpson, 1968b: 252-277 [reproduction]; Wendt, 1970: 3500-B [cytology]; Sindelar, 1970: 3771-B [cytology]; Bagby, 1970: 579-594 [ultrastructure]; Kunen et al., 1970:565-576 [physiology]; Madri et al., 1970: 1-5 [biochemistry]; Swartz, 1972: 17 [ecology]; Bagby, 1972: 217-244 [ultrastructure]; Bito, 1972: 65 [biochemistry]; Bose et al., 1972: 217-222 [biochemistry]; Turner & Burger, 1973: 509-510, textfig.! [cytology]; Weinbaum & Burger, 1973: 510-512, [biochemistry]; Stempien, 1973: 363; Sutherland, 1974: 859-873 [ecology]; Turner et al., 1974: 35 [cytology]; Bose, 1974: 476-490 [chemistry]; Jefferts et al., 1974: 244-247 [biochemistry]; Reiswig, 1975:493-502 [physiology]; Reed et al., 1976: 153-169 [cytology]; Jumblatt et al., 1976: 73-86 [cytology]; Morales & Litchfield, 1976: 206-216 [biochemistry]; Burger, 1977: 357-376 [cytology]; Greenberg et al., 1977: 95-102 [cytochemistry]; Morales, 1977: 5043 [biochemistry]; Morales & Litchfield, 1977: 570-576 [biochemistry]; Simpson, 1978: 31-42 [morphology]; Burkart eta!., 1979: 239-246 [cytology]; Leith, 1979: 212-223 [cytology]; Litchfield & Liaaen-Jensen, 1980: 359-365 [biochemistry]; Jumblatt et al., 1980: 1038-1042 [biochemistry-cytology]; Saxegaard eta!., 1981: 325-327 [biochemistry]; Lee & Nicol, 1981: 445 [chemistry]; Biernbaum, 1981: 85-96 [ecology]; Liaaen-Jensen et al., 1982: 170-171 [biochemistry]; Misevic & Burger, 1982: 200 [biochemistry-cytology]; Misevic eta!., 1982: 6931-6936 [biochemistry-cytology]; Kuhns et al., 1980: 73-79 [cell reaggregation]; Dunham et al., 1983: 4756 [cell reaggregation]; Rice & Humphreys, 1983: 6394-9 [biochemistry]; Akiyama & Johnson, 1983: 687-694 [biochemistry]; Collier, 1983: 428-432 [biochemistry]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Dunham et al., 1985: 2914 [cytology]; Sharma et al., 1985: 241 [biochemistry]; Knight & Fell, 1987: 263 [physiology]; Misevic eta!., 1987: 5870 [cytology]; Stanley-Samuelson, 1987:92 [chemistry]; Sliwka et al., 1987: 245 [chemistry]; Knight & Fell, 1987: 253 [cytology]; Misevic & Burger, 1988: 134-152; Fell, Knight & Rieders, 1989: 195; Misevic & Burger, 1988: 134; 1990a: 307; 1990b: 20577 [cytology]; Kuhns, Misevic & Burger, 1990: 358 [biochemistry]; Leamon & Fell, 1990: 265 [cytology]; Ayanoglu et al., 1990: 597; Lam et al., 1991: 372 [biochemistry]; Misevic & Burger, 1990c: 81 [chemistry]; Fell, 1990: 497 [ecology]; Misevic et al., 1990: 182 [ontogeny]; Spillmann et al., 1993: 13378 [chemistry]; Aho eta!., 1993: 7288 [genetics]. Not Microcionaprolifera; de Laubenfels, 1949a: 12, text-figs 8-10 (Pearse & Williams, 1951: 135); Johnson, 1971: 110-111, text-fig.14 (Van Soest, 19846: 93). cf. Microciona prolifera, in part; Vosmaer, 1935a: 612-613 [excessive lumping of species]. Clathria (Clathria) prolifera; Van Soest, 19846: 91-93, textfig.35, table 4 [North Carolina]. Not Clathria prolifera Burton, 1940: 109 [see C. burtoni, nom.nov.]. Spongia ostacina Rafinesque, 1819: 150. Spongia urceolata Desor, 1851: 67. Clathria delicata Lambe, 1896: 12, 160, 192, p1.2, fig.2 [St.Lawrence Gulf]; Lambe, 1900: 160; Whiteaves, 1901: 18 [Canada]; Hentschel, 1912: 365; Hentschel, 1929: 971; Hartman, 1958: 37. Thalysias delicata; de Laubenfels, 1936a: 105. Esperiopsis obliqua, in part; de Laubenfels, 1947: 5. MATERIAL. HOLOTYPE: BMNH missing; representative specimens: USNM23562, ZMAPOR38, MCZH6907, PMNH754, PMNH1912E, NMCIC1900-2874. NW. Atlantic. Clathria (Clathria) pyramidalis (Brondsted, 1924) MicrocionapyramidalisBrondsted, 1924:466, text-figs 21a-e [Slipper Is]; Dawson, 1993: 37 [note]. Dictyociona pyramidalis; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953a: 528; Bergquist & Fromont, 1988: 104. MATERIAL. HOLOTYPE: UZM (not located). New Zealand. Clathria (Clathria) rectangulosa Schmidt, 1870 Clathria rectangulosa Schmidt, 1870: 60 [Tortugas, Florida]; Vosmaer, 1880: 149; Ridley & Duncan, 1881: 485; Wiedenmayer, 1977: 261, table 52 [imperfectly known]; Desqueyroux-Faundez & Stone, 1992: 36, 103. cf. Clathria coralloides; Vosmaer, 1880: 149. MATERIAL. HOLOTYPE: BMNH1870.5.3.68. NW. Atlantic - Caribbean. Clathria (Clathria) rhaphidotoxa Stephens, 1915 Clathria rhaphidotoxa Stephens, 1915: 445-447, p1.38, fig.2, p1.40, fig. 15 [Saldanha Bay]; Levi, 1963: 57-58, text-fig. 65, p1.8H [St. Helena and Saldanha Bays]. Thalysias raphidotoxa; de Laubenfels, 1936a: 105. MATERIAL. HOLOTYPE: RSME 1921.143.1451 (fragment BMNH 1953.11.11.144). S Africa. Clathria (Clathria) sarai sp.nov. Clathria elastica Sara, 1978: 70-73,text-figs 44-46 [Cape San Sebastiono, Tierra del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 50 [Tierra del Fuego, Argentina]. Not Clathria elastica Levi, 1963: 52. MATERIAL. HOLOTYPE: IZUG116, fragment: MNHNDCL604. Note: C. elastica Levi, 1963 has seniority. SW Atlantic. Clathria (Clathria) saraspinifera sp. nov. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWV ^ 177 Clathria spinifera Sara, 1978: 67-70, text-figs 41-43 [Rio Grande, Tierra del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 50 [Tierra del Fuego, Argentina]. Not Rhaphidophlus filifer var. spinifera Lindgren, 1897: 483. Not Rhaphidophlus spinifer Thiele, 1903a: 958, p1.28, fig. 23; Hallmann, 1912: 177. MATERIAL. HOLOTYPE: IZUG147, fragment: MNHNDCL2755. SW. Atlantic. Note: C. spinifera (Lindgren, 1897) has seniority. Clathria (Clathria) sartaginula (Lamarck, 1814) Spongia sartaginula Lamarck, 1814: 383,362 [locality unknown]. Pseudanchinoe sartaginula; de Laubenfels, 1936a: 109 [note]. Clathria sartaginula; Topsent, 1930:45, p1.4, fig. 1 . MATERIAL. HOLOTYPE: MNHNDT527. Unknown. Clathria (Clathria) shirahama Tanita, 1977 Clathria shirahama Tanita, 1977: 38, p1.2, fig.9, text-fig.6 [Kii-Shirahama]; Hoshino, 1981: 161. MATERIAL. HOLOTYPE: MMBS. Japan. Clathria (Clathria) spinispicula Tanita, 1968 Clathria spinispicula Tanita, 1968: 48-49, p1.1, fig.6, textfig.8 [Ariake Sea]; Rho et al., 1972: 5, p1.4, figs 9-10 [South Korea]; Hoshino, 1981: 161 [Ariake Sea]; Sim & Bakus, 1988: 25 [Korea]; Sim & Byeon, 1989: 38 [Korea]. MATERIAL. HOLOTYPE: MMBS. Japan, S. China Sea. Clathria (Clathria) spongodes Dendy, 1922 Clathria spongodesDendy, 1922: 69, p1.6, fig.1, p1.14, fig.2 [Amirante]; Vacelet et al., 1976: 70-71, text-fig. 48, p1.3, fig.e [Madagascar]. Damoseni spongodes; de Laubenfels, 1936a: 110 [note]. Clathria madrepora Dendy, 1922: 68-69, p1.5, fig.3, p1.14, fig.1 [Seychelles]; Sim & Kim, 1988: 25, p1.2, figs 1-2 [Korea]; Sim & Byeon, 1989: 38 [Korea; possible misidentification]. Thalysias madrepora; de Laubenfels, 1936a: 105. Clathria spongiosa Burton, I959a: 245, text-fig. 26 [Red Sea]; Vacelet et al., 1976: 70 [with question]. MATERIAL. HOLOTYPE: BMNH1921.11.7.58. W. Indian Ocean, ? S. China Sea. Clathria (Clathria) surculosa (Esper, 1797) Spongia surculosa Esper, 1797: 39, p1.65 A3 ['East Indies']. Clathria surculosa; Ehlers, 1870: 23,31. MATERIAL. HOLOTYPE: Unknown. Indonesia. Clathria (Clathria) terranovae Dendy, 1924 Clathria terrae-novae Dendy, 1924a: 353-354, p1.12, fig.5, p1.14, figs 9-13 [North Cape, New Zealand]; Dawson, 1993: 37 [note]. Dictyociona terrae-novae; Burton, 1932a: 324 [Falkland Is]; Burton, 1940: 112-114, p1.5, figs 3-4 [Argentina]; de Laubenfels, 1953a: 528; Koltun, 1964a: 72-73 [Antarctica]. Clathria terranova; Koltun, 1976: 188; Rho & Sim, 1976:74, p1.6, figs 3-4 [Seogwipo, South Korea; possible misidentification]; Boury-Esnault & van Beveren, 1982: 107-108, p1.18, fig.69, text-fig.31 [Kerguelen Is]; Bergquist & Fromont, 1988: 109, p1.50, figs d-f, p1.51, fig.a [N. New Zealand]; Sim & Byeon, 1989: 38 [Korea]. MATERIAL. HOLOTYPE: BMNH1923.10.1.132, paratypes BMNH1923.10.1.133, 134. Antarctica, Subantactic, New Zealand, SW Atlantic. Clathria (Clathria) textile (Carter, 1876) Cornulum textile Carter, 1876: 309. Clathria textile; Vosmaer, 1880: 154 [Shetland Is; imperfectly known]. MATERIAL. HOLOTYPE: BMNH1882.7.28.75. NE Atlantic. Clathria (Clathria) tortuosa Uriz, 1988 Clathria tortuosa Uriz, 1988a: 86-87, pls 22a, 42a-b, textfig.62 [Namibia]. MATERIAL. HOLOTYPE: ABIP6B-11d. SW Africa. Clathria (Clathria) toxistricta Topsent, 1925 Clathria toxistricta Topsent, 1925:656-658, text-fig. 13 [Gulf of Naples]; Sara, 1960a: 462 [Ischia]; Levi, 1960a: 62-63 [Naples]; Boury-Esnault, 1971: 324 [Banyuls]; PulitzerFinali, 1983: 610 [list]. MATERIAL. HOLOTYPE: MOM, fragments-MNHNDT327, 1244. Mediterranean. Clathria (Clathria) toxistyla (Sara, 1959) Microciona toxistyla Sara, 1959: 17, text-fig.6 [Naples]; Siribelli, 1960:9-10, text-fig. 3B [Naples]; Sara, 1961:47 [Adriatic Sea]; Cimino et al., 1979: 3619-3622 [biochemistry]. Clathria toxistyla; Melone, 1963: 5-7, p1.1, fig.2, text-fig.2 [Adriatic Sea; erect form of Microciona toxistyla]; Sara, 1964: 229 [Ligurian Sea]; Sara & Melone, 1963: 20 [Adriatic Sea]; Labate, 1964: 334 [Adriatic Sea]; PulitzerFinali, 1983: 569, 610, fig.66 [Mediterranean]. MATERIAL. HOLOTYPE: ZSN1000. Mediterranean. Clathria (Clathria) toxivaria (Sara, 1959) Microciona toxivaria Sara, 1959: 14, p1.1C, text-fig.5 [Naples]; Siribelli, 1960: 8, text-fig.3A [Naples]. Clathria toxivaria; Melone, 1963: 2-3, p1.1, fig.1, text-tig.1 [Adriatic; erect forms of Microciona toxivaria]; PulitzerFinali, 1977: 61 [Bay of Naples]; Sara, 1964: 229-230 [Ligurian Sea]; Sara & Melone, 1963: 20-21 [Adriatic]; Labate, 1964: 334 [Adriatic]; Boury-Esnault, 1971: 323 [Banyuls]; Pulitzer-Finali, 1983: 568, 610 [Mediterranean]. MATERIAL. HOLOTYPE: ZSNGG920, paratype ZSNGG923. Mediterranean. Clathria (Clathria) toxotes (Schmidt, 1868) Scopalina toxotes Schmidt, 1868: 12, 26, 39, 40, 44, p1.5, fi g.5 [Canal of Zara, Adriatic]; Schmidt, 1870: 2, 56; Pagenstecher, 1872:43; Hyatt, 1877: 500; Schmidt, 1880: 81; Vosmaer, 1880: 118-119; Fristedt, 1885:37; Vosmaer, 1885b: 353; Carter & Hope, 1889: 101; Schulze & Lendenfeld, 1889: 9; Heider, 1895: 281; Kieschnick, 1896: 533; Thiele, 1903a: 959; Svarcevskij, 1906: 342; Levi, 1960a: 55. ? Microciona toxotes; de Laubenfels, 1936a: 111 [imperfectly known]. cf. Microciona armata; Vosmaer, 1935a: 627. MATERIAL. HOLOTYPE: unknown, possibly LMJG. Mediterranean. MEMOIRS OF THE QUEENSLAND MUSEUM 178^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Clathria) typica Kirkpatrick, 1904 Clathria typica Kirkpatrick, 1904: 148 [Natal]; Hallmann, 1912: 208 [anomalous species]. ? Tenacia clathrata; Carter, 1875: 195 [nomen nudum]; Carter, 1878: 160,163; sensuV osmaer, 1935a: 628. Not Echinonema typicum Carter, 1881a: 362. MATERIAL. HOLOTYPE: BMNH1902.11.16.31. Note: imperfectly known; listed in BMNH register as C. typica Carter. S Africa. Clathria (Clathria) ulmus Vosmaer, 1880 Clathria ulmus Vosmaer, 1880: 151 [locality unknown]; Ridley, 1884a: 444. Thalysias ulmus; de Laubenfels, 1936a: 105 [note]. MATERIAL. HOLOTYPE: possibly RNHL. Unknown locality. Clathria (Clathria) unica Cuartas, 1993 Clathria unica Cuartas, 1993: 112 [Argentina]. MATERIAL. HOLOTYPE: Division Invertebrados del Museo de Ciencias Naturales de La Plata, Argentina. SW Atlantic. Clathria (Clathria) vasiformis (de Laubenfels, 1953) Thalyseutypon vasiformis de Laubenfels, 1953a: 525, textfig.4 [Gulf of Mexico]; Little, 1963:50 [note]. Clathria vasiformis, Van Soest, 1984b: 109, table 4. Myxilla beani; Topsent, 1892b: 23; Topsent, 1894a: 8, 9, 25; Hanitsch, 1894: 179. cf. Microciona prolifera; Vosmaer, 1933: 610. MATERIAL. HOLOTYPE: BMNH1930.7.3.339. Note: Synonym of Antho involvens (Levi, 1960a: 76). Echinonema caespitosa Carter, 1885f: 352 [Port Phillip, Victoria]. Plumohalichondria caespitosa; Dendy, 1896: 41. MATERIAL. HOLOTYPE: BMNH1886.12.15.453. Note: referred to Anchinoidae, Plumohalichondria. Antherochalina concentrica Lendenfeld, 1887b: 788, p1.22, fig.42 [Port Molle (Airlee Beach), Q1d.]. Cymbastela concentrica; Hooper & Bergquist, 1992: 114. MATERIAL. HOLOTYPE: AMZ1993 (lectotype), fragment: BMNH1886.8.27.451, 460 (paralectotypes). Note: referred to Axinellidae, Cymbastela. Clathria corallorhizoides Fristedt, 1887: 460, p1.25, figs 7377, p1.29, fig.23. MATERIAL. HOLOTYPE: unknown, fragment: BMNH1910.1.1.1445. Note: referred to Myxillidae, synonym of Lissodendoryx complicata (Lundbeck, 1905: 166). Antherochalina elegans Lendenfeld, 1887b: 787, p1.22, ML4-232. E Pacific. fig.40. Syringella elegans; Burton, 1934a: 558. Raspailia (Syringe/la) elegans; Hooper, 1991: 1262. MATERIAL. HOLOTYPE: BMNHI886.8.27.452. Note: referred to Raspailiidae, Raspailia. Clathria (Clathria) zoanthifera Levi, 1963 Echinonema incrustans Carter, 1885f: 353 [Port Phillip, . Pandaros vasiformis; Wiedenmayer, 1977: 144 [note]. MATERIAL. HOLOTYPE: USNM23403, paratype MLUM- Clathria zoanthifera Levi, 1963: 58, text-fig.66, p1.9D [Cape of Good Hope]. MATERIAL. HOLOTYPE: UCT (fragment IVINHNDCL607). S Africa. TRANSFERS List of other species described in Clathria (Clathria), or one of its synonyms, but now transferred to another genus. ClathrM australis Lendenfeld, 1888: 222 [Port Phillip, Victoria]. Wilsonella australis; Hallmann, 1912: 239. Not Clathria australis Whitelegge, 1901: 90. Not Plumohalichondria australis; Whitelegge, 1901: 90, p1.11, fig.14. MATERIAL. LECTOTYPE: AMZ957. Note: referred Crellithe, synonym of Crella incrustans var. arenacea (Hallmann, 1914c: 411). Plumohalichondria australis Whitelegge, 1901: 90-92, p1.11, fig.14a,b [New South Wales coast]; Whitelegge, 1907: 492 [New South Wales coast]. MATERIAL. HOLOTYPE: AMG9042. Note: referred Crellidae, synonym of Crella incrustans (Hallmann, 1912: 160). Isodictya beanii Bowerbank, 1866: 274, 334, 335 [Britain]; Gray, 1868: 164; Schmidt, 1870: 77; Bowerbank, 1874: 147, p1.58, figs 1-6. Amphilectus beanii; Vosmaer, 1880: 115. Clathria beanii; Ridley, 1881: 485, 486; Bowerbank, 1882: 13, 23, 150; Topsent, 1890c: 203. Victoria]. Plumohalichondria incrustans; Dendy, 1896: 42. Plumohalichondria mammillata; Carter, 1885f: 355; Ridley & Dendy, 1887: 156, p1.30, fig.4, p1.47, fig.4. Crella incrustans-, Hallmann, 1912: 152-156, p1.23, figs 2-3, p1.24, text-figs 28-34. MATERIAL. HOLOTYPE: BMNH1886.12.15.123, paratypes 1886.12.15.249, AME651, AMZ957. Note: referred to CrelCrella. Cornulum johnsoni de Laubenfels, 1934: 15. Clathria johnsoni; Van Soest, 1984b: 104 [possible synonym of Clathria]. MATERIAL. HOLOTYPE: USNM Note: referred to Coelosphaeridae, possibly Cornulum (imperfectly known). Echinonema levis Lendenfeld, 1888:220 [Port Jackson, New South Wales]. Plumohalichondria australis, in part; Whitelegge, 1901: 65, 91,92; Whitelegge, 1902a: 212. Crella incrustans var. levis; Hallmann, 1912: 164-167, textfigs 33-34. MATERIAL. LECTOTYPE: AMZ959, paralectotype AMG9708. Note: referred to Crellidae, synonym of Crella incrustans (Hallmann, 1912: 164). Clathria loveni Fristedt, 1887: 458, p1.25, figs 70-72, p1.30, fig.24 [Cape Jakan, Siberian Arctic Ocean]; Lambe, 1900: 160. [?] Esperella loveni; Lambe, 1895: 123, p1.4, fig.1 (W coast, North America). Esperia loveni; de Laubenfels, 1936a: 120 [note]. MATERIAL. HOLOTYPE: unknown, fragments BMNH1927.2.22.2. Note: referred to Mycalidae, Mycale. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 179zyxwvutsrqponm Plumohalichondria microcionides Carter, 1876: 236, p1.12, fig.!!, p1.15, fig.30 [between Scotland and Faroe Is]. Clathria microcionides; Vosmaer, 1880: 154. Plocamionida microcionides; Alander, 1942: 53 [Sweden]. Not Hymeraphia microcionides Carter, 1876: 390. Not Plocamia microcionides; Topsent, 1891a: 529, 544-545. MATERIAL. HOLOTYPE: BMNH1954.3.9.173. Note: referred to Anchinoidae, Plocamionida. Clathria mollis Kirkpatrick, 1903: 249-250, p1.5, fig. 15, p1.6, fig. 16 [East London Coast, Natal]. Pronax mollis; Levi, 1963: 66. Not Wilsonella mollis; Hallmann, 1912: 243. Not Clathria dura var. molls; Hentschel, 1911: 370. MATERIAL. HOLOTYPE: BMNH1902.11.16.32. Note: referred to Anchinoidae, Pronax. MATERIAL. HOLOTYPE: LMJG 15517; fragment BMNI-1 1867.3.11.29. Note: referred to Dictyoriellidae, Dictyonella. Clathriaprocumbens Brondsted, 1923; Brondsted, 1926:329 [probably a typographical error for Clathrina (Calcarea), and possibly synonym of Ascetta procumbens Lendenfeld, 1885a]. Antherochalina quercifolia Keller, 1889: 383-384, p1.23, fig.34 [Red Sea]. Querciclona quercifolia; de Laubenfels, 1936a: 46 [note]. MATERIAL. HOLOTYPE: ZMB429, fragment BMNH1908.9.24.179. Note: referred to Axinellidae, Phakellia. Clathria raphida Hechtel, 1976:244; Van Soest, 1984b: 153. Note: cited in a list of Brazilian endemic sponges; attributed to Boury-Esnault (1973); probably a misprint for Cliona rhaphida Boury-Esnault, 1973. Clathria morisca Schmidt, 1868: 9, 41, 43, p1.2, fig.7 [Algiers, Mediterranean]; Vosmaer, 1880: 150-151 [Algiers]; Antherochalina renieroides Lendenfeld, 1887b: 788, p1.28, Topsent, 1902: 329; Topsent, 1938: 11; Desqueyrouxfigs 18, 23 [New Zealand]. Faundez & Stone, 1992: 10, 35. MATERIAL. HOLOTYPE: BMN1-11886.8.27.449. Referred to Diciyoclathria morisca; Topsent, 1920b: 18-21 [re-examinaAxinellidae, synonym of Phakellia flabellata (Carter) (see tion of holotype]; Topsent, 1925: 660-661, p1.8, fig.! [Gulf of Hooper, 1991). Naples]; Topsent, 1928a: 301-302, p1.3, fig.3 [Porto Santo, Azores]; Levi, 1959: 134, text-fig.27, p1.5, fig.! [Rio de Oro, Hymeraphia tuberosocapitata Topsent, 1890b: 68 [Azores, Gulf of Guinea]; Levi, 1960b: 761-762, text-fig.15 [var. Atlantic]; Topsent, 1892a:113, p1.11, fig.6 [Atlantic]. anisotyla; SW. Cape of Naze, W. Africa]; Sara, 1960a: 462 Cionanchora tuberosocapitata; de Laubenfels, 1936a: 108 [Ischia, Mediterranean]. [note]. Artemisina mediterranea Babic, 1921: 87 [Adriatic]; Babic, Clathria tuberosocapitata; Van Soest, 1984b: 7 [generic synonymy for Cionanchora]. 1922: 258, text-fig.3; Burton, 1930a: 528.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Myxilla banyulensis, in part, Topsent, 1892b: 23; Topsent, Discorhabdella tuberosocapitata; Boury-Esnault, Pansini & 1902: 351, 363, 366; Cotte, 1903:423. Uriz, 1992: 2-6. cf. Clathria coralloides and C. compressa; Vosmaer, 1935a: MATERIAL. HOLOTYPE: MOM040323, fragment 626 [intermediate form]. MNHNDT938. referred to Hymedesmiidae, Discorhabdella. MATERIAL. HOLOTYPE: MNHN DT2170, fragment BMNH 1868.3.2.21. Note: Synonym of Antho involvens (Levi, Clathria (Wilsonella) Carter, 1885 1960a: 57). Hooper & Wiedenmayer, 1994 Clathriaoroides Schmidt, 1864:35, p1.4, figs 1-2 [Quarnerno, Adriatic]; Carter, 1875: 195; Vosmaer, 1880: 155; Dendy, 1889a: 41; Desqueyroux-Faundez & Stone, 1992: 10, 35, 103. ? Oroidea adriatica; Gray, 1867: 520. Chalinopsis oroides; Schmidt, 1870: 60. Ophlitaspongia oroides; Bowerbank, 1874: 10. Age/as oroides; Riitzler, 1965: 34 [Banyuls]; Boury-Esnault, 1971: 322 [Banyuls]; Laubier, 1966 [Banyuls]; PulitzerFinali, 1983: 534 [Mediterranean]. MATERIAL. HOLOTYPE: LMJG 15957, fragment BMNH1868.3.2.22. Note: referred to Agelasidae, Agelas. Artemisina paradoxa Babic, 1921:87; Babic, 1922:258-261, p1.8, fig.6, text-fig.c [Adriatic]; Levi, 1960a: 85-86 [Adriatic; with question] Clathria paradoxa; Burton, 1930a: 528. Dictyoclathria morisca; Topsent, 1925: 660; Ristau, 1978: 585-586 [note on affinities]. MATERIAL. HOLOTYPE: unknown. Note: synonym ofAntho involvens (Topsent, 1925: 660). Clathria pelligera Schmidt, 1864: 34, p1.3, fig.13 [Lesina, Adriatic]; Desqueyroux-Faundez & Stone, 1992: 10, 36, 103. Rhaphidostyla pelligera; Burton, 1935b: 652; Sara, 1958: 246-247, fig.15 [Gulf of Naples]. Stylotella pelligera; Topsent, 1925: 638; Boury-Esnault, 1971: 328 [Banyuls]. Dictyonella pelligera; Pulitzer-Finali, 1983: 545. Wilsonella Carter, 1885f: 366 (not Hallmann, 1912:242). Clathriopsamma Lendenfeld, 1888: 227. Aulenella Burton & Rao, 1932: 345. TYPE SPECIES. Wilsonella australiensis Carter, 1885f: 366 (by monotypy). DEFINITION. Sand grains and foreign spicules partially or completely replacing coring spicules inside fibres; coring spicules same or very similar geometry to auxiliary spicules located outside fibres; skeletal architecture reticulate. REMARKS. Microcionids with sand and foreign debris incorporated into fibres ('arenaceous sponges') are a predominant feature of S Australian waters. Of the 17 species in Wilsonella or a synonym, 14 are valid and 6 are known to live in this region including 2 new species. This group of arenaceous microcionids is well known as Clathriopsamma (Hooper, 1990a; Hooper & Levi, 1993a). However, the inclusion of C. (W) australiensis in this group means that Wilsonella (1885) takes precedence over Clathriopsamma (1888). MEMOIRS OF THE QUEENSLAND MUSEUM 180zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ A FIG. 81. Clathria (Wilsonella) abrolholensis sp.nov. (holotype NTMZ3218). A, Auxiliary style. B, Echinating acanthostyles. C, Palmate isochelae. D, Wing-shaped toxas. E, Section through peripheral skeleton (hatches foreign spicules). F, Australian distribution. G, Preserved holotype. H, Holotype. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 181 Clathria (VVilsonella) abrolhosensis sp. nov. (Figs 81-82, Plate 2C) MATERIAL. HOLOTYPE: NTMZ3218 (fragment QMG300584): N. edge of Pelsart Is, Houtman Abrolhos, WA, 28°47.2'S, 113°58.5'E, 10.vii.I987, 22m depth, coll. J.N.A. Hooper (SCUBA). HABITAT DISTRIBUTION. 22m depth; on an Acropora reef; Houtman Abrolhos, WA (Fig. 81F). DESCRIPTION. Shape. Fistulose, with multiple fistules composed of erect, bulbous-cylindrical digits, single or fused together with adjacent digits, each tapering at base and apex, thickest in apical portion, attached directly to substrate without stalk; each digit between 75-140mm long, up to 45mm maximum diameter. Colour Pale beige-yellow alive (Munsell 2.5Y 8/4), dark brown in ethanol. Oscules. Single, large, apical oscule on apex of each digit, 10-15mm diameter in life, with slightly raised membraneous lip surrounding oscule; oscules nearly completely contractile upon preservation; pores not observed in life or preserved. Texture and surface characteristics. Compressible, spongy, relatively easily torn; surface glabrous, even, without any ornamentation. Ectosome and subectosome. Discrete, erect bundles of auxiliary styles, relatively closely packed together, arising from ends of peripheral fibres, with tangential layer of auxiliary styles connecting adjacent bundles; erect spicules protrude only a short distance through surface; mesohyl matrix light in ectosomal region; subectosomal region slightly cavernous, reticulate. Choanosome. Almost regular, ovoid reticulation of fibres and spicule tracts forming ovoid, square or rectangular meshes, 220-360vm diameter; spongin fibres moderately light, relatively homogenous in size, 40-70Rm diameter, but with differentiated primary and secondary tracts; primary ascending fibres cored by multispicular tracts of auxiliary styles, interconnected by secondary, pauci- or multispicular transverse tracts of auxiliary styles; fibres relatively heavily echinated by acanthostyles; sparse core of detritus in primary ascending fibres only, mostly foreign spicules; mesohyl matrix moderately heavy, lightly pigmented; choanocyte chambers small, oval, 40-901im diameter, usually lined by isochelae. Megascleres. Principal spicules absent. Auxiliary spicules coring fibres and forming dermal skeleton moderately thin, straight or slightly curved near base, with slightly subtylote bases, hastate or slightly telescoped points, and apical and basal spination on most spicules. Length 1244151.7)-164,m, width 3-(4.1)-6p.m. Echinating acanthostyles short, thin, claviform, sharply pointed, slightly subtylote bases, lightly and evenly spined, with relatively large recurved spines. Length 71-(76.3)-86Rm, width 3-(4.7)-7p,m. Microscleres. Isochelae of 'typical' palmate form, with straight shaft, lateral alae completely fused to shaft, very long, broad front ala. Length 14-(15.7)-18Rm. Toxas wing-shaped, slender, with pronounced central curvature, slightly reflexed arms. Length 48488.4)-11211m, width 0.841.6)-2.0p.m. ETYMOLOGY. For Houtman Abrolhos. REMARKS. This species is a sibling species of C. (Wilsonella) australiensis having basal and apical spines on auxiliary styles. It was first assigned to the Western Australian subspecies C. australiensis spinulata Hentschel (1911) but subsequent re-examination of Hentschel's (1911) syntype (ZMB4446) found that he omitted several crucial characters from his description (and that his species was not substantially different from typical C. australiensis). Conversely, C. (W) abrolhosensis differs from that species in several respects, warranting its recognition as a distinct taxon. Growth form in C. (W) abrolhosensis is endolithic, consisting of discrete bulbous-cylindrical digits arising from a partially burrowing, encrusting massive base, and with large terminal oscules (superficially resembling syconoid calcarean growth forms) (whereas C. (W) australiensis has lobate or clavulate morphologies); there is only one class of auxiliary spicule (as compared with two); fibre reticulation is small, close-set, nearly regular, and predominantly ovoid (whereas in C. australiensis meshes are elongate and cavernous); spongin fibres are poorly developed (compared with well developed fibres); there is very little detritus in fibres, and these are completely confined to primary ascending fibres and more-or-less restricted to foreign spicules without sand (C. (W) australiensis has abundant detritus in all fibres, including sand grains, and this is a prominent feature of the skeleton); and spicule dimensions differ between both species. Further comparisons are given below under remarks for C. (W) australiensis. This species strictly belongs to Clathria (Dendrocia) in having only a single undifferentiated category of structural megasclere within MEMOIRS OF THE QUEENSLAND MUSEUM 182zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 82. Clathria (Wilsonella) abrolholensis sp.nov. (holotype NTMZ3218). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Spined base and point of auxiliary style. E, Echinating acanthostyles. F, Acanthosty le spines. G, Wing-shaped toxas. H, Palmate isochelae. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 183 TABLE 15 Comparison between present and published records of Clathria (Wilsonella)australiensis Carter. All measurements are given in rim, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE choanosomai auxiliary styles Lectotype Paralectotype (BMNH1886. (BMNH1886. 12.15.43) 12.15.284) 92(119.8)152 x 2.5(3.6)4 105-(125.6)152 x 2.5(3.2)4 Holotype of C. lobosa (AMG9053) Holotype of T. ramsayi (AMG8820) Specimen Specimen Specimen Paratype of (N=1) (Abrolhos Is) (I-evi,1967) var. spinulata (NWAustralia) (New (NCIQ66 (ZMB4446) C4266C) Caledonia) 92(122.6)146 x 1.5(2.7)-4 106-(127.4)173 x 2.5(3.7)-5 112(118.8)135 x 2-(3.3)5 121-(149.0)163 x 2.5(3.7)4.5 89(111.4)175 x 2.5(4.1)-8 Subect^mal 92(122.6)148 x 2-(2.9)auxiliary styles 4 101(118.3)155 x 1.5(2.9)4 94(119.6)98(117.2)141 x 2-(2.7)- 139 x 2-(2.8)3.5 4 108(120.1)134 x 2.5(3.7)-6 118(154.8)95(123.1)172 x 3-(4.1)- 164 x 2-(2.8)4.5 5 Echinatini 49-(59.8)-68 acanthosty es x 2-(3.6)-4.5 45-(64.0)-77 x 2-(3.8)-4.5 54-(62.6)-78 x 2-(3.5)-4.5 46(58.4)-70 x 2.5-(3.4)4.5 46-(57.7)-66 x 4-(6.1)-9 59-(67.6)-74 x 3-(3.9)-6 11-(14.7)-18 12-(13.2)-16 12-(13.8)-16 12-(13.4)-15 12-(13.8)-16 72-(89.4)105 x 1-(1.6)2.5 0.8 Chelae Toxas 35-(61 .2)-89 28-(53.4)-75 32-(52.3)-74 24-(45.4)-75 x 0.5-(0.7)-1 x 0.5-(0.8)-1 x 0.5-(0.8)-1 x 0.5-(0.8)-1 fibres, but it is included here in Clathria (Wilsonella) having detritus in fibres and spined terminations on auxiliary spicules similar to C. (W.) australiensis. This further illustrates the unclear generic boundary between many nominal genera of Microcionidae, and supports the recognition of a wide definition for Clathria. Clathria (Wilsonella) australiensis (Carter, 1885) (Figs 83-84, Plate 2B, Table 15) Wilsonella australiensis Carter, 1885f: 366; Hallmann, 1912: 239; Hallmann, 1920: 768; Burton, 1934a: 599. Clathria australiensis; Dendy, 1896: 33; Whitelegge, 1901: 66, 84, 85, p1.11, fig.12; Hooper & Levi, 1993a: 1242, table 4; Hooper & Wiedenmayer, 1994: 275. Clathria australiensis var. spinulata Hentschel, 1911: 374-375, text-fig. 47; Dendy, 1922: 71. Clathriopsamma lobosa Lendenfeld, 1888: 149; Whitelegge, 1901: 85; Hallmann, 1912: 239. Thorecta ramsayii Lendenfeld, 1888: 149. Sigmatella corticata var. elegans Lendenfeld, 1888: 199-201; Lendenfeld, 1889b: p1.40, fig.7. Not Clathria australiensis; Levi, 1967b: 22, p1.2, fig.D, text-fig.6. Not Ophlitaspongia australiensis Ridley, 1884a: 442. Not Echinochalina australiensis; Thiele, 1903a: 961962. MATERIAL. LECTOTYPE: BMNH1886. 12.15.43 (fragment AMG2805): Port Phillip, Vic, 38 ° 09'S, 144 ° 52'E, 12m depth, coll. J.B. Wilson (dredge). PARALECTOTYPE: BMNH1886.12.15.284: same locality. HOLOTYPE of C. lobosa: AMG9053: Port Jackson, NSW, 33 °51'S, 151 ° 16'E. HOLOTYPE of T. ramsayi: AMG8820: same locality. SYNTYPES of var. spinulata: ZMB4446, HM numbers unknown: 125-180 x4 51(60.4)-81 x 2.5-(3.7)5.5 65-80 x 4 14-(15.2)-17 12-(14.6)-19 14-15 28-(39.4)-59 x 0.5-(0.6)- 43-(55.6)-88 x 0.5-(0.8)1.5 40-75 x 1-2 Bunbury and Middleton Beach areas, WA, 33 ° 20'S, 115 °36'E, coll. W. Michaelsen & R. Hartmeyer (dredge). OTHER MATERIAL: NSW - QMG301447, QMG301458, AMG975, BMNH1886.12.15.288, AMZI412, fragment NTMZ1526, AMZ3176, AMZ3199, AMZ3140, AMZ4283. VICTORIA NMVRN748. S. AUST. - SAMTS4098 (fragment NTMZ1646), SAMTS4116 (fragment NTMZ1627). WESTERN AUSTRALIA - WAM623-81(1) (fragment NTMZ1710), QMG300622 (NCIQ66C-4266-C). HABITAT DISTRIBUTION. Subtidal to 160m depth; on rock, sand and algal bed substrates; known only from Australia: throughout temperate Australian waters - from Houtman Abrolhos, Perth, Bunbury (WA) (Hentschel, 1911; present study), Nuyts Archipelago, St Vincent Gulf (SA) (present study), Port Phillip (Vic) (Carter, 1885f; present study), Bega, Jervis Bay, Port Hacking, Port Jackson, Botany Bay, N. Sydney, Tweed River region (NSW) (Lendenfeld, 1888 Whitelegge, 1901; present study), and extending into the tropics as far as Low Isles, GBR, Qld. (Burton, 1934a) (Fig. 83G). DESCRIPTION. Shape. Lobate, lobate-digitate, club-shaped, thickly lamellate, or thickly encrusting-bulbous growth forms, up to 140mm high, 110mm wide, with subspherical, tubular, bulbous or flabellate digits, up to 75n un high, 45mm wide, 25mm thick. Colour. Pale orange alive (Munsell 7.5 YR 8/6), pale pink, brown or yellow preserved. Oscules. Large oscules, up to 4mm diameter, on apical or lateral margins of surface digits/lobes. Texture and surface characteristics. Surface optically even, microscopically rugose, with a whitish arenaceous, slightly hispid dermal membrane. - MEMOIRS OF THE QUEENSLAND MUSEUM 184zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 83. Clathria(Wilsonella)australiensis (Carter) (NTMZ1627). A, Subectosomal auxiliary subtylostyles. B, Choanosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Oxhorn toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype BMNH1886.12.15.43. I, NMVRN748. Ectosome and subectosome. Ectosome membraneous, without specialised dermal megasclere, varying from densely arenaceous, with most or all dermal megascleres obscured by large sand grains and spicule fragments, or with light palisade of erect brushes of subectosomal auxiliary subtylostyles, arising from ascending primary choanosomal fibres; sand grains at surface slightly larger than those in choanosome. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 185 FIG. 84. Clathria (Wilsonella) australiensis (Carter) (A-B, AMG9053, QMG301447). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Palmate and modified isochelae. G, Oxhorn toxas. H-I, Continuum in basal and apical spination of auxiliary subtylostyles. MEMOIRS OF THE QUEENSLAND MUSEUM 186^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA differences in interpretation of character importance between the present study and that of Hallmann (1920) (see also Hooper, 1990a). Unlike Clathria (Dendrocia), in which there is only a single category of coring and extra-fibre megasclere, most Clathria (Wilsonella) have more than one form of auxiliary style, one coring the fibres (choanosomal megascleres) and one outside of fibres (subectosomal megascleres). In some cases (e.g., C. (W) australiensis, C. (W) ensiae sp. nov.), these spicules are only slightly different in geometry, although showing clear differences in patterns of spination; in others (e.g., C. (W) reticulata, C. (W) mixta) these spicules have different geometry; whereas in one (C. (W) abrolhosensis) there are no apparent differences which is interpreted as a convergence or subsequent loss of a spicule category. Clathria (Dendrocia) and Clathria (Wilsonella) can also be distinguished by their skeletal architecture being predominantly plumose in the former and reticulate in the latter. In material listed above most of the larger auxiliary styles without spines on points appeared to be located within spongin fibres (i.e., choanosomal spicules), whereas most of the auxiliary styles with spines on both bases and points were found predominantly outside fibres, strewn within the mesohyl, and in the dermal skeleton (i.e., subectosomal spicules). However, this observation is difficult to verify in all cases because of the dense core of sand particles in fibres. In this species both sorts of spicules are classed as auxiliary styles due to their very similar geometry: true principal styles are absent (i.e., Wilsonella s.s.). The principal diagnostic characteristics of C. (W) australiensis and its affinities with other species are discussed elsewhere (see Table 19 and remarks for C. (W) tube rosa). This species differs from most Australasian Clathria (Wilsonella) in having spinose extremities on both the bases and points of quasidiactinal auxiliary subtylostyles. In this regard it is similar to its sibling species, C. (W) abrolhosensis from the Houtman Abrolhos, REMARKS. Clathria (W) australiensis is the WA (which is sympatric with C. australiensis) only species retained by Hallmann (1920) in Wil- and the allopatric species C. (W) rugosa, from sonella; other species were transferred to either New Caledonia (Hooper & Levi, 1993a; Table Clathria or Paradoryx, depending on whether 19). Spicule geometry (megascleres and toxas) they had palmate or arcuate-like isochelae are useful in distinguishing these species of microscleres. Some of these, however, are further Clathria (Wilsonella) (Table 19). Levi's (1967b) material from New Caledonia, allocated here to Clathria (Dendrocia) or Clathria (Thalysias), depending on their ec- described as C. australiensis, was referred to C. tosomal skeletons, choanosomal skeletal struc- (W) rugosa (Hooper & Levi, 1993a), based on ture and spicule diversity (i.e., there are differences in spicule geometry (especially shape Choanosome. Choanosomal skeleton irregularly reticulate, with clearly differentiated primary and secondary spongin fibres, forming a vaguely longitudinal reticulation with cavernous meshes; spongin fibres well developed; primary ascending fibres marginally thicker, producing ascending lines abundantly cored by detritus, lightly cored by auxiliary styles, heavily echinated by acanthostyles, particularly at fibre nodes; smaller secondary spongin fibres mainly transverse, connecting with primary elements, with no or little detritus, paucispicular tracts of auxiliary styles and lightly echinated by acanthostyles; auxiliary megascleres coring fibres occupy only a small proportion of fibre diameter; detritus in fibres consists of scattered sand grains and spicule fragments; mesohyl matrix heavy; choanocyte chambers oval to eliptical, some appear to be paired, and chambers lined by microscleres. Megascleres (Table 15). Choanosomal auxiliary styles coring fibres differ from subectosomal auxiliary styles only in being slightly thicker and lacking characteristic apical microspines of the latter. Coring spicules relatively thin, straight, smooth, mostly hastate, with slightly subtylote and occasionally microspined bases. Subectosomal auxiliary styles, dispersed between fibres and in dermal skeleton, straight, usually hastate, subtylote bases with microspines on both points and bases. Echinating acanthostyles small, evenly spinose or with granular, vestigial spines, slightly subtylote bases, fusiform points. Microscleres (Table 15). Palmate isochelae relatively large, unmodified. Toxas oxhorn, uncommon, rare in some specimens, often forming trichodragmata, usually with wide, angular, central curves and slightly reflexed points. Larvae. Ovoid, incubated parenchymella larvae, up to 350p.m diameter seen in some material. Associations. Commensal polychaetes common in many specimens; Abrolhos specimen with epizootic zoanthids on surface. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO ^ 187 and spination of auxiliary spicules), and the latter having flabellate-digitate growth forms. The quasidiactinal (strongylotes) modifications of some of the subectosomal auxiliary spicules in both species, with weakly spined points and bases, is a unique trait within the Wilsonella group, but is also known in a few other species of Microcionidae (C. (Dendrocia) pyramida, C. (Thalysias) major, C. (Clathria) chelifera). These quasidiactinal spicules are convergent in geometry with true tornotes and strongyles found in other groups such as Iophonidae (see remarks for Megaciella and Acarnus in the introductory section above). Hentschel (1911) created a subspecies spinulata for material from WA, indicating that it differed substantially from typical forms of the species in spicule dimensions (particularly isochelae), and supposedly lacked toxas. However, Hentschel's type has toxas and isochelae dimensions are within the size range of other S Australian populations, and all other aspects (growth form, fibre characteristics, amount and form of detritus in fibres) are identical between populations. Hentschel (1911) suggested that spinulata differed from other populations in having spines on both the points and bases of auxiliary spicules, but this feature occurs in all other populations, and the WA population is conspecific with C. (W) australiensis. Clathria (W.) australiensis is widely distributed throughout temperate Australia, from the Houtman Abrolhos on the W coast (30 °S), around S Australia to the Tweed River (28 °S). Burton (1934a) recorded this species from Low Isles, (16°S), but his voucher material has not been examined, and his description is not detailed enough to differentiate between C. (W.) australiensis and C. (W) rugosa from New Caledonia. Recent collections from this region do not include the species. Examination of many other specimens (cited above), from many locations throughout Australia, showed that C. (W) australiensis is a heterogeneous species. Most regional populations differ from each other slightly in spicule geometry or spicule size, but there is not enough available material of any of these populations to determine whether these differences are consistent within populations, nor are there any features in any of these regional populations worthy of distinguishing separate taxa (cf. Hentschel, 1911). Widespread sampling of regional populations, collection of samples for biochemical and genetic studies, and determination whether or not observed morhological dif- ferences correlate with any fixed genetic differences is a worthwhile study for the future. Clathria (Wilsonella) claviformis Hentschel, 1912 (Figs 85-86, Table 16, Plate 2D) Clathria claveformis Hentschel, 1912: 366-367, p1.19, fig.29. MATERIAL. HOLOTYPE: SMF 1504 (fragment MNHNDCL2238): Sungi Manumbai, near Kapala Sungi, E. side of Aru I., Arafura Sea, Indonesia, 6°S, 134 ° 50'E, 28.iii.1908, 23m depth, coll. H. Merton (dredge). OTHER MATERIAL: NT - NCIQ66C0528-0, NTMZ3082. HABITAT DISTRIBUTION. 18-23m depth; sand, rock and dead coral substrate; Timor Seas (Fig. 85G). Also Arafura, SE. Indonesia. DESCRIPTION. Shape. Claviform, 160-220mm high, 60-150mm wide, with small cylindrical base and stalk, and one or more lobate, clubshaped, or bulbous digits, up to 80mm long, 65mm diameter. Colour. Pale 'dusty' beige and red-brown mottled colour alive (Munsell 2.5Y 8/6 and 5R 8/4), yellow-grey preserved. Oscules. Large oscules, up to 5min diameter, on ends of bulbous digits, occasionally in crevices between digits. Texture and surface characteristics. Surface even, with distinct, partially arenaceous, skinlike covering, and with several longitudinal ridges on sides of digits and between bulbous digits; texture rubbery, compressible, sandy. Ectosome and subectosome. Ectosome lightly and evenly arenaceous, hispid, with plumose tufts of subectosomal auxiliary subtylostyles protruding through surface, sometimes in dense brushes; subectosome cavernous, with sparsely cored, arenaceous, ascending primary spongin fibres forming large meshes in periphery. Choanosome. Choanosomal skeleton irregularly reticulate, cavernous, with light spongin fibres and spicule tracts; spongin fibres divisible into primary and secondary components; primary fibres relatively large, with very little fibre spongin content, cored by both large and small sand grains, Foraminifera and paucispicular tracts of choanosomal principal styles, becoming sinuous during towards periphery; secondary fibres consist of uni- or paucispicular spicule tracts bound together by collagenous spongin and debris; primary fibres moderately echinated, slightly heavier towards periphery; secondary fibres with 188zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 85. Clathria (Wilsonella) claviformis Hentschel (holotype SMF1504). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype. ^ 189 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR FIG. 86. Clathria (Wilsonella) claviformis Hentschel (holotype SMF1504). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Palmate isochelae. E, Echinating acanthostyle. F, Acanthostyle spines. G, Base of principal subtylostyle. H, Bases of auxiliary subtylostyles. I, Accolada toxas. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 190zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA no or few echinating acanthostyles; mesohyl matrix heavy, granular, with abundant microscleres and detritus scattered throughout, usually lining small oval choanocyte chambers. Megascleres (Table 16). Choanosomal principal styles thin, slightly curved, fusiform, with rounded or slightly subtylote, lightly microspined bases. Subectosomal auxiliary subtylostyles hastate, thin, mostly straight, with slightly subtylote bases, usually microspined. Echinating acanthostyles slightly subtylote, with few, dispersed, large spines on base and apex, usually with aspinose neck; spines located on point usually more recurved than those on base. Microscleres (Table 16). Palmate isochelae large, abundant, with some twisted examples. Toxas thin, with slight angular central curvature and straight, unreflexed points. REMARKS. C. (W) claviformis is relatively easily differentiated from other arenaceous species by its skeletal architecture, fibre characteristics and spiculation (see remarks for C. (W) tube rosa below, and compare spicule dimensions between species in Table 19). Hentschel's (1912) mention of similar isochelae geometry in C. (W) clavifonnis and other microcionid species such as C. (Dendrocia) pyramida is misleading, since the latter species has arcuate-like chelae whereas those of C. (W.) claviformis are simply palmate. The nearest relative of C. (W) claviformis is probably C. (W) tube rosa, especially in specific features of its arenaceous ectosome. Clathria (VVilsonella) ensiae sp. nov. (Figs 87-88, Table 17, Plate 2E) MATERIAL. HOLOTYPE: NTMZ3561 (NCIQ66C2384-0 (fragment QMG05004): Marion Reef, off Edithburg, S. Yorke Peninsula, SA, 35°09.5'S, 137°48.0'E, 10.ii.1989, 6m depth, coll. NCI (SCUBA). PARATYPE: NTMZ3821 (NCIQ66C-3744-L) (fragment QMG300270): Trap Reef, Bicheno, Tas., 41°51.7'S, 148°18.6'E, 30m depth, 26.ii.1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION. 6-30m depth; on sand covered rock substrate, with algae and seagrasses on patch reef; Yorke Peninsula (SA), E coast (Tas) (Fig. 87F). DESCRIPTION. Shape. Erect, digitate, flabellate growth form, 205-350mm high, 70-150mm wide, with multiple branches usually aligned in one plane, composed of long, slender, flattened or cylindrical, bifurcate digits, 70-190mm long, up TABLE 16. Comparison between present and published records of Clathria (Wilsonella) claviformis Hentschel. All measurements are in p,m, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Choanosomal principal styles Subectosomal auxiliary styles Echinating acanthostyles Holotype Specimens (N=2) (SMF1504) 2584308.6)-364 x 6- 235-(287.2)-338 x 7(8.4)-10 (9.6)-13 164-(292.0)-371 x 3- 169-(289.2)-375 x 3(5.1)-8 (5.2)-7 64-(74.0)-82 x 479-(88.8)-103 x 4(6.4)-8 (6.4)-8 Chelae 16-(18.4)-22 14-(16.8)-20 Toxas 28-(140.8)-266 x 0.8-(1.1)-1.5 44-(113.6)-2I8 x 0.8-(0.9)-1.2 to 18nun diameter, frequently fused near their basal ends, attached to a common base or on a short cylindrical stalk. Colour. Red-brown alive (Munsell 2.5R 8/8), darkening in air (2.5R 6/10), brown preserved. Oscules. Large oscules, up to 4nun diameter, scattered evenly over all surfaces of digits, in life slightly raised above the surface with a membraneous lip. Texture and surface characteristics. Surface even, optically smooth, broken only by raised oscules; texture firm, compressible, rubbery. Ectosome and subectosome. Ectosome heavily arenaceous, with peripheral spongin fibres fully packed with mostly sand particles and some foreign spicule fragments (holotype; vice versa in paratype), and with sparse plumose tracts of subectosomal auxiliary styles, confined completely below surface; spongin fibres in subectosomal region ascend to surface, plumose, fully arenaceous, with plumose brushes of subectosomal auxiliary spicules. Choanosome. Choanosomal skeleton irregularly reticulate in axis, plumo-reticulate near periphery, clearly divided into primary, ascending fibres, 60-130Rm diameter, and secondary, connecting, transverse spongin fibres, 30-70p..m diameter; primary fibres fully arenaceous, incorporating both sand and foreign spicule fragments, and a sparse core of choanosomal auxiliary styles amongst the debris; secondary fibres without sand particles, with some foreign spicules, and also with a light core of choanosomal auxiliary spicules; echinating acanthostyles not abundant on fibres, usually echinating fibres at acute angles, directed towards surface; mesohyl matrix moderately heavy, with few foreign spicules and ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 191 TABLE 17. Comparison between present and published records of Clathria (Wilsonella) ensiae sp.nov. All measurements are given in 1.1m, denoted as range (and mean) of spicule length x spicule width (N=25). Subectosomal auxiliary styles Holotype (NCIQ66C2384I) 69-(111.4)-132 x 3.5-(4.2)-6 106-(130.7)-147 x 1.5-(2.6)-3.5 97-(132.0)-154 x 2(2.4)-3.5 1888) (Figs 89-90, Table 18) F,chinating acanthostyles 28-(40.7)-54 x 2.5(3.1)-4 48-(61.1)-70 x 3.5(4.3)-5.5 Clathriopsamma reticulata Lendenfeld, 1888: 227; Chelae 11-(14.2)-16 13-(14.4)-16 Toxas absent SPICULE Choanosomal principal styles Paratype (NCIQ66C3744L) 87-(108.6)-120 x 4(4.4)-5.5 could also be included in C. (Dendrocia) due to close resemblance between choanosomal and subectosomal styles, both classed here as auxiliary spicules (i.e., Wilsonella s.s.). However, like C. (W) australiensis, those styles coring fibres differ subtly in their terminations from those styles outside of fibres. Clathria (Wilsonella) reticulata (Lendenfeld, Hallmann, 1920: 771. Clathria reticulata; Hooper & Wiedenmayer, 1994: absentzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB - very little sand; choanocyte chambers large, 180480p.m diameter, oval or eliptical. Megascleres (Table 17). Choanosomal auxiliary megascleres straight or slightly curved towards base, slightly subtylote bases, hastate, abruptly pointed; very similar geometry to subectosomal auxiliary styles but shorter, slightly thicker. Subectosomal auxiliary styles short, slender, straight or rarely slightly curved at centre, slightly subtylote bases, hastate, stepped points. Echinating acanthostyles short, slender, straight, slightly subtylote bases, fusiform points, spined all over but spines slightly heavier on base and point. Microscleres (Table 17). Palmate isochelae relatively long, slender, unmodified. Toxas absent. ETYMOLOGY. Phonetic acronym in the National Cancer Institute (NCI), in appreciation of the AIMS group who provided the author with unrestricted access to all their sponge collections. REMARKS. There are some differences between the two specimens of C. (W) ensiae in the size of acanthostyles (Table 17). Similarly, primary spongin fibres of the holotype are predominantly cored with sand particles, whereas in the paratype foreign spicules are more abundant than sand, but in all other respects both these specimens are identical, and these observed differences are considered to be relatively minor. C. (W) ensiae differs from other species of the Wilsonella group primarily in its flattened-flabelliform, erect, bifurcate, branching growth form and in having auxiliary styles with peculiar hastate, telescoped points. Other features such as skeletal structure and spicule dimensions can also be used to distinguish allied species (Table 19). Like C. (W) australiensis, the present species 275. Not Echinochalina reticulata; Whitelegge, 1907: 506, p1.45, fig.25; Hallmann, 1912: 287, p1.30, fig.2, text-fig.66. Not Dictyocylindrus reticulatus Carter, 1881a: 377. Not Rhaphidophlus reticulatus; Hallmann, 1912: 177. MATERIAL. LECTOTYPE: AMG9135 (dry): E. coast of Australia, no other details known. PARALECTOTYPES: AMZ457: E coast of Australia, no other details known. BMNH1925.11.1.576 (dry): Manly Beach, NSW, 33°49'S, 151°18'E, no other details known. HABITAT DISTRIBUTION. Ecology unknown; central E coast (NSW) (Fig. 890). DESCRIPTION. Shape. Subspherical, reticulatebranching growth form, 80-150mm high, 5595nun wide, composed of lobate, bifurcating, sometimes anastomosing tubular digits with rounded margins, 30-50nun long, up to 18mm diameter. Colour. Dark brown in ethanol. Oscules. Not seen (available material dry and surface contracted). Texture and surface characteristics. Surface shaggy, reticulate; texture brittle in dry state. Ectosome and subectosome. Ectosomal skeleton lightly arenaceous, with plumose brushes or individual choanosomal principal styles protruding, together with a paratangential layer of subectosomal auxiliary subtylostyles, lying near bases of principals, and echinating acanthostyles projecting into these. Choanosome. Choanosomal skeleton irregularly reticulate, with moderately heavy, large spongin fibres forming relatively wide ovoid meshes, lined by very large, typically curved oxeote toxas; spongin fibres not easily divisible into primary or secondary components based on fibre diameter, although primary, ascending fibres contain plumose, paucispicular tracts of both principal 192zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 87. Clathria (Wilsonella) ensiae sp.nov. (holotype NTMZ3561). A, Choanosomal auxiliary subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Palmate isochelae. E, Section through peripheral skeleton. F, Australian distribution. G, Paratype QMG300270. and auxiliary megascleres; secondary, transverse, connecting fibres without coring spicules; both sorts of spongin fibres contain a light core of detritus, especially small sand grains; echinating acanthostyles very abundant, including peripheral fibres; mesohyl matrix heavy, darkly pigmented, with abundant microscleries, especially bundles of whispy, sinuous toxas (toxodragmata); extra-fibre auxiliary megascleres organised into ascending subdermal tracts, with few loose spicules scattered between fibres. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 193 FIG. 88. Clathria (Wilsonella) ensiae sp.nov. (holotype NTMZ3561). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Palmate isochela. E, Echinating acanthostyles. F, Acanthostyle spines. G, Base of principal subtylostyle. H, Bases of auxiliary subtylostyle. MEMOIRS OF THE QUEENSLAND MUSEUM 194^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA BC n E m..zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA LO CV 23 FIG. 89. Clathria (W ilsonella) reticulata Lendenfeld (lectotype AMG9135). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Oxeote toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Paralectotype BMNH1925.11.1.576. ^ 195 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ FIG. 90. Clathria (Wilsonella) reticulata Lendenfeld (paralectotype AMZ457). A, Choanosomal skeleton. B, Peripheral skeleton. C-D, Fibre characteristics. E, Echinating acanthostyle. F, Acanthostyle spines. G, Base of principal subtylostyle. H, Bases of auxiliary subtylostyles. I. Oxeote toxa. J, Palmate isochelae. 196zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Megascleres (Table 18). Choanosomal principal subtylostyles well differentiated from auxiliary spicules, mostly straight, fusiform, with profusely microspined, slightly subtylote bases. Subectosomal auxiliary subtylostyles thin, fusiform, straight, slightly curved, sometimes sinuous, with minutely rnicrospined, subtylote bases. Echinating acanthostyles with rounded bases, relatively even spination, although basal and distal portions slightly more heavily spined than points. Microscleres (Table 18). Palmate isochelae differentiated into 2 size classes, smaller with approximately 75% of contort forms. Toxas extremely abundant, long, thin, sinuous, characteristically oxeote, with only very slight or no central curvature and straight tapering points. REMARKS. This species is poorly known from 3 specimens but differentiated from other species in the Wilsonella group having only a very light core of detritus in fibres, usually composed of small sand and spicule particles, differentiated primary and secondary fibres, and prominent bundles of sinuous toxas lining aquiferous chambers (Fig. 90). Affinities with other species (Table 19) are discussed elsewhere (remarks under C. (W) tuberosa and C. (W) australiensis). In some respects (growth form, geometry of some spicules, presence of two sizes of isochelae with contort forms) the species is closest to C. (W) tuberosa, but toxa geometry is quite different between these two species. Clathria (Wilsonella) tuberosa (Bowerbanlc, 1875) (Figs 91-93, Table 19, Plate 2F) Microciona tube rosa Bowerbank, 1875: 281, 282, 286; Vosmaer, 1935a: 607. Clathria tuberosa; Ridley, 1881: 121; Ridley, 1884a: 444-445, p1.42, fig.d; Hentschel, 1912: 365-366; Hooper & Wiedenmayer, 1994: 275. MATERIAL. HOLOTYPE: BMNH1877. 5.21.1312: Straits of Malacca, Malaysia, vicinity of 2 ° N, 102 ° E, coll. Capt. Parish (dredge). OTHER MATERIAL: INDONESIA - SMF978 (fragment MNHNDCL2346). QLD - BMNH1881.10.21.325, BMNH1882.2.23.198, 253, 283, 334. NT - AMZ4559 (RRIMP FN989), NTMZ777, NTMZ809, NTMZ920, NTMZ933, NTMZ946, NTMZ1091, NTMZ2087, NTMZ2400, NTMZ2708, QMG303336, NTMZ2157, NTMZ2189, NTMZ1959, NTMZI980, NTMZ1987, NTMZ2020, NTMZ2098, QMG303428, NTMZ2107, NTMZ234, NTMZ235, NTMZ236 - NTMZ2496, NTMZ540, NTMZ107, NTMZ112, NTMZ128, NTMZ554. WESSEL ISLANDS; NT - NTMZ3955. TABLE 18. Comparison between present and published records of Clathria (Wilsonella) reticulata (Lendenfeld). All measurements are given in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). Lectotype (AMG9135) Paralectotype (AMZ457) Paralectotype (BMNH 1925.11.1.576) Choanosomal 173-(209.4)258 x 7-(9.1)principal styles 11 182(240.2)289 x 8-(9.3)11 227-(251.5)296 x 8-(9.4)ii SPICULE Subectosomal I63-(236.8)1984243.4)187-(259.0)auxiliary 324 x 3-(4.0)-6 322 x 3-(4.2)-5 309 x 3-(4.6)-7 styles 52-(61.7)-71 x 54-(63.6)-75 x 65-(67.8)-72 x Echinating acanthostyles 4-(7.2)-9 6-(7.3)-9 3-(6.5)-9 Chelae I 5-(6.6)-9 4-(5.8)-7 6-(6.1)-9 Chelae II 12-(14.4)-17 13-(14.8)-17 14-(15.2)-17 Toxas 238-(402.7)684 x 0.8(2.4)-4 45-(231.3)433 x 0.5(1.9)-3.5 302-(427.7)593 x I -(2.2)-4 HABITAT DISTRIBUTION. Semi-encrusting on rock, dead or live coral heads, epizootic on other sponges and gorgonians; usually associated with shallow coral reef habitats; 2-19m depth range; prevalent in the tropical, Australian and Indo-Malay shallow water macrobenthic community, extending as far south as 13°S latitude: Torres Strait (FNQ) (Ridley, 1884a), Bynoe Harbour, Darwin Harbour, Coral Bay, Port Bremer, Wessel Is (NT). Also Straits of Malacca (Bowerbank, 1875; Ridley, 1881) and Aru Is, Indonesia (Hentschel, 1912). DESCRIPTION. Shape. Subspherical, predominantly bulbous growth form, 60-135mm diameter, less often club-shaped with apical lobate digits, or pseudo-vasiform on low stalk with convoluted, apical, lobate digits; surface projections (or branches) rounded lobate, relatively close-set, attached to common centre, which in turn is usually attached to substrate by a small peduncle; lobate digits usually bifurcate with rounded margins. In life lobes prominently bulbous, evenly rounded; after preservation lobes become slightly flattened and angular. Colour. Live pigmentation dusty pale pink-red (Munsell 2.5R 6/10) to pink (5RP 8/6), with a darker choanosome (5RP 7/8); lighter ectosomal colouration due to arenaceous nature of ectosome; dessicated colouration darkens to brown (2.5Y 8/2), red-brown (5Y 8/4), or red-purple (5RP 3/6), as paler ectosome collapses. In situ, subdermal ridges and canals red-pink in life, showing darker choanosomal pigmentation. Oscules. Exhalant pores variable in diameter, ranging from 1-3mm, each with prominent, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 197 Clat hria ( Wilsonella) t uberosa (Bowerbank) (NTMZ2157). A, Choanosomal principal subtylostyle. B, FIG. 91. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Echinating acanthostyle. C, Subectosomal auxiliary subtylostyles. D, Longer accolada and smaller wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Known Australian distribution. H, Specimen of Hentschel (1912) SMF978. I, Specimen of Ridley (1884) BMNH1882.2.23.198. J, NTMZ107 in sit u. 198zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 92. Clathria (Wilsonella) tuberosa (Bowerbank) (NTMZ2708). A, Choanosomal skeleton. B, Fibre characteristics (x164). C, Echinating acanthostyles. D, Acanthostyle spines. E, Bases of principal subtylostyles. F, Palmate isochelae. G, Base of auxiliary subtylostyle. H, Longer accolada toxas. I. Smaller wing-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 199 SEASON TOTAL SAMPLES SAMPLES WITH LARVAE WET 3 2 PREDRY 5 5 DRY 10 0 PREWET 10 5 FIG. 93. Clathria (Wilsonella) tube rosa (Bowerbank). Seasonal production of incubated larvae in populations from NW Australia. membraneous, raised lip (collapses upon preservation); oscules confined to apex of lobate digits, with subdermal canals and grooves radiating away from pores in cruciform pattern. Texture and surface characteristics. Surface optically smooth, even in situ, distinctly membranous; ectosomal membrane transparent or slightly opaque when intact, stretched across adjoining lobes, with darker subdermal pigmentation and fibre reticulation clearly visible below; subdermal grooves and minute subdermal canals produce a more-or-less microscopically reticulate surface; upon dessication ectosome collapses to become optically reticulate, distinctly arenaceous, with convoluted ridges and conules, and large amounts of clear mucous usually produced; texture stiffly compressible, arenaceous, harsh to touch, minutely hispid. Ectosome and subectosome. Ectosomal skeleton heavily arenaceous, with delicate traces of sand coring peripheral fibres, through which protrude sparse tufts of subectosomal auxiliary styles, usually raised on low surface conules; special ectosomal megascleres absent; subectosomal auxiliary subtylostyles also form tangential or paratangential tracts perpendicular to dermal crust, intermingled with foreign particles auxiliary spicules variable in size, but no distinct localisation of smaller or larger forms; subectosomal region obscured by abundant sand grains coring peripheral subdermal fibres; individual extra-fibre auxiliary styles are intermingled amongst sand and fibres in subdermal region, sometimes forming dense paratangential plumose brushes, ascending to ectosome, but usually producing sparse tangential subdermal tracts; subdermal tracts clustered tightly around fibres and sand matrix, bound together with abundant collagen; on peripheral fibres, choanosomal principal spicules produce plumose brushes, sometimes protruding through surface, but usually only obvious in places where ectosome has collapsed and peripheral fibres are closest to surface. Choanosome. Choanosomal skeleton irregularly reticulate, with light spongin fibres fully cored by sand grains and fewer choanosomal principal megascleres, the latter in rows of 6-10 abreast in larger fibres; spicule fragments also common amongst detritus, particularly haplosclerid oxeas; spongin fibres heavily echinated by acanthostyles; fibre branching produces irregular oval meshes, 50-(334)-600p.m diameter, with irregular eliptical choanocyte chambers (38121p,m diameter), with light mesohyl matrix and abundant microscleres, without sand or any megascleres; spongin fibres not clearly divisible into primary or secondary elements, but thinner fibres (30-551J,m diameter) have coring megascleres more visible (fewer detrital particles); larger spongin fibres 70-(104)-230p„m diameter; megascleres core fibres in paucispicular tracts, slightly more heavily aggregated in thicker fibres but partially obscured by sand particles; mesohyl matrix in axis light, with little foreign debris or auxiliary megascleres. Megascleres (Table 19). Choanosomal principal subtylostyles slightly curved towards basal end, occasionally straight, with heavily microspined bases, tapering to sharp fusiform points. Subectosomal auxiliary subtylostyles straight, variable in size, usually with microspined, prominently subtylote bases, sharply tapering, fusiform points. Acanthostyles very variable in length and width, straight, subtylote, fusiform, evenly spi nose with granular spines (thinner spicules) or heavy thorn-like spines (thicker spicules). Microscleres (Table 19). Palmate isochelae typically very abundant, incompletely divided into two size classes, with some twisted smaller examples. Toxas abundant, thin, usually long, without reflexed points, only slightly curved at centre, although smaller examples may have more angular central curvature; occurring individually or more often in toxodragmata within mesohyl matrix. Larvae. Incubated parenchymella larvae were recorded in only 17% of specimens, collected from Darwin and Cobourg Peninsula regions, NT, during May, September, October and December, suggesting a possible breeding period during the wetter months (Fig. 93). Larvae orange-brown pigmented, oval - elongate, ranging from 165280 x 110-160iLm. All larvae contained juvenile megascleres scattered throughout central portion of mesohyl, usually with heavy collagen. In the few adult sponges seen incubating larvae, the ^ MEMOIRS OF THE QUEENSLAND MUSEUM 200zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA mesohyl was usually lightly orange in colour, whereas in non-fecund specimens the mesohyl matrix was invariably pigmented light brown. Associations. Scyllid poly chaete worms (Typosyllis spongicola) relatively common (31% of specimens examined). Morphological variation. Gross morphology: characteristic, relatively consistent, varying only in elongation of basal stem and clump of digits on apex, ranging from low spherical bulbs (69%), club-shaped (16%), or convoluted, semivasiform growth forms (15%). Live colouration: consistent, only slight variation in pinkish hue. Extosomal skeletal structure: extends from densely arenaceous with few dermal spicule brushes (39%), arenaceous with abundant protruding plumose subectosomal brushes (35%), to arenaceous with plumose brushes of subectosomal spicules and single choanosomal styles erect on surface (26%). Subectosomal skeletal structure: varies from well-developed, plumose ascending spicule tracts composed of subectosomal auxiliary styles (27%), few plumose extra-fibre brushes with most extra-fibre spicules closely bound by collagen (31%), or with all extra-fibre spicules simply bound closely but external to spongin fibres (42%). Choanosomal skeletal structure: relatively consistent, ranging from simply irregularly reticulate (54%) to regularly reticulate (46%), with oval meshes (88%) or less commonly longitudinally elongated meshes (12%). Mesohyl matrix: lightly pigmented (58%), moderately heavily pigmented (23%), or heavily pigmented collagen (19%). Echinating acanthostyles: forming dense (42%), moderately dense (50%), or only lightly echinating fibres (8%). Megasclere geometry: consistent; majority of choanosomal subtylostyles with microspined bases, 8% of specimens with smooth subtylote bases. Acanthostyles vary only in dimensions. Microsclere geometry: consistent although proportion of contort versus unmodified TABLE 19. Comparison between morphological characteristics of some Australasian arenaceous Clathria (Wilsonella) species, based on present and published records. All measurements are given in p.m. MATERIAL CHARACTER 1 2 3 4 5 6 Shape massive subspherical tubular elongate subspherical tubular clay iform digitate flabellate digitate fan Digits lobate lobate clubs/bulbs cylindrical or flattened cylindrical lobate Live colour pale orange pale orange red-brown red-brown unknown unknown Skeletal architecture irreg. relic irreg. relic regular relic irreg. retic. irreg. relic. Differentiated primary/seconda ry fibre system irreg. relic, axis, plumo-,regular relic. subect. yes yes yes yes yes no Choanosomal styles 89-175 x2.5-8 smooth or spined base 96-214x4-6.5 smooth or spined base 235-364 x6-13 spined base 69-132 x3.5-6 smooth base 172-388 x6-16 smooth base 192-298 x6-11 spined base Subectosomal styles 92-172 x1.5-5 spined apex and base 99-148 x2.5-3.5 spined apex and base 164-375 x3-8 spined base 97-154 x1.5-3.5 smooth base 162-541 x3-7 smooth base 160-325 x2.5-5.5 spined base 49-85x2-6 52-63x3-5.5 64-103x4-8 28-70x2.5-5.5 63-94x4-8 78-94x3-9 11-19 12-15 14-22 11-16 14-18 12-18 absent absent absent absent absent 3-8 twisted 28-89 x 0.5-1.5 angular centre 43-59 x0.8-1.5 angular centre 28-266 x 0.8-1.5 slightly angular centre absent 57-74 x1-2 rounded centre 91-435 x0.8-2 straight at centre Acanthostyles Large chelae Small chelae Toxas Material: 1. W. australiensis (Carter) - present study. 2. C. lobosa Lendenfeld (1888:149) (=W. australiensis) zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFED holotype AMU G9053; Port Jackson,NSW. 3. W. clavifonnis (Hentschel) - present study. 4. W. ensiae, sp.nov. - present study. 5. W. ramosa (Lindgren, 1897:482- schizotype BMNH 1929.11.26.48. 6. W. mixta (Hentschel, 1912:298) - holotype SMF 974. - 201 ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR fibres. Nevertheless, the two species are closely related. Clathria (W) australiensis has choanosomal fibres divided into ascending and radial primary elements, fully cored by detritus, with less heavily cored secondary transverse elements. Subectosomal auxiliary subtylostyles have spines on both bases and points, typically with a prominent terminal spine/point and also smaller spines occuring at least part of the way along the shaft. Choanosomal subtylostyles are also auxiliary spicules, with smooth or spined bases only. Acanthostyles are pointed and tapering; there is only one size of isochelae; and toxas have strongly curved (rounded or sharply angular) central curves. Clathria (W) rugosa is a sister species of C. (W) australiensis, differing in having prominent subectosomal drainage canals ('astrorhizae'). Subectosomal auxiliary styles are also spined on both ends, but spines are perched only on the very extremity of the spicule point (not on the shaft); acanthostyles have bulbous points; there is onlyzyxwvutsrqp small isochelae varies from 0-20% of contort spicules (12%), 20-40% (19%), 40-60% (24%), 60-80 (35%), to 80-10% of spicules (12%). Spicule dimensions: Few specimens atypical but variation apparently random with no statistical significance between specimens irrespective of seasonal or geographical distribution of samples. REMARKS. C. (W) tuberosa is distinctivein the field: pink colour, bulbous growth form, soft texture. However, it is more difficult to differentiate descriptively. Pertinent differences are: Choanosomal architecture and fibre characteristics of C. (W) reticulata are identical to those of C. (W) tuberosa; acanthostyles are as equally abundant in both species, but many acanthostyles have heavier spines on the distal part than on points; and there are two sizes of isochelae, 75% of the smaller being contort. Thus, the major features distinguishing the two taxa are the straight or sinuous oxeote toxas in C. (W) reticulata, which are never present in C. (W) tuberosa, and the light deposits of debris into MATERI AL CHARACTER Shape 7 8 9 fan palmate- massive spherical massive spherical digitate tubular globular cylindrical Digits 10 11 elongate massive spherical subspherical globular tubular cupriform lobate lobate lobate lobate pale pink or pink- pale pink or pink- pale pink or pink- pale pink or pink- red red red red Live colour bt.red-orange Skeletal architecture irreg.retic. irreg.retic. irreg.retic. irreg.retic. yes none none none 134-159x4.5-8 167 - 278 x4 - 11 144 - 286x4 - 8 l72-302x6-8.5 smooth base most spined bases most spined bases most smooth bases Differentiated primary/ seconda ry fibre system Choanosomal styles 162-206 x2.5-4 Subectosomal styles spined apex and Acanthostyles 58-91 x4-7 none 1334226.3)-343 x4-(8.2)-14 most spined bases Small chelae lobate tubular unknown partial 173-296 x7-11 all spined bases 163-324 x3-7 154-324 x2-5 148-223 x2-5 zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA x1.5-(4.1)-8 159 - 333 x2-6 spined bases spined bases spined bases 44-97 x4-9 43-75 x3-8 69-88 x3.5-5 base spined bases spined bases (6.7)-11 52-75 x3-9 12 - 17 11 - 19 11 - 16 13 - 19 10 - (13.3) - 18 14 - 17 absent 4-8 twisted 4-7 twisted 6-8 rare 4-(6.1)-9 twisted 4-9 twisted 24-122 x0.8-3 Toxas reticulate branches 129-(228.9)-375 60-(79.9)-112 x4- Large chelae 12 slight curve at centre 55-255 x0.5-1.5 rounded centre 95-215 x0.5-1.5 angular& rounded centre 30-(140.3)-388 81-94 x1-2 rounded centre x0.4-(1.I )-2 45-684 x0.5-4 sinuous oxeote rounded centre Material: 7. W. rugosa (Hooper & Levi, 1993) - New Caledonian population (included for comparative purposes). 8. W. t uberosa (Bowerbank, 1875:281); holotype BMNH 1877.5.21.1312; Straits of Malacca (19=25). 9. W. t uberosa; Ridley, 1884a:444; BMNH 1881.10, 21.325, 1882.2.23.198, 253, 283, 334; Tones Strait (N=125). 10. W. t uberosa; Hentschel, 1912: 365; SMF 978; Arafura Sea (N= 25). 11. W. t uberosa; present material - NW Australia (N= 650). 12. W. r et iculat a (Lendenfeld) - .present st udy. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC 202zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM a single small category of isochelae; and toxas are only slightly curved at their centre. Clathria (W) clavtformis, C. (W) ensiae, and C. (W) ramosa are different from these other species in spicule geometry, spicule size, skeletal architecture and fibre characteristics. These IndoAustralasian species are relatively easy to distinguish from their gross morphology and field characteristics although their skeletal characters are usually at least partially obscured by the incorporation of sand into fibres. With the exception of C. (W.) australiensis species of Clathria (Wilsonella) have relatively restricted, mostly allopatric distibutions: C. (W) tube rosa is from N Australia and SE Asia; C. (W) claviforrnis from the Arafura and Timor Seas; C. (W) mixta from 2 disjunct populations in the Arafura Sea and the S. Arabian coast; C. (W) ramosa from the Java Sea; C. (W) ensiae in S Australia and Tasmania; C. (W) reticulata SE Australia. C. (W.) australiensis ranges from SW Australia to S Queensland, whereas its cryptic sibling C. (W) rugosa is restricted to New Caledonia. OTHER SPECIES OF CLATHRIA (WILSONELLA) Clathria (Wilsonella)cercidochela (Vacelet & Vasseur, 1971) Clathriopsammacercidochela Vacelet& Vasseur, 1971: 104105, text-fig.62, p1.3, fig.! [Tulear, Madagascar]. MATERIAL. HOLOTYPE: MNHNDJV24. W Indian Ocean. Clathria (Wilsonella) ferrea (de Laubenfels, 1936) Fisherispongiaferrea de Laubenfels, 1936b: 460, fig.44. [Atlantic coast of Panama]. Clathria (Microciona) ferrea; Van Soest, 1984b: 101-103, text-fig.40, table 4 [Curacao]. Clathria ferrea; Zea, 1987: 172, text-fig.60 [Colombian Caribbean]. Microciona ferrea; Pulitzer-Finali, 1986: 149 [West Indies]. MATERIAL. HOLOTYPE: USNM22239. Caribbean. Clathria (VVilsonella) foraminifera (Burton & Rao, 1932) Aulenella foraminifera Burton & Rao, 1932: 345-346, p1.18, fig.11, text-fig.11 [Gaspar Straits, Java Sea]. MATERIAL. HOLOTYPE: IMP790/1. E Indonesia. Clathria (VVilsonella) lindgreni sp. nov. Clathria ramosa Lindgren, 1897: 482-483; Lindgren, 1898: 308-309, p1.17, fig.9, p1.18, fig.15, p1.19, fig.16 [Belitung I., Java Sea]; Hentschel, 1912: 367. Thalysias ramosa; de Laubenfels, 1936a: 105. Not Rhaphidophlus ramosus Kieschnick, 1896: 533; Kieschnick, 1900: 53-54, p1.45, figs 47-50. Not Echinoclathria ramosa; Hallmann, 1912: 277, p130, fig.3. Not Wilsonella ramosa; Hallmann, 1912: 243, 298. Not Colloclathria ramosa Dendy, 1922: 74-76. cf. Microciona prolifera tropus spinosa; Vosmaer, 1935a: 642. MATERIAL. HOLOTYPE: NHRM (fragment BMNH1929.11.26.48). Indonesia. Clathria (Thalysias) ramosa (Kieschnick, 1896) has priority. Clathria (Wilsonella) litos Hooper & Levi, 1993 Clathria (Clathriopsamma) litos Hooper & Levi, 1993a: 1243-1246, figs 9-10 [New Caledonia]. MATERIAL. HOLOTYPE: QMG301269. SW Pacific. Clathria (Wilsonella) mixta Hentschel, 1912 Clathria mixta Hentschel, 1912: 298, 367, 368, p1.13, fig.8, p1.19, fig.30 [Aru I., Arafura Sea]; Burton, 1959a: 244 [S. Arabian coast]. Thalysias mixta; de Laubenfels, 1936a: 105. cf. Clathria lobata or Clathria ulmus; Vosmaer, 1935a: 649. MATERIAL. HOLOTYPE: SMF 974 (fragment MNHNDCL2280). Indonesia, Arabian Gulf. Clathria (Wilsonella) pseudonapya (de Laubenfels, 1930) Clathriopsamma pseudonapya de Laubenfels, 1930: 28; de Laubenfels, 1932: 96-97, text-fig.57 [Pacific Grove, California]; Sim & Bakus, 1986: 10 [California]. MATERIAL. HOLOTYPE: USNM21436. PARATYPE BMNH1929.8.22.19. NE Pacific rim. Clathria (Wilsonella) rugosa Hooper & Levi, 1993 (Table 19) Clathria (Clathriopsamma) rugosa Hooper & Levi, 1993a: 1237-1243, figs 7-8, tables 4-5 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300278 (fragment NTMZ3880). PARATYPE QMG300696 (fragment NTMZ3889). SW Pacific. Clathria (Microciona) Bowerbank, 1862 Microciona Bowerbank, 1862b: 1109. [Abila] Gray, 1867: 539 [preocci. [Aaata] de Laubenfels, 1930: 271preocci. Anaata de Laubenfels, 1932: 89. Axocielita de Laubenfels, 1936a: 118. Cionanchora de Laubenfels, 1936a: 108. Fisherispongia de Laubenfels, 1936b: 460. Folitispa de Laubenfels, 1936a: 119. Hymantho Burton, 1930a: 503. Hymeraphia, in part, Hentschel, 1912: 377; not Hymeraphia Bowerbank, 1864: 189. Leptoclathria Topsent, 1928a: 298. Ophlitaspongia Bowerbank, 1866: 14; not Ophlitaspongia of authors. Paratenaciella Vacelet & Vasseur, 1971: 103. Pseudanchinoe Burton, 1929a: 433. Seriatula Gray, 1867: 515 Sophax Gray, 1867: 521. Wetmoreus de Laubenfels, 1936a: 112. DEFINITION. Persistently encrusting growth form, with hymedesmoid skeletal architecture consisting of a basal layer of spongin, typically REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 203 with ascending, plumose, non-anastomosing, spongin fibre nodes, and megascleres embedded and erect on basal layer; ectosomal skeleton with only a single undifferentiated category of auxiliary megasclere. TYPE SPECIES. Microciona atrasanguinea Bowerbank, 1862b: 1109 (by subsequent designation of Bowerbank, 1864: 188). REMARKS. Of 118 named species described in, or subsequently referred to Microciona or one of its synonyms, 103 appear to be valid, and 7 are recorded from Australasia, including 2 new species. Clathria (Microciona) aceratoobtusa (Carter, 1887) (Figs 94-95, Table 20, Plate 3C) Microciona acerato-obtusa Carter, 1887: 62, 67, 83, p1.5, figs 7-10; Dendy, 1896: 18; Hentschel, 1911: 348, text-fig. 32a-f. Axocielita aceratoobtusa; de Laubenfels, 1936a: 118. Clathria aceratoobtusa; Rudman & Avern, 1989: 335; Hooper & Wiedenmayer, 1994: 266. cf. Microciona prolifera; Vosmaer, 1935a: 608, 637. MATERIAL. HOLOTYPE: LFM (confirmed destroyed during WWII): Kadan Kyun (King I.), Mergui Archipelago, Andaman Sea, Burma. NEOTYPE: NTMZ3676: NW. side of N. I., Ko Wao Yai Group, vicinity of Ko Samui, Gulf of Thailand, 9°46.7'N, 99°40.3'E, 12m depth, 6.vi.1990, coll. J.N.A. Hooper (SCUBA). OTHER MATERIAL: NSW - NTMZ2835 (fragment QMG300543), NTMZ3125. QLD QMG303089, QMGL713 (fragment NTMZ1536). SAHUL SHELF, WA - QMG301083, QMG301188. INDONESIA - BMNH1946.11.25.244. HABITAT DISTRIBUTION. Coral rubble, rock and bivalve substrata; intertidal-14m depth; Shark Bay, Cartier I., Hibernia Reef, Sahul Shelf (WA) (Hentschel, 1911, present study); Cairns and Shelburne Bay (FNQ) (present study); Sydney and Iluka (NSW) (Rudman & Avern, 1989, present study) (Fig. 940). Also IndoMalay Archipelago — Andaman Sea (Burma) (Carter, 1887), Gulf of Thailand (present study). DESCRIPTION OF NEOTYPE. Thinly encrusting on bivalves, up to lnun thick; colour orangered alive (Munsell 1 OR 6/12); firm texture; oscules not seen; surface microscopically hispid, with choanosomal principal styles protruding up to 100p,m from ectosome; subectosomal auxiliary styles lie paratangential to surface, in bundles or individually; choanosomal skeleton leptoclathriid, with principal styles and echinating acanthostyles embedded in and perpendicular to basal spongin fibres; principal styles form plumose brushes, and both sorts of spicules also scattered individually in skeleton; mesohyl matrix heavy, dark brown, granular, with incorporated detritus, numerous toxas and auxiliary styles dispersed; principal choanosomal styles long, fusiform, rounded or very slightly subtylote, with smooth or minutely spined bases (length 175-548p.m, width 11 22 p,m); subectosomal auxiliary subtylostyles polytylote, with microspined swollen bases (length 264-387p.m, width 1.5-4.5p,m); echinating styles short, slightly curved, robust, with prominently swollen, usually rnicrospined bases and smooth shafts (length 128-183p,m, width 5-12p,m); palmate isochelae small, relatively homogeneous in size, with many twisted forms (9-14p.m long); toxas short, thickest at centre, tapering to sharp, slightly reflexed points (length 58-92p,m, width 25.5p,m). - DESCRIPTION. Shape. Thinly encrusting, contiguous or discrete mats on rock or coral substrata, covering up to 120mm2, 0.4-2mm thick. Colour. Bright orange-red alive (Munsell lOR 6/12-14), grey-brown in ethanol. Oscules. Small exhalant apertures unevenly distributed over surface, up to 1.5mm diameter, slightly raised or flush with surface; small membraneous lip surrounding oscules when alive, collapsing in air. Minute inhalant pores irregularly dispersed, producing slightly reticulate appearance. Texture and surface characteristics. Firm, mucusy alive, minutely hispid; surface with irregularly dispersed, bifurcate subdermal drainage canals meandering from oscules. Ectosome and subectosome. Ectosomal skeleton hispid, with points of large choanosomal principal styles protruding up to 200p.m from surface, occurring individually or in paucispicular plumose brushes of about 5 spicules; subectosomal auxiliary styles usually lie paratangential to surface, sometimes forming tangential tracts lying immediately subdermal; auxiliary styles arise from skeleton at oblique angles, rarely protruding through ectosome; tracts of auxiliary spicules originate in basal half of skeleton, with 8-12 spicules per tract. Choanosome. Choanosomal skeletal architecture hymedesmoid in thin sections, microcionid in thicker regions, with a relatively thick layer of heavy spongin fibre lying on basal substrate, 2243 um diameter; bases of principal and echinating styles embedded in basal spongin, perpendicular to substrate, individually or in plumose bundles; ^ MEMOIRS OF THE QUEENSLAND MUSEUM 204zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA D1 FIG. 94. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Microciona) acerat oobt usa (Carter) (NTMZ2835). A, Choanosomal principal subtylostyles. B, Polytylote subectosomal auxiliary subtylostyles. C, Echinating subtylostyles. D, Oxhorn toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 205 FIG. 95. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Microciona) acerat oobt usa (Carter) (QMG303089). A, Choanosomal skeleton. B, Close view of hymedesmoid skeleton. C, Bases of choanosomal principal subtylostyles. D, Polytylote base of auxiliary subtylostyle. E, Palmate isochela. F, Oxhorn toxa. MEMOIRS OF THE QUEENSLAND MUSEUM 206^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA TABLE 20. Comparison between present and published records of Clathria (Microciona) aceratoobtusa (Carter). All measurements are given in p,m, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Choanosomal principal styles Subectosomal auxiliary styles Neotype (NTMZ3676) 175(386.0)548 x11(15.5)-22 264-(324.3)387x 1.5(3.8)4.5 Hentschel (1911) up to 408 x 8 168 x 4 Specimens (N=6) 235-(352.8)492 x12.5(17.2)-22 217(379.8)443 x 3.5(4.3)-6 97-( 194 x 5.2(6.8)-9 128-(149.9)183 x 5-(7.2)12 from 64 x 5 Chelae 9-(12.2)-14 9-12 10.5-(13.5)-16 Toxas 58-(71.5)-92 x 2-(3.1)-5.5 35-92 x 3-4 18-(57.3)-84 x 0.8-(2.2)-3.5 styles in thick sections basal spongin fibres form small erect nodes, 22-35 p.m thick, up to 48p,m high, enveloping bases and parts of spicule shafts; mesohyl matrix heavy, granular, darkly pigmented, incorporating irregularly dispersed sand grains and other foreign debris, numerous toxas occurring singly or in dragmata, more-or-less ascending tracts of subectosomal auxiliary megascleres, and fewer isochelae; choanocyte chambers minute, ovoid but rarely seen, 1218p,m diameter, mostly obscured by heavy collagen; large subectosomal cavities, 110-145p.m diameter, visible where inorganic substrate is fragmented and discontiguous. Megascleres (Table 20). Choanosomal principal subtylostyles long, thick, fusiform, typically curved in basal third, with slightly subtylote mostly smooth, less often microspined bases. Subectosomal auxiliary subtylostyles usually long, straight, thin, fusiform, with prominently swollen, smooth or microspined bases. Echinating subtylostyles entirely smooth or occasionally with lightly microspined bases, small, thick, fusiform, slightly curved or straight. Intermediates between echinating and principal styles also occur. Microscleres (Table 20). Palmate isochelae small, with long lateral alae fused to shaft for most of its length; chelae relatively common, of a single size class, homogeneous in size and geometry, approximately 70% with contort shafts. Toxas very abundant, oxhorn, with slightly rounded central curves, straight or slightly reflexed points; central part thickest whereas tips taper to fine points. Associations. On the NSW coast this species has been found in association with pairs of nudibranchs grazing on the sponge, Rostanga arbutus (AMC151078, 154589) (W.B. Rudman, pers.comm.). These predators are identical in their live colouration to the sponge, presumably utilising the sponge's carotenoid pigments. REMARKS. This species was originally recorded from Mergui Archipelago and by Hentschel (1911) from Shark Bay, WA. The holotype was destroyed during WWII (its absence from the LFM collections has been checked by Shirley Stone, BMNH, pers.comm.); the neotype comes from an area in Thailand relatively close to the type locality. The first record of the species in the Pacific Ocean is also made here. Previous published descriptions of this species are relatively poor and non-discriminatory; some attributes of the type material are still uncertain. Carter (1887) did not give any spicule dimensions, but his figures indicate that Mergui specimens are very similar to present material. There are some minor differences between my material and descriptions by Carter (1887) and Hentschel (1911). Hentschel's specimens from Shark Bay were thickly encrusting with stoloniferous, mammiform surface processes. Choanosomal architecture varied from leptoclathriid, with a thin layer of spongin lying on the substrate, to microcionid in thicker regions, with fibre nodes and single, non-anastomosing columns of spongin arising from the substrate. Principal styles were fusiform, prominently subtylote, often with microspined bases. Palmate isochelae were frequently contort. By comparison, Carter (1887) reported the holotype had principal styles with hastate or styloid points, and their bases were completely smooth and only slightly subtylote. Similarly, there was no mention in Carter's description whether isochelae were modified (contort). Vosmaer (1935a) expressed doubts about the conspecificity between Carter's and Hentschel's material based on alleged differences between them in megasclere and microsclere geometries, but this criticism is unfounded. Both Carter and Hentschel reported that their specimens were thinly encrusting on living and dead serpulid worm tubes, bivalves and gastropods; colour was brown to beige preserved; toxa geometry was distinctive and identical; and echinating megascleres were entirely smooth. De Laubenfels (1936a) erected Axocielita for this species, having smooth echinating REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 207 TABLE 21. Spicule dimensions of of Clathria (Microciona) antarctica (Topsent), giving comparisons between nominotypical material and other type material. All measurements are given in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). Holotype of S. SPICULE Part of type of M. toxtfera basispinosa (MNHNDT1612) (BNMN1933.3.1739) Choanosomal principal styles 409-(519.9)-676 x 9- 293-(498.3)-618 x 9(13.8)-22 (12.6)-18 Subectosomal auxiliary styles 2I3-(424.9)-899 x 4- 252-(360.9)-503 x 4(10.5)-16 (7.2)-10 52-(110.8)-2I4 x 2.5-(6.8)-10 78-(130.1)-265 x 3(8.7)-11 Chelae absent absent Toxas 31-(46.1)-84 x 0.8(1.8)-3.0 18-(27.8)-35 x 1.5(2.3)4.0 Echinating acanthostyles megascleres (i.e., like Axociella) but also with a Microciona-like encrusting growth form. This is surprising given that Ophlitaspongia seriata, a north Atlantic species, also has an encrusting growth form, microcionid architecture and smooth echinating spicules (which Simpson (1968a) subsequently showed was a synonym of Microciona based on cytology and other characters). These arguments demonstrate that the apparent generic boundaries between microcionids based on growth form and spicule spination are tenuous at best. QMGL713 from the Cairns region, encrusting on an ascidian, has skeletal architecture, fibre characteristics and spiculation closely comparable to other material of C. (M.)aceratoobtusa but lacks toxas completely. It is difficult to confirm the identity of this specimen (i.e., because the species is largely characterised by the geometry of its toxas), but given that all other characters are the same it is included here. Although only known from few specimens it is likely that this species is widespread in the Indowest Pacific shallow-water fauna. Clathria (Microciona) antarctica (Topsent, 1917) (Figs 96-97, Table 21) Stylostichon toxiferum Topsent, 1913a: 621-622, p1.4, fig.7, p1.6, fig.14. Not Hymeraphia toxifera Hentschel, 1912: 382. Anchinoe toxifera var. antarctica; Topsent, 1917: 43, p1.4, fig.5, p1.6, fig.5. Pseudanchinoe toxifera; Burton, 1929a: 433-434; Burton, 1932a: 325; Burton, 1934b: 39; Burton, 1940: 115; Koltun, 1964a: 72; Koltun, 1976: 155, 188, figs 11-12. Clathria toxifera; Van Soest, 1984b: 129. Pseudanchinoe toxiferum; Koltun, 1976: 155, 188. Stylostichon tuberculata Burton, 1934b: 35, p1.3, fig.2, text-figs 6-9; Koltun, 1976: 188. Microciona basispinosa Burton, 1934b: 38-39, p1.5, fig.2, text-figs 11-12; Burton, 1938b: 17; Koltun, 1964a: 76; Desqueyroux, 1972: 31, figs 103-107; Desqueyroux & Moyana, 1987: 49; Dawson, 1993: 36. Clathria antarctica; Hooper & Wiedenmayer, 1994: 266. MATERIAL. HOLOTYPE: MNHNDT1612: Gough I., S. Atlantic, 40°20'S, 95°6.3'W, 200m depth, 22.iv.1904, coll. R.R.V. 'Scotia' (dredge). HOLOTYPE of M. basispitzosa: NHRM997 III (fragment BMNH 1933.3.17.39): Port Albemarle, Falkland Is, 18-30m depth, 11.ix.1902, Swedish Antarctic Expedition (dredge). HABITAT DISTRIBUTION. Deeper water rock reefs; 16-610m depth (Koltun, 1976); Antarctica - Discovery Inlet, Ross Sea, McMurdo Sound, Graham Land, Victoria Land, Enderby Land; Subantarctic — Macquarie I. (Fig. 96F). Also SW Atlantic; SW and SE Pacific — Tierra del Fuego, Falkland Is, Shag Rocks, South Georgia, Gough I., Argentina, Chile, Kerguelen, New Zealand. DESCRIPTION. Shape. Growth forms range from thickly encrusting to massive, subspherical. Colour. Brown in ethanol. Oscules. Small, 2mm diameter, on apex of surface conules. Texture and surface characteristics. Compressible; smooth surface with scattered prominent conules, translucent surface. Ectosome and subectosome. Moderately dense plumose brushes of auxiliary subtylostyles, of a single size category, arising from ends of erect fibre nodes, protruding through surface and interdispersed with long principal spicules. Choanosome. Skeletal architecture microcionid, with hymedesmoid basal layer of spongin lying on substrate echinated by erect acanthostyles of various sizes, and erect fibre nodes at 200-40011An intervals; fibre nodes non-anastomosing, forming discrete skeletal columns in choanosome; erect fibre columns, 40-100p,m diameter, cored by long choanosomal subtylostyles, usually protruding through fibres in plumose bundles or individually, and also heavily echinated by smaller acanthostyles in their basal portion only; fibres form single, discrete columns of spongin and spicules for most of their length but diverge into 2 or more branches in subectosomal region, ultimately producing ectosomal spicule brushes at their ends; mesohyl matrix light, choanocyte chambers 30-50p.m diameter, numerous spherical cells, and dispersed auxiliary spicules outside MEMOIRS OF THE QUEENSLAND MUSEUM 208zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ E D. 0 C\J Clat hria ( Microciona) ant arct ica (Topsent) (holotype MNHNDT1612). A, Choanosomal principal FIG. 96. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Wing-shaped toxas. E, Section through skeleton. F, Antarctic distribution. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 209 fibres also form plumose columns; mesohyl collagenous. Megascleres (Table 21). Choanosomal principal subtylostyles very long, slender, with fusifonn points, slightly curved towards apical end, subtylote or slightly subtylote bases, smooth or microspined bases. Subectosomal auxiliary subtylostyles short, robust, straight or slightly curved near basal end, hastate points or at least less fusiform than principal spicules, with subtylote microspined bases. Echinating acanthostyles variable size range, the larger ones clearly intermediate between principal spicules and smaller spined spicules; spicules straight or slightly curved, moderately heavily spined, evenly spined, spination becoming vestigial on larger spicules, fusiform points, subtylote bases. Microscleres (Table 21). Chelae absent. Toxas short, thick, wingshaped, with wide central curvature, curved at slight angle, slightly reflexed arms. REMARKS. Hentschel's (1912) toxifera has seniority over Topsent's (1913a) name, and hence the next available name antarctica (Topsent, 1917) is used for this species (Hooper & Wiedenmayer, 1994). Koltun (1976) proposed that M. basispinosa Burton was conspecific with S. toxiferum Topsent, and this is now confirmed. Koltun's (1976) proposed synonymy of this species and S. tuberculata Burton has not yet been corroborated (types not yet found). His proposal to include C. (Clathria) pauper Brondsted, 1927, in this taxon is rejected, the latter species having a plumoreticulate skeleton (as opposed to exclusively plumose skeleton), FIG. 97. Clathria (Microciona) antarctica (Topsent) (fragment of different pattern of spination on holotype BMNH1933.3.17.39a). A, Echinating acanthostyles and toxa. acanthostyles, two toxa morB, Plumose skeletal structure. phologies (C. (M.) antarctica MEMOIRS OF THE QUEENSLAND MUSEUM 210^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA having only one), and spicule sizes differing substantially between the two species (compare Table 21 and description of C. (C)pauper above). Clathria (Microciona) grisea (Hentschel, TABLE 22. Comparison between present and published records of Clathria (Microciona) grisea (Hentschel). All measurements are given in gm, denoted as range (and mean) of spicule length x spicule width (N=25). 1911) (Figs 98-99, Table 22) Holotype (ZMB4435) Specimen (N=1) Choanosomal principal styles 135-(198.2)-212 x 6(10.4)-12 241-(268.5)-298 x 11-(13.2)-16 Subectosomal auxiliary styles 195-(218.9)-242 x 4- 246-(264.4)-283 x 4(5.8)-8 (6.3)-8 954103.4)-116 x 6- 109-(124.6)-158 x 8(9.4)-11 (7.8)-10 SPICULE Leptosia grisea Hentschel, 1911: 353, text-fig.35. Microcionagrissa [lapsus]; de Laubenfels, 1936a: 111. Clathria grisea; Hooper & Wiedenmayer, 1994: 266 MATERIAL. HOLOTYPE: ZMB4435: NW. of Middle Bluff, Shark Bay, WA, 2548'S, 11326'E, 7-8m depth, 21.ix.1905, coll. W. Michaelsen & R. Hartmeyer (dredge). OTHER MATERIAL: WA- NTMZ2863 (fragments QMG300054, PIBOC 04-295). HABITAT DISTRIBUTION. 7-25m depth; growing on bivalves and Acropora cf. robusta; Shark Bay and Pelsart Is, Houtman Abrolhos (WA) (Fig. 98F). DESCRIPTION. Shape. Thinly encrusting, up to 3mm thick (holotype) or long cylindrical digitate sponge, 480mm long, 70mm maximum width, with few, slightly flattened, bifurcate, cylindrical branches, up to 40mm diameter, and short basal, holdfast attachment. Colour. Red-brown alive (Munsell lOR 4/10), brownish-grey preserved. Oscules. Large oscules, up to 4mm diameter, irregularly distributed on lateral sides of branches in ramose material. Texture and sutface characteristics. Surface smooth, unornamented, with distinct skin-like detachable covering; texture compressible, rubbery. Ectosome and subectosome. Ectosome heavily collagenous, up to 1801.1,m thick, including a light crust of arcuate isochelae, and with tangential fibres running longitudinally along surface; subectosomal skeleton consists of plumose brushes of auxiliary subtylostyles, in bundles, standing erect or semi-erect but not protruding beyond surface; subectosomal spicule bundles arise from ends of principal spicules erect on the substrate (in holotype) or from peripheral fibres (in specimen). Choanosome. Holotype — choanosomal skeleton hymedesmoid, with a basal layer of spongin fibre, acanthose bases of principal styles and smaller acanthostyles embedded in basal spongin, standing erect upon substrate. Specimen — choanosomal skeleton irregularly plumose, slightly reticulate, with sinuous, heavy spongin fibres, up to 220..m diameter, cored by both subectosomal auxiliary subtylostyles and principal styles, and echinated by plumose brushes of both choanosomal principal styles and echinating Echinating acanthostyles Chelae I 15-(16.8)-19 14-(15.2)-17 Chelae II 20-(22.9)-28 23-(25.5)-28 absent absent Toxas acanthostyles poking through fibres into mesohyl; mesohyl matrix light, choanocyte chambers elongate-oval, up to 150p,m diameter, with abundant arcuate isochelae dispersed throughout. Megascleres (Table 22). Choanosomal principal styles thick, long, slightly curved at centre, subtylote, with heavily spined bases and sparsely microspined shafts. Subectosomal auxiliary subtylostyles, in dermal skeleton within choanosomal fibres, long, thick, straight, with fusiform points and very slightly subtylote, smooth bases. Echinating acanthostyles long or short, relatively slender, straight, slightly subtylote, heavily spined all over spicule except for aspinose point. Microscleres (Table 22). Arcuate-like isochelae divided into two size classes, without intermediates; larger chelae with very thick, strongly curved shaft, small rounded lateral alae attached to shaft for most of its length, front ala completely free; smaller chelae with slightly curved shaft, long lateral alae only partially attached to shaft. Toxas absent. REMARKS. There are some notable differences in skeletal structure and spicule sizes between the encrusting holotype and the branching specimen described above (Table 22), but the two specimens agree so closely in spicule diversity and geometry that they are obviously conspecific. These differences may be due to the holotype being immature, having smaller spicule dimensions and a hymedesmoid skeleton, whereas the larger branching specimen still retains the ascending plumose (non-anastomosing) fibre nodes, typical of the Microciona condition. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 211 The major distinguishing features in C. (M.) grisea are the arcuate isochelae and spined principal spicules. On this basis it is surprising that de Laubenfels (1936a) did not refer it to Anaata which he created specifically for this purpose (i.e., to include species with acanthose principal spicules, echinating acanthostyles, smooth subectosomal styles and arcuate isochelae). The holotype of Leptosia grisea has a hymedesmoid choanosome, without plumose fibre nodes (and therefore strictly a member of Leptoclathria), and only a single category of auxiliary spicule. Anaata, Leptoclathria and Microciona are considered synonyms of Clathria. Clathria (M.) grisea belongs to Hallmann's (1912) spicata group of species (see comments for C. (Thalysias) lendenfeldi), having a spicate arrangement of principal and echinating spicules, which protrude through fibres in a plumose manner (Hooper et al., 1990). Clathria (Microciona) illawarrae sp. nov. (Figs 100-101, Plate 3D) MATERIAL. HOLOTYPE: QMG304572: Shellharbour, Illawarra, NSW, 34 ° 35'S, 150°52'E, 10.vi.1993, coll. L. Miller (SCUBA). HABITAT DISTRIBUTION. Shallow subtidal; on rock reef, growing over bivalves and coralline algae; central E coast (NSW) (Fig. 100G). DESCRIPTION. Shape. Thinly encrusting, 0.31.5mm thick, following contours of substrate. Colour. Pale yellowish-orange alive (Munsell 2.5Y 8/10), beige in ethanol. Oscules. Minute, less than 2mm diameter, scattered over surface, with slightly raised surrounding membraneous lip; pores very small covering entire surface. Texture and surface characteristics. Soft, compressible, easily torn; porous, opaque, even, fleshy surface, without any sculpturing or other ornamentation. Ectosome and subectosome. Single category or large subectosomal auxiliary subtylostyles form paratangential plumose brushes, protruding only slightly through surface but extending well into mesohyl. Choanosome. Skeleton microcionid, with thin basal layer of spongin lying on substrate, 3040p..m thick, containing incorporated sand grains; erect spongin fibre nodes arise at approximately 200Rm intervals along basal spongin, 20-4011m thick, 100-150p,m long, cored by erect choanosomal principal styles in uni- or multispicular tracts, up to 5 spicules per bundle, form- ing perfectly erect or slightly plumose brushes ascending to but not protruding through surface; fibre nodes discrete, not anastomosing with adjacent nodes, but some principal spicules from adjacent nodes cross within mesohyl; paratangential plumose brushes of auxiliary spicules located in several places within mesohyl, forming a tangential tract near basal spongin layer, forming stellate brushes midway along erect fibre nodes, and forming plumose paratangential brushes near surface; echinating acanthostyles relatively sparse on both basal spongin and erect fibre nodes; mesohyl matrix moderately heavy with microscleres dispersed throughout; choanocyte chambers not seen. Megascleres. Choanosomal principal styles long, thin, slightly curved or whispy near point, often bent in distal third of spicule, with smooth tapering hastate bases and fiisiform points, occasionally slightly telescoped. Length 62-(129.4)-165Rm, width 3-(3.6)-4.5p,m. Subectosomal auxiliary subtylostyles long, thin, straight, with elongated smooth subtylote bases and hastate points. Length 1764206.5)228p,m, width 1.0-(2.3)-3.0Rm. Echinating acanthostyles short, relatively thick, cylindrical, usually thickest above basal constriction, slightly spined, aspinose slightly constricted neck, slightly swollen base, rounded or fusiform point. Length 36-(53.2)-68p,m, width 2-(3.8)-6Rm. Microscleres. Palmate isochelae very small, with greatly reduced lateral alae, often no more than ridge on shaft, and small front ala complete; sometimes asymmetrical ends. Length 445.9)Toxas small, thick, u-shaped or forceps shaped, with angular central curve and non-reflexed arms. Length 6-(8.1)-11m, width 0.540.8)1.01/m. ETYMOLOGY. For the type locality. REMARKS. This species is one of the most thinly encrusting microcionids, with most sections no more than 300jLm thick. Its choanosomal skeleton is typical of Microciona (erect fibre nodes arising from a hymedesmoid basal skeleton), but subectosomal auxiliary spicules have an unusual distribution within the skeleton forming both stellate brushes around the fibre nodes and basal tangential tracts near the substrate. The species is also unusual amongst thinly encrusting microcionids in that there are no subsurface drainage canals associated with the aquiferous system, whereasthe surface is smooth, porous and fleshy. These live surface features, the MEMOIRS OF THE QUEENSLAND MUSEUM 212zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 98. Clathria (Microciona) grisea (Hentschel) (NTMZ2863). A, Choanosomal principal acanthostyles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Arcuate isochelae. E, Section through peripheral skeleton. F, Australian distribution. G, NTMZ2863. H, Section through fragment of holotype ZMB4435. skeletal structure, spicule geometry, and spicule sizes differentiate this species from other Microciona in the Indo-west Pacific. Clathria (Mierociona) lizardensis sp. nov. (Figs 102-103, Plate 3E) MATERIAL. HOLOTYPE - QMG304121: Blue Lagoon, Lizard I., Cairns Section, Great Barrier Reef, 14°41.0'S, 145 °27.5'E, 9m depth, 03.iv.1994, coll. J.N.A. Hooper, L.J. Hobbs, J.A. Kennedy & S.D. Cook (SCUBA). HABITAT DISTRIBUTION. Coral reef, patch reef in lagoon, under coral overhangs, on ledges or exposed coral heads, growing on live coral, coral rubble at base of reef or on dead bivalves; 9-12m depth; Lizard I. (FNQ) (Fig. 102G). DESCRIPTION. Shape. Thinly or thickly encrusting, bulbous in life, usually following contours of substrate, up to about 101nm thick, collapsing and less than 4mm thick when preserved. Colour. Pale red alive (Munsell 2.5R 6-5/10), light brown in ethanol. Oscules. Large, up to 4mm diameter alive, surrounded by raised membraneous lip, usually situated on apex of bulbous (flaccid) surface, with drainage canals radiating towards pores; oscules and drainage canals not visible in preserved material. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 213 FIG. 99. Clathria (Microciona) grisea (Hentschel) (NTMZ2863). A, Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal principal acanthostyle. D, Spination on principal acanthostyle. E, Echinating acanthostyle. F, F,chinating acanthostyle spines. G-H, Smaller and larger arcuate isochelae. Texture and surface characteristics. Soft, slimy, easily peeled from substrate; smooth, fleshy flaccid surface in life, with slightly sculptured subectosomal drainage canals visible in live sponge; in preserved material surface uneven, regularly papillose. Ectosome and subectosome. Membraneous, collagenous, rarely intact in histological sections, with some detritus; tips of choanosomal principal MEMOIRS OF THE QUEENSLAND MUSEUM 214^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A FIG. 100. Clathria (Microciona) illawarrae sp.nov. (holotype QMG304572). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, U-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 215 FIG. 101. Clathria (Microciona) illawarrae sp.nov. (holotype QMG304572). A, Hymedesmoid basal skeleton. B, Erect spongin fibre. C, Echinating acanthostyles. D, Acanthostyle spines. E, Reduced palmate isochelae. F, U-shaped toxa. 216zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM CD Clat hr ia ( Micr ociona) lizar densis sp.nov. (holotype QMG304121). A, Choanosomal principal style/ FIG. 102. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Accolada-U-shaped toxas. D, Echinating acanthostyles. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. styles arising from fibre endings, and subectosomal auxiliary subtylostyles protrude through surface in preserved material, but probably do not when alive; no special ectosomal skeleton but plumose bundles of subectosomal subtylostyles clustered on intact parts of surface skeleton, usually lying just below the ectosome. Choanosome. Microcionid skeletal structure, with very thick, relatively long spongin fibre nodes, 450-1900p,m long, up to 420p.m diameter, arising from hymedesmoid basal spongin fibre, 70-23011in diameter, lying directly on substrate; fibre nodes discrete, erect, without any anastomoses between adjacent nodes, 300-770p.m ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 217 FIG. 103. Clathria (Microciona) lizardensis sp.nov. (holotype QMG304121). A, Choanosomal skeleton. B, Ascending fibre node. C, Echinating acanthostyles. D, Acanthostyle spines. E, Palmate isochela. F, Accolada and u-shaped toxas. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 218zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA apart, unbranched except at apex of each node which bifurcates 1 or more times; fibres dark brown, with heavy spongin, cored by multi- or paucispicular plumose tracts of choanosomal principal styles, with 1 or more principal styles protruding from apex of each fibre node; fibres moderately heavily echinated by acanthostyles dispersed evenly over each ascending fibre node and all basal fibres; exterior surface of most fibres often with small amount of collagen and plumose bundles of auxiliary spicules lying paratangential to surface (but this is probably an artifact of preservation, the mesohyl region collapsing around the fibres, whereas in life the areas between fibres is likely to contain a more structured aquiferous system); mesohyl matrix granular, containing some detritus, abundant auxiliary spicules, and rare microscleres; choanocyte chambers difficult to see in preserved material, small, oval, up to 40v.m diameter. Megascleres. Choanosomal principal styles long or short, thick, straight, cylindrical or clubshaped, variable basal terminations from tapering hastate, evenly rounded or faintly subtylote, fusiform points. Length 183-(272.3)-345p,m, width 8-(l2.3)-1611m. Subectosomal auxiliary subtylostyles long, very slender, usually straight, rarely curved, sometimes sinuous, with well developed smooth subtylote bases, fusiform points. Length 211(306.2)-428p.m, width 2-(3.8)-6p.m. Echinating acanthostyles relatively long, thick, prominently subtylote, fusiform pointed, with more-or-less evenly dispersed very small spines, but abundant larger spines concentrated only on base and point, giving appearance of aspinose shaft. Length 81-(94.3)-112Rm, width 446.6)111.1.m. Microscleres. Isochelae palmate, unmodified, uncommon, moderately large, with long broad front ala, reduced lateral alae completely fused to shaft, front and lateral alae approximately the same size. Length 16-(22.4)-28p,m. Toxas uncommon, accolada to u-shaped, long or short, very thin, with slight central curvature, straight arms or faintly reflexed arms. Length 22-(85.5)-112Rm, width 1.0-(1.2)-1.511m. ETYMOLOGY. For the type locality. REMARKS. This species has typical `microcionid' skeletal structure, with long, discrete, virtually unbranched spongin fibre nodes arising from a hymedesmoid basal fibre skeleton. The species is also remarkable for the thickness and density of its spongin fibres, which are even heavier than those found in C. (T) corneolia from New Caledonia (which was named for this character). The external colouration and bulbous surface processes seen in C. (M.) lizardensis are also reminiscent of C. (T) corneolia, although spicule geometry, spicule size and skeletal structure differ substantially between the two species (Hooper & Levi, 1993a), and they do not appear to be otherwise closely related. In its live external appearance this species could also be mistaken for C. (M.) aceratoobtusa, but that species has entirely smooth echinating styles, curved principal styles, oxhorn toxas and abundant isochelae. Comparisons with other Indo-west microcionids are discussed in the remarks for C. (M.) aceratoobtusa. OTHER SPECIES OF CLATHR1A (MICROCIONA) Clathria (Microciona) adioristica (de Laubenfels, 1953) Dictyociona adioristica zyxwvutsrqponmlkjihgfedcbaZYXWVU de Laubenfels, 1953a: 526-528, textfig.5 [ Gulf of Mexico] ; Wells et al.,1960: 217-218, textfigs 21,24 [ North Carolina] . Clathria (Microciona) adioristica; Van Soest, 1984b: 104, 108-109, table 4 [ affinity with Clathria obliqua, possible synonymy] . MATERI AL. HOLOTYPE: USNM23403, paratype MLUMML4-214. NW. Atlantic, Caribbean. Clathria (Microciona) affinis (Carter, 1880) Microciona affinis Carter, 1880a: 41, 151, 153, p1.4,fig.15 [ Gulf of Manaar, Ceylon] ; Carter, 1881a: 368,384; Carter, 1882b: 111; Ridley & Dendy, 1887: 110; Dendy, 1889a: 38; Vosmaer, 1935a: 608; Burton, 1959a: 247 [ S. Arabian coast, Zanzibar] . Not Microciona affinis; de Laubenfels, 1936a: 111. Not Hymeraphia affinis Topsent, 1889: 43, fig.8. MATERI AL. HOLOTYPE: LFM dest royed, fragment BMNH1936.3.4.597. Gulf of Manaar, Arabian Gulf. Clathria (Microciona) africana (Levi, 1956) Microciona africana Levi, 1956b: 402-403, text-fig.8 [ Dakar, Sengal] . MATERI AL. HOLOTYPE: MNHNDCL1276. NW. Africa. Clathria (Microciona) angularis (Sara & Siribelli, 1960) Microciona angularis Sara & Siribelli, 1960: 69-71, fig.18 [ Bay of Naples] ; Siribelli, 1960: 12, fig.5C [ Naples] ; Sara & Siribelli, 1962: 47 [ Gulf of Naples] ; Pulitzer-Finali, 1983: 610. MATERI AL. HOLOTYPE: I MZUN100.4. Mediterranean. Clathria (Microciona) anonyma (Burton, 1959) Microciona anonyma Burton, 1959a: 250-251, fig.30 [ Zanzibar area, I ndian Ocean] REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 219 MATERIAL. HOLOTYPE: BMNHI936. 3.4.575. Central E Africa. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT125. Mediterranean. Possible synonym of Clat hria spinarcus (Carter & Hope) (Maldonado, 1992: 1152). Clathria (Microciona) armata (Bowerbank, 1862)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Microciona) atrasanguinea (Bower- Microcionaarm at a Bowerbank, 1862a: 779, 1858, p1.24, figs 26-28; Bowerbank, 1864:41, p1.4, figs 96-98; Bowerbank, 1866: 124, 129-131, 141 [Britain]; Gray, 1867: 535; Norman, 1869:330; Schmidt, 1870:76; Bowerbank, 1874: 60, p1.23, figs 17-24; Carter, 1874b: 405, p1.21, fig.27; Carter, 1874c: 456-457; Carter, 1876: 310; Carter, 1880a: 40-41, 151 [Gulf of Manaar]; Koehler, 1886a: 62 [English Channel]; Topsent, 1888: 117, 124, 125, 141, 156, 158, p1.6, fig.8; Topsent, 1891a: 528 [Roscoff]; Stephens, 1916:234 [W coast, Ireland]; Stephens, 1921; Vosmaer, 1935a: 6-7, 666; Alander, 1942: 62 [Sweden]; Levi, 1956b: 399-400, text-fig.6 [Dakar, Senegal]; Koltun, 1959: 181-182, textfig. 141 [White Sea]; Levi, 1960a: 73, text-figs 16, 17 [Roscoff,Sidmouth]; Poggiano, 1965: 3, 7, 11-14, textfig.5, tables 1, 2 [Italy]; Pulitzer-Finali, 1983: 573-574, 610, text-fig. 69 [Mediterranean]; Wintermann-Kilian & Kilian, 1984: 134 [Colombia]; Ackers, Moss & Picton, 1992: 143 [Ireland]. Microciona arm at us; Bowerbank, 1882: 7, 18, 53; Sollas, 1884: 614; Vosmaer, 1884a: 121; Vosmaer, I885b: 353; Koehler, 1886a: 11, 55; Carter, 1889a: 287; Carter & Hope, 1889: 99, 101-106; Dendy, 1889c: 17; Hope, 1889: 333, 336, 337; Chatin, 1890: 889; Topsent, 1890c: 202, 204; Topsent, 1892a: 17; Topsent, 1893d: 445; Norman, 1892: 6, 11; Hanitsch, 1894: 176; Topsent, 1894a: 8,23; Topsent 1899: 105; Topsent, 1900: 255; Topsent, 1904a: 189; Loisel, 1898:38; Minchin, 1898:529; Minchin, 1909:215; Stephens, 1912: 27; Ferrer Hernandez, 1914: 41. Clat hria arm at a; Topsent, 1925:649 [discussion]; Van Soest & Stone, 1986: 45 [Norway]. Scopalina arm at a; Wright, 1868: 224. Am philect us arm at us; Vosmaer, 1880: 118-119; Vosmaer, 1889: 353; Svarcevskij, 1906: 342; Babic, 1921: 87-88 [Adriatic]; Babic, 1922: 261-262, text-fig.6. Esperia arm at a Fristedt, 1885: 36-38. Not Microciona arm at us; de Laubenfels, 1936a: Ill. Microciona svarchevskyi de Laubenfels, 1936a: 111; Levi, 1960a: 73. cf. Microciona prolifera; Vosmaer, 1935a: 607. MATERIAL. HOLOTYPE: BMNH1910.1.1.66 (1930.7.3.209). Caribbean, NE Atlantic, Gulf of Manaar, NW Africa, Mediterranean. Many of these records are suspect given their disjunct distribution and this taxon is likely to consist of a species complex. Clathria (Microciona) ascendens (Cabioch, 1968) Microcionaascendens Cabioch, 1968a: 239, text-fig.11 [Ros- bank, 1862) Microciona at rasanguinea Bowerbank, 1862c: 824, 1109, 1110, 1135, p1.30, fig. 1, p1.74, fig.2 [British Seas]; Bowerbank, 1864: 188, 286, p1.33, fig.368, p1.34, fig.369; Bowerbank, 1866: 7, 124, 138-141 [Britain]; Bowerbank, 1874: 63, p1.24, figs 14-19; Topsent, 1888: 141, 157; Topsent, 1890c: 202,204; Topsent, 1891a: 528 [Roscoff]; Topsent, 1892c: 17 [Banyuls]; Dendy, 1922: 60, p1.13, fig. la-e [Egmont Reef]; Burton, 1934b: 37, text-fig.10; Burton, 1938a: 30, p1.4, fig.24 [Madras]; Lilly et al., 1953 [recordLough Inc, Ireland]; Levi, 1960a: 72-73, text-fig.I5 [English Channel, Atlantic]; Sara & Siribelli, 1962: 47 [Gulf of Naples]; Levi, 1965: 18-19, text-fig.21 [Red Sea]; Simpson, I968a: 33, text-fig.! [Plymouth,England]; Juniper & Steele, 1969: 161 [Portsmouth, England]; Van Soest & Weinberg, 1980: 10 [Lough Inc. Ireland]; BouryEsnault, 1971: 326 [Banyuls]; Ackers, Moss & Picton, 1992: 142 [Ireland]. Microciona at rosanguinea; Gray, 1867: 535; Norman, 1869: 330; Schmidt, 1870: 76; Carter, I870a: 332, 339, 340; Carter, 1871a: 272, 274; Carter, 1871b: 8; Carter, 1872a: 106, 111, p1.10, figs 17-20; Carter, 1874c: 457; Carter, 1875: 195; Carter, 1876: 308; Carter, 1880a: 38-41, 151 [Gulf of Manaar]; Carter, 1880b: 59 [Indian Ocean]; Carter, 1881a: 384 [record]; Bowerbank, 1882: 7, 18, 54; Koehler, 1885: 53, 55; Vosmaer, 1885b: 209; Koehler, 1886a: 61, 62 [English Channel]; Carter, 1887b: 355; Carter & Hope, 1889: 102, 104-106; Dendy, 1889c: 18; Hope, 1889: 334; Topsent, 1889: 39; Topsent, 1890c: 202; Topsent, 1891d: 232; Topsent, 1893d: 445; Topsent, 1894a: 8, 10, 23; Norman, 1892: 6; Hanitsch, 1894: 176; Heider, 1895: 280; Loisel, 1898: 38; Topsent, 1900: 255 [note]; Woodland, 1908: 140, 145; Minchin, 1909: 217; Burton & Rao, 1932: 344-345; [coasts of Bengal, Burma, India and Arabian Sea]; de Laubenfels, 1936b: 448-449 [Panama]; Pulitzer-Finali, 1983: 610; Rodriguez SolOrzano et al., 1991: 177 [Galicia, Spain]. Microciona at rasanguineum ; Cuenot, 1903: 4 [Arcachon]. Clat hria ( Microciona) at rasanguinea; Van Soest, 1993: 103 [Mauritius]. Am philect us at rasanguineus; Vosmaer, 1880: 115. Plum ohalichondriaat rasanguinea; Hanitsch, 1890: 207-208, 210 [England]. Scopalina lophyropoda; Schmidt, 1868: 26, 40. Scopalina at rosangut hea; Schmidt, 1866a: 149; Schmidt, 1866b: 15. cf. Microciona prohfera; Vosmaer, 1935a: 604, 607. coff, France]; Rodriguez SolOrzano et al., 1979: 44, 59-60, text-fig.15 [Galicia, Spain]. MATERIAL. HOLOTYPE: RMBS. NE Atlantic. MATERIAL. HOLOTYPE: BMNH1930.7. 3.225, paratypes BMNH1930.7.3.226, 1910.1.1.68. Caribbean, NE Atlantic, Mediterranean, Red Sea, Arabian Gulf, W Indian Ocean, W India, Gulf of Manaar, Bay of Bengal, Andaman Sea. Clathria (Microciona) assimilis Topsent, 1925 Clathria (1Vlicrociona) basifixa Clat hria assim ilis Topsent, 1925: 649; Topsent & Olivier, (Topsent, 1913) 1943: 1 [no diagnosis; Adriatic]. Pseudanchinoe assim ilis; de Laubenfels, 1936a: 109 [note]. Microcionaassim ilis; Levi, 1960a: 76 [Adriatic, Naples, Marseilles]; Siribelli, 1960: 18, text-fig.7C [Naples]; Poggiano, 1965: 3, table 1; Vacelet, 1969: 207, text-fig.46 [Mediterranean]; Pulitzer-Finali, 1977: 61 [Bay of Naples]; Pulitzer-Finali, 1983: 610; Pansini, 1987: 170 [Alboran Sea]. cf. Clat hria com pressa; Topsent, 1925: 649. Ophlit aspongia basifixa Topsent, 1913b: 39 [Norway]; Bur- ton, 1935c: 74 [Japan; probable misidentification]; de Laubenfels, 1954: 162 [note]. Clat hria ( Microciona) basifixa; Van Soest & Stone, 1986:45 [Norway]. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT1957. NE Atlantic. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 220zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Hymantho bitoxa Burton, 1930a: 503, text-fig.2 [Norway]; Alander, 1942: 63 [Sweden]. Microciona levis; Fristedt, 1887: 416. Clathria bitoxa; Van Soest, 1984b: 90 [generic synonymy]; Van Soest & Stone, 1986: 47 [note]. Microciona bitoxa Rodriguez SolOrzano & Rodriguez Babio, 1993: 62 [Iberian Peninsula]. MATERIAL. HOLOTYPE: BMNH1910.1. 1.787.v. NE Atlantic. Microciona affinis; de Laubenfels, 1936a: 111. Clathria (Microciona)affinis; Van Soest, 1984b: 93-95, 108, text-fig.36, table 4 [Curacao, West Indies]. Clathria cf. affinis; Kobluk & Van Soest, 1989: 1216 [Bonaire]. Not Microciona affinis Carter, 1880a: 41, p1.14, fig. 15; Vosmaer, 1933: 608. MATERIAL. HOLOTYPE: MNHNDT1841, paratype MNHNDT3584. Caribbean, NE Atlantic. Clathria (Microciona) affinis (Carter, I880a) has priority. Clathria (Microciona) brepha (de Laubenfels, 1930) Clathria (Microciona) carnosa (Bowerbank, 1862) Clathria (Microciona) bitoxa (Burton, 1930) Aaata brepha de Laubenfels, 1930: 27 [California] Anaata brepha; de Laubenfels, 1936a: 91, text-fig.53. Clathria brepha; Van Soest, 1984b: 7 [generic synonymy]. MATERIAL. HOLOTYPE: USNM2I427. PARATYPES BMNH1929.8.22.36, 57. NE Pacific. Clathria (Microciona) brondstedi sp. nov. Hymedesmia pennata Brondsted, 1932: 12 [Faeroe Is]. Anaata pennata; de Laubenfels, 1936a: 109. Clathria pennata; Van Soest,1984b: 7 [generic synonymy for Anaata]. Not Desmacella pennata Lambe, 1895: 129. MATERIAL. HOLOTYPE: UZM (not located). NE Atlantic. Clathria ( Microciona) pennata (Lambe, 1895) has seniority. Clathria (Microciona) bulboretorta (Carter, 1880) Microcionabulboretorta Carter, I880a: 41, 42, 151, 153, p1.4, fig.3a-e [Gulf of Manaar, Ceylon]; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: LFM destroyed. Gulf of Manaar. Clathria (Microciona) bulbotoxa Van Soest, 1984 Clathria (Microciona)bulbotoxa Van Soest, 1984b: 103-104, p1.7, figs 5-8, text-fig.41, table 4 [Curacao, West Indies]. Microciona bulbotoxa; Pulitzer-Finali, 1986: 149-150 [West Indies]. MATERIAL. HOLOTYPE: ZMAPOR4789. Caribbean. Clathria (Microciona) calla (de Laubenfels, 1934) Axociella calla de Laubenfels, 1934: 16 [Puerto Rico]. Avocielita calla; de Laubenfels, 1954: 149 [note]; Sim & Byeon, 1989: 40, p1.5, figs 3-5 [Korea; probable misidentification]. Clathria calla; Boury-Esnault, 1973: 286, text-fig.46 [Brazilian Basin]; Zea, 1987: 170, text-fig.59, p1.2, fig. 3 [Colombian Caribbean]. Clathria(Microciona)calla; Van Soest,1984b: 100-101, p1.7, fig.1, text-fig.39, table 4 [Curacao, Florida; affinity with Clathria coralloides from Mediterranean]. Microciona calla; Pulitzer-Finali, 1986: 150 [West Indies]. Microciona rarispinosa Hechtel, 1965: 42-44, text-fig.8 [Port Royal, Jamaica]; Wintermann-Kilian & Kilian, 1984: 135 [Colombia]. Tenaciella obliqua; Alcolado, 1976:5; Alcolado, 1980: 10. MATERIAL. HOLOTYPE: USNM. Caribbean, tropical SW Atlantic. Clathria (Microciona) campecheae nom. nov. II ymeraphia affinis Topsent, 1889: 43, fig.8A [Banc de Campeche]; Topsent, 1904a: 162-3 [Azores]. Microciona camosa Bowerbank, I 862a: 804, 1110 [Britain]; Bowerbank, 1866: 133; Vosmaer, 1935a: 607 ? Halichondria incrustans; Schmidt, 1866a: 150 MATERIAL. HOLOTYPE: BMNH1930.7. 3.203, fragment BMNH1910.1.1.666. NE Atlantic, tropical SW. Atlantic. Clathria (Microciona) claudei sp. nov. Microciona acanthotoxa Levi & Levi, 1989: 81, fig.49 [Philippines]. MATERIAL. HOLOTYPE: MNHNDCL341 I. Philippines. Clathria acanthotoxa (Stephens) has seniority. Clathria (Microciona) cleistochela Topsent, 1925 Clathria cleistochela Topsent, 1925: 650-651, fig.9 [Gulf of Naples]. Microciona cleistochela; de Laubenfels, 1936a: 1 1 1 [note]; Levi, 1960a: 72, fig. I 4 [ Naples,Banyuls]; Siribelli, 1960: 12-14, fig.5B [Naples]; de Laubenfels, 1951b: 214 [Black Sea]; Pulitzer-Finali, 1983: 610; Boury-Esnault & Lopes, 1985: 193-194, fig.42 [Azores]. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT329. Mediterranean, NE Atlantic. Clathria (Microciona) coccinea (Bergquist, 1961) Microciona coccinea Bergquist, 1961a: 38, fig.8a,b [N. New Zealand]; Bergquist & Sinclair, 1968: 427, 428, fig. 1 a [morphology and larvae]; Bergquist & Sinclair, 1973: 43; Bergquist et al., 1970: 248, 254; Evans & Bergquist, 1977: 195-196; Bergquist & Fromont, 1988: 102-103, p1.47, fig.f, p1.48, fig.a; Rudman & Avern, 1989: 335; Dawson, 1993: 36 [note]. Not Thalisias coccinea Duchassaing & Michelotti, 1864: 84, p1.18, fig.5 [St.Thomas]; Wiedenmayer, 1977a: 253, table 49. MATERIAL. HOLOTYPE: NMNZ unregistered. New Zealand. Clathria (Microciona) ctenichela (Alander, 1942) M wrociona ctenichela Alander, 1942: 61-62, p1.15, fig.20 [Sweden]. Clathria (Microciotia) ctenichela; Van Soest & Stone, 1986: 44-45 [Norway]. MATERIAL. HOLOTYPE: ZMA. NE Atlantic. Clathria (Microciona) dendyi (Bergquist & Fromont, 1988) Microciona dendyi Bergquist & Fromont, 1988: 100-102, p1.47, figs d,e [Slipper 1.]; Dawson, 1993: 37 [note]. MATERIAL. HOLOTYPE: NMNZPOR114. New Zealand. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 221 Clathria (Microciona) densa (Burton, 1959) Microciona densa Burton, 1959a: 248, text-fig.28 [S Arabian coast] MATERIAL. HOLOTYPE: BMNH1936.3. 4.456. Arabian Gulf. Clathria (Microciona) dianae (Schmidt, 1875) Suberites dianae Schmidt, 1875: 116, p1.1,fig.1 [Norway]; Czerniawsky, 1880: 70. Microciona dianae; Thiele, 1903b: 394, 395, 398, p1.21, fig.28a-e; Vosmaer, 1935a: 608, 630; de Laubenfels, 1936a: 1 1 1 [note]. Not A rtemisina dianae Topsent, 1907: 69; Vosmaer, 1935a: 630. Clathria dianae; Van Soest & Stone,1986: 47 [note]. MATERIAL. HOLOTYPE: unknown. NE Atlantic. Clathria (Microciona) ditoxa (Stephens, 1916) Eurypon ditoxa Stephens, 1916:239-240 [W coast of Ireland]; Stephens, 1921: 51, p1.5; Burton, 1959b: 44-45 [Iceland]. Microciona ditoxa; Levi, 1960a: 66 [W coast of Ireland, Atlantic]. Dictyociona ditoxa; de Laubenfels, 1936a: 110 [note]. MATERIAL. HOLOTYPE: INMSR151.9.1916. NE Atlantic. Clathria (Microciona) duplex Sara, 1958 Clathria duplex Sara, 1958: 262-264, fig.24 [Gulf of Naples]. Microciona duplex; Levi, 1960a: 69 [Naples]; Siribelli, 1960: 14-15, fig.6A [Naples]; Sara, 1963: 210 [Gulf of Policastro]; Pulitzer-Finali, 1983: 610; Rodriguez SolOrzano et al., 1991: 179, fig.3 [Galicia, Spain]. MATERIAL. • HOLOTYPE: IMZUN93.26.x.58. Mediterranean. Clathria (Microciona) echinata (Alcolado, 1984) Axociella echinata Alcolado, 1984: 7 [Cuba]. Clathria echinata; Kobluk & Van Soest, 1989: 1216; Meesters et al., 1991: 194 [Curacao, Bonaire]. Clathria (Microciona) simpsoni Van Soest, 1984b: 97-99, p1.7, figs 2-4, text-fig.38, table 4 [Puerto Rico, Curacao]; Pulitzer-Finali, 1986: 150 [West Indies]. Clathria simpsoni; Zea, 1987: 168, text-fig.58, p1.3, fig.1 [Colombian Caribbean]. MATERIAL. HOLOTYPE: Cuba. Holotype of simpsoni: ZMAPOR3332. Caribbean. Clathria (Microciona) elliptichela (Alander, 1942) Microciona elliptichela Alander, 1942: 58-61 [Sweden]. Clathria elliptichela; Van Soest & Stone, 1986:45 [note]. MATERIAL. HOLOTYPE: ZMA. NE Atlantic. Clathria (Microciona) fallax (Bowerbank, 1866) Microciona falla.v Bowerbank, 1866: 124, 128, 129, 135 [Hastings, Britain]; Vosmaer, 1935a: 607; Curtis, 1970: 260-261 [cytology]; Ackers, Moss & Picton, 1992: 147 [Ireland]. Sophax fallav; Gray, 1867: 521. MATERIAL. LECTOTYPE-BMNH 1910.1. 1.71. PARALECTOTYPE BMNH1930.7.3.198, fragment USNM5047. NE Atlantic. Clathria (Microciona) fascispiculifera (Carter, 1880) Microciona fascispiculifera Carter, 1880a: 44, 45, 151, 153, p1.4, fig.7a-g [Gulf of Manaar]; Hallmann, 1916c: 637 [note]; [?] Vosmaer, 1935a: 608. Damoseni fascispiculifera; de Laubenfels, 1936a: 110. MATERIAL. HOLOTYPE: LFM destroyed, no extant fragment in BMNH. Gulf of Manaar. Clathria (Nlicrociona) fraudata (Bowerbank, 1874) Microciona fraudata Bowerbank, 1874: 273, 275, 277, p1.83, figs 7-11 [Polperro, Fowey Hbr.]; Vosmaer, 1935a: 607. MATERIAL. HOLOTYPE: BMNH1930.7.3. 205. NE Atlantic. Clathria (Microciona) frogeti (Vacelet, 1969) Microciona frogeti Vacelet, 1969: 208, text-fig.47 [Mediterranean]. Microciona fregeti [sic.]; Pulitzer-Finali, 1983: 610 [list]. MATERIAL. HOLOTYPE: MNHN missing. W Mediterranean. Clathria (Microciona) gradalis Topsent, 1925 Clathria gradalis Topsent, 1925: 651-653 [Gulf of Naples]; Topsent & Olivier, 1943: 1 [Monaco]; Sara, 1958: 258260, text-fig.22 [Gulf of Naples]; Sara, 1960a: 461 [Ischia]. Clathria gradalis var. atoxa; Topsent, 1928a: 299, p1.10, fig.14 [Boavista I., Senegal]. Microciona gradalis; de Laubenfels, 1936a: 111; Levi, 1960a: 75 [W Mediterranean]; Sara & Siribelli, 1960: 67 [Bay of Naples]; Siribelli, 1960: 16, text-fig.6B [Naples]; Sara & Siribelli, 1962:47-48 [Gulf of Naples]; Poggiano, 1965:3, table 1; Cabioch, 1968a: 244 [Roscoff, N. France]; Vacelet, 1969: 207 [W Mediterranean]; Pulitzer-Finali, 1977: 63 [Bay of Naples]; Pulitzer-Finali, 1983: 610. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT328. Mediterranean, NE Atlantic, NW Africa. Clathria (Microciona) haematodes (de Laubenfels, 1957) Microciona haematodes de Laubenfels, 1957: 240, text-fig.6 [Oahu, Hawaii]; Bergquist, 1977: 67 [Hawaii]. MATERIAL. HOLOTYPE: USNM23533. Hawaii. Clathria (Microciona) haplotoxa (Topsent, 1928) Leptoclathria haplotoxa Topsent, 1928a: 298, p1.10, fig.16 [Madeira I.]; [cf.] Topsent, 1934b: 24. Microciona haplotoxa;Topsent, 1934a: 92-93 [Gulf of Gabes, Tunisia]; Levi, 19566: 400-402, text-fig.7 [Dakar, Senegal]; Levi, 1960a: 70 [Madeira, Tunisia]; PulitzerFinali, 1983: 610. MATERIAL. HOLOTYPE: MNHNDT1101. Mediterranean, NE Atlantic, NW Africa. Clathria (Microciona) hentscheli sp. nov. Ilymeraphia lendenfeldi Hentschel, 1912: 378-379, p1.20, fig. 35 [Mimien Bay, Am I., Arafura Sea]. Eurypon lendenfeldi; de Laubenfels, 1936a: 110. Not Clathria lendenfeldi Ridley & Dendy, 1886: 474. MATERIAL. LECTOTYPE: SMF 1705. Indonesia. Clathria (Thalysias) lendenfeldi Ridley & Dendy has priority. 222zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Clathria (Microciona) heterotoxa (Hentschel, 1929) Microciona heterotoxa Hentschel, 1929: 891-892, 970, p1.14, fig.5 [White Sea]; Koltun, 1959: 182-183, text-fig.142 [Arctic, USSR]. Dictyociona heterotoxa; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953a: 527. MATERIAL. HOLOTYPE: HM or ZMB (not located). Arctic. Clathria (Microciona) hymedesmioides Van Soest, 1984 Clathria (Microciona) hymedesmioides Van Soest, 1984b: 104-105, p1.7, figs 9-10, text-fig.42, table 4 [Curacao]. MATERIAL. HOLOTYPE: ZMAPOR4790. Caribbean. Clathria (Microciona) ixauda (Levi, 1969) Microciona ixauda Levi, 1969: 965, text-fig.7a [Vema Seamount]. MATERIAL. HOLOTYPE: MNHNDCL1415. S Atlantic. Clathria (Microciona) jecusculum (Bowerbank, 1866) Hymeniacidon jecusculum Bowerbank, 1866: 198 [Harris I., Hebrides]. Microciona jecusculum; Bowerbank, 1874: 273-275, p1.83, figs 1-6; Carter, 1876: 237 [Cape St. Vincent; Faroe Is]; Vosmaer, 1933: 607 [imperfectly known]. MATERIAL. HOLOTYPE: unknown; fragments BMNH1954.3.9.176, 177. NE Atlantic. Clathria (Microciona) kentii (Bowerbank, 1874) Microciona kentii Bowerbank, 1874: 311, 312, 317-319, p1.89, figs 9-13 [Jersey, Strangford Lough]; Vosmaer, 1935a: 607. MATERIAL. HOLOTYPE: BMNH1910.1.1.77, fragment USNM5044. NE Atlantic. Clathria (Microciona) laevis (Bowerbank, 1866) Microciona laevis Bowerbank, 1866: 124, 127-128 [Britain]; Stephens, 1917: 12, p1.1, fig.3 [N. of Bolus Head, Ireland]; Vosmaer, 1935a: 607; Burton, 1959b: 43 [Iceland]. Not Microciona laevis; Fristedt, 1887: 415. Abila laevis; Gray, 1867: 539. Hymantho laevis; de Laubenfels, 1936a: 111; Alander, 1942: 63 [Sweden]. MATERIAL. HOLOTYPE: BMNH1930.7.3. 215. NE Atlantic. Clathria (Microciona) laevissima (Dendy, 1922) MATERIAL. HOLOTYPE: USNM22827. W central Pacific. Clathria (Microciona) leighensis sp. nov. Microciona rubens Bergquist, 1961a: 38, text-fig.9 [NNew Zealand]; Bergquist & Green, 1977b: 289-302 [ontogeny]; Bergquist & Fromont, 1988: 103, p1.48, figs b-c [N New Zealand]; Dawson, 1993: 37 [note]. Not Thalassodendron rubens Lendenfeld, 1888: 223. MATERIAL. HOLOTYPE: NMNZ unregistered. New Zealand. C. (Clathria) rubens (Lendenfeld, 1888) has priority. Clathria (Microciona) levii (Sara & Siribelli, 1960) Microciona levii Sara & Siribelli, 1960: 71-73, text-fig.19 [Bay of Naples]; Siribelli, 1960: 6-8, text-fig.2 [Naples]; Poggiano, 1965:3, table 1; Pulitzer-Finali, 1983:610 [list]. MATERIAL. HOLOTYPE: IMZUN31. Mediterranean. Clathria (Microciona) longispiculum (Carter, 1876) Microciona longispiculum Carter, 1876: 231, 237, 238, 470, p1.12, fig. 1 h, p1.15, fig.31a-c [N. Scotland]; Vosmaer, 1935a: 608 [insufficiently known]. MATERIAL. HOLOTYPE: BMNH1887.10. 29.3. NE Atlantic. Clathria (Microciona) longistyla (Burton, 1959) Microciona longistyla Burton, 1959a: 249-250, text-fig.29 [S. Arabian coast]; Sim & Kim, 1988: 26 [Korea]; Sim & Byeon, 1989: 40 [Korea; possible misidentifications]. MATERIAL. HOLOTYPE: BMNH1936.3.4. 583. Arabian Gulf, ? S China Sea. Clathria (Microciona) macrochela (Levi, 1960) Microciona macrochela Levi, 1960a: 70, text-fig.13 [Roscoff]. MATERIAL. HOLOTYPE: MNHNDCL940. NE Atlantic. Clathria (Microciona) microjoanna (de Laubenfels, 1930) Microciona microjoanna de Laubenfels, 1930: 27 [Carmel, California]; de Laubenfels, 1932: 93-95, text-fig.55 [California]; Bakus, 1966:433-435, text-fig.4, table 4 [San Juan Arch., Washington State]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Bakus & Green, 1987: 72 [S. California]. MATERIAL. HOLOTYPE: USNM21468. PARATYPE USNM21469, fragment BMNH1929.8.22.28. NE Pacific. Clathria (Microciona) micronesia (de Laubenfels, 1954) Hymedesmia laevissima Dendy, 1922: 81-82, p1.15, fig.1 [Mauritius]. Folitispa laevissima; de Laubenfels, 1936a: 119 [note]. MATERIAL. HOLOTYPE: BMNH1921.11. 7.69. W Indian Ocean. Microciona micronesia de Laubenfels, 1954: 145-146, textfig.93 [Majuro Atoll]. MATERIAL. HOLOTYPE: USNM22833. W central Pacific. Clathria (Microciona) lajorei (de Laubenfels, 1954) Clathria (Microciona) microxea (Vacelet & Vasseur, 1971) Anaata lajorei de Laubenfels, 1954: 147-148, text-fig. 95 [Ailing-lap-lap Atoll]. Clathria lajorei; Van Soest, 1984a: 129 [generic synonymy]. Paratenaciella microxea V acelet & Vasseur, 1971: 103, textfig.61 [Tulear, Madagascar]. MATERIAL. HOLOTYPE: MNHNDIV27. W Indian Ocean. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 223 Clathria (Microciona) mima (de Laubenfels, 1954) Ophlitaspongia mima de Laubenfels, 1954: 161-162, textfig.105 [Majuro Atoll, central W. Pacific]; WintermannKilian & Kilian, 1984: 135 [Colombia]. MATERIAL. HOLOTYPE: USNM22839. NW central Pacific, Caribbean. Clathria (Microciona) namibiensis (Uriz, 1984) Microciona namibiensis Uriz, 1984b: 111-113, text-figs 3 A-B, 5D; Uriz, 1988a: 87 [Namibia]. MATERIAL. HOLOTYPE: ABIPB-12. SW Africa. Clathria (Microciona) normani (Burton, 1930) Hymantho norrnani Burton, 1930a: 503, text-fig.! [Norway]. Clathria normani; Van Soest, 1984b: 90 [generic synonymy]; Van Soest & Stone, 1986: 46-47 [note]. MATERIAL. HOLOTYPE: BMNH1910.1.1. 791.iii. NE Atlantic. Clathria (Microciona) novaezealandiae (Brondsted, 1924) Microciona novaezealandiae Brondsted, 1924: 463-464, textfig.19 [Slipper Is]; Bergquist & Fromont, 1988: 103-104; Dawson, 1993: 37. Wetmoreus novaezealandica; de Laubenfels, 1936a: 112. MATERIAL. HOLOTYPE: UZM not located, no fragment in BMNH. New Zealand Clathria (Microciona) osismica (Cabioch, 1968) Microciona osismica Cabioch, 1968a: 240-244, text-fig. 12 [Roscoff, France]. MATERIAL. HOLOTYPE: RMBS. NE Atlantic. Clathria (Microciona) parthena (de Laubenfels, 1930) Microciona parthena de Laubenfels, 1930:27 [California]; de Laubenfels, 1932: 95-96, text-fig.56 [California]; Henkart et al., 1973: 3045-3050, text-figs 1-5 [biochem.]; Cauldwell et al., 1973: 3051-3058, text-figs 1-5 [biochem.]; Lee & Gilchrist, 1985: 24-32 [biocheml; Sim Bakus, 1986: 9 [California]; Bakus & Green, 1987: 72-73 [S. California]. MATERIAL. HOLOTYPE: USNM21383. PARATYPE BMNH1929.9.30.6. NE Pacific. Clathria (Microciona) pennata (Lambe, 1895) Desmacella pennata Lambe, 1895: 129, p1.4, figs 6a-d [Sooke, Vancouver I., Canada]; Ricketts & Calvin, 1948: 34 [Vancouver I., Canada]. Ophlitaspongia pennata; de Laubenfels, 1927: 265-266, textfigs 1-4 [Laguna Beach and Monterey Pen., California]; de Laubenfels, 1930: 28; Burton, 1930a: 521; de Laubenfels, 1932: 103, text-fig.62 [var. californiana]; Hewatt, 1946: 193 [California]; Hartman (in Light, 1954): 20 [California]; de Laubenfels, 1954: 162 [note]; Burton, 1959a: 247 [key to species]; de Laubenfels, 1961: 198, fig.1 [California and Washington State]; Bakus, 1966: 435-440, textfig.5, tables 5-6 [et var., San Juan Arch., Washington]; Anderson, 1973:5668 [associates]; Lee & Gilchrist, 1985: 24-32 [biochem.]; Sim & Bakus, 1986: 10; Bakus & Green, 1987:73 [var. californiana]; Sim & Byeon, 1989: 37, p1.1, figs 1-4 [var. californiana; Korea]; Lee & Klontz, 1991: 61 [chemistry]. Tylodesma pennata; Koltun, 1959: 96 text-fig.51, 1-3 [S. Kuriles]. Biemna pennata; Koltun, 1958: 54. Ophlaaspongia affinis basifixa; Burton, 1935c: 74 [Possiet Bay, Sea of Japan]. Microciona pennata; Simpson, 1968a: 40, p1.15 [Washington State]. Clathria (Ophlitaspongia)pennata; Rudman & Avem, 1989: 335 [associates; probable misidentification of sponge]. Not Pandaros pennata Duchassaing & Michelotti,1864: 88. Not Hymedesmia pennata Brondsted, 1932: 12 [see C. brondstedi sp. nov.]. MATERIAL. HOLOTYPE: USNM7488. PARATYPE NMCIC1900-2826. NE Pacific, NW. Pacific, Japan. Clathria (Microciona) plana (Carter, 1876) Microciona planum Carter, 1876: 238, 472 [Cape St. Vincent, Hebrides]. Microciona plana; Topsent, 1889: 41-42 [Banc de Campêche]; Norman, 1892: 9. Amphilectus planus; Vosmaer, 1880: 121; Dendy, 1889: 18. Clathria planer; Topsent, 1894b: 30; Van Soest, 1984b: 108 [unrecognisable]. cf. Microciona prolifera; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: BMNH1890.4. 10.13. NE Atlantic. Clathria (Microciona) plinthina (de Laubenfels, 1954) Microciona plinthina de Laubenfels, 1954: 144-145, textfig.92 [Ailing-lap-lap Atoll]. MATERIAL. HOLOTYPE: USNM22949. W central Pacific. Clathria (Microciona) poecilosclera (Sara & Siribelli, 1960) Microciona poecilosclera Sara & Siribelli, 1960: 73-75, textfig.20 [Bay of Naples]; Siribelli, 1960: 18, text-fig.7B [Naples]; Poggiano, 1965:3, table 1; Pulitzer-Finali, 1983: 610. MATERIAL. HOLOTYPE: IMZUN350. Mediterranean. Clathria (Microciona) polita (Ridley, 1881) Hymedesmia polita Ridley, 1881: 121-122, p1.10, fig.9 [Sandy Point, Magellan Straits; supposed affinities with Clathria tuberosa (Bowerbank)]. Microciona polita; Topsent, 1900: 113 [English Channel; identified with doubt]. MATERIAL. HOLOTYPE: BMNH1879.12.27. 22. SW Atlantic, NE. Atlantic. Clathria (Microciona) primitiva (Koltun, 1955) Microciona prirnitiva Koltun, 1955a: 16-17, text-fig.6 [Behring Sea]; Burton, 1959: 43 [Iceland]; Koltun, 1959: 184, text-fig. 144 [USSR]. Not Clathriella primitiva Burton, 1935c: 73. MATERIAL. HOLOTYPE: ZIL, fragments BMNH1932.1.1.241, 572. Boreal region. Clathria (Microciona) proxima (Lundbeck, 1910) Hymedesmia proxima Lundbeck, 1910: 81 [Denmark Strait]. Anaata proxima; de Laubenfels, 1936a: 109 [imperfectly known]. 224zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Clathriaproxima; Van Soest, 1984b: 7 [generic synonymy for Anaata]. MATERIAL. HOLOTYPE: ZRS. NE Atlantic. Clathria (Microciona) pugio (Lundbeck, 1910) Hymedesmia pugio Lundbeck, 1910: 94 [Denmark Strait]. Anaata pugio; de Laubenfels, 1936a: 109. Clathria pugio; Van Soest, 1984b: 7 [generic synonymy for Anaata]. MATERIAL. HOLOTYPE: ZRS. NE Atlantic. Clathria (Microciona) pustulosa (Carter, 1882) Halichondria pustulosa Carter, 1882a: 285, p1.11, fig.1 [vicinity of Pantagonia and Falkland Is]. Not Halichondria pustulosa; Carter, 1886g: 450 [Port Phillip Bay, Victoria]. Anaata pustulosa; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: BMNH not found, fragment BMNH1954.3.9.233. SW Atlantic. Clathria (Microciona) quadriradiata (Carter, 1880) Microciona quadriradiata Carter, 1880a: 42, 43, 151, 153, p1.4, figs 4a-d [Gulf of Manaar]; [?] Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: LFM destroyed. Gulf of Manaar. Clathria (Microciona) rhopalophora (Hentschel, 1912) Hymeraphia rhopalophora Hentschel, 1912: 380, p1.20, fig.37 [Aru I., Arafura Sea]. Microciona rhopalophora; Burton, 1959a: 248 [Maldives]; Thomas, 1970b: 206, text-fig.7 [Cocos-Keeling Basin, Gulf of Manaar]. Eurypon rhopalophora; de Laubenfels, 1936a: 111. MATERIAL. HOLOTYPE: HM (fragment 5MF992). Indonesia. Clathria (Microciona) scotti Dendy, 1924 Clathria scotti Dendy, 1924a: 352, p1.10, fig.!, p1.14, figs 5-8 [E of North Cape, New Zealand]. Pseudanchinoe scold; de Laubenfels, 1936a: 109 [note]; Bergquist & Fromont, 1988: 110-111, p1.51, figs b-d [N. New Zealand]; Dawson, 1993: 39 [note]. MATERIAL. HOLOTYPE: BMNH1923.10. 1.128, paratypes BMNH1923.10.1.129-131, AMZ2568. New Zealand. Clathria (Microciona) seriata (Grant, 1826) Spongia seriata Grant, 1826: 116. Halichondria seriata; Johnston, 1842:74, 125, 197, 248, 258, p1.14, fig.2; Gray, 1848: 12,16. Halic lona seriata; Bowerbank, 1861: 235; Bowerbank, 1862a: 769, 824, p1.29, fig.10. Chalitza seriata; Bowerbank, 1864: 24,2275, p1.17, fig.287; Bowerbank, 1866: 139, 294, 361, 376-378; Wright, 1868: 228; Norman, 1869:298-299; Schmidt, 1870: 3, 77; Carter, 1871b: 196. Clathriaseriata; Schmidt, 1866b: 10, 24, p1.1, fig.7; Vosmaer, 1935a: 619 [uncertain affinity]; Hanitsch, 1889: 158; Hanitsch, 1890: 205-207 [England]; Babic, 1921: 84 [Adriatic]; Babic, 1922: 244-245, text-fig.T [Adriatic]. Seriatula seriata; Gray, 1867: 515. Desmacidon seriata; Schmidt, 1868: 12. ? Desmacodes seriatus; Vosmaer, 1880: 107. Ophlitaspongia seriata; Bowerbank, 1874:6, 167, p1.65, figs 1-4; Carter, 1875: 196; Bowerbank, 1882: 14, 24, 120, 186-188; Carter, 1883b: 314; Vosmaer, 1885: 357; MacMunn, 1888: 12, 14, 20; Dendy, 1889c: 14; Topsent, 1890c: 204; Topsent, 1891a: 529; Norman, 1892: 8; Minchin, 1900: 20, fig.32; Thiele, 1905: 450-451; Kirkpatrick, 1907: 274; Kirkpatrick, 1908a: 26; Weltner, 1910a: 23; Hallmann, 1912: 254; Stephens, 1912: 3, 28; Stephens, 1916: 234 [Ireland]; Ferrer Hernandez, 1914: 43; Lilly et al., 1953 [Ireland]; de Laubenfels, 1954: 161162 [note]; Burton, 1959a: 247 [key to species of Ophlitaspongia]; Levi, 1960a: 64-65, text-fig.9 [Atlantic]; Levi, 1963: 59-60, text-fig. 69, p1.9B-C [Cape Town, South Africa]; Bergquist & Sinclair, 1968: 427, 428, textfig.1B [larvae, New Zealand]; Bergquist & Hogg, 1969: 207, 210; Fry, 1970: 135-157 [ecology]; Fry, 1971: 155178 [larvae]; Bergquist &Sinclair, 1973:37-39; Fry, 1973: 159-170 [ecology]; Van Soest & Weinberg, 1980: 10 [Lough Ine, Ireland]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Rudman & Avern, 1989: 335 [associates]; Ackers, Moss & Picton, 1992: 147 [Ireland]; Dawson, 1993: 38 [note]. Echinoclathria seriata;Topsent, I893d: 445; Hanitsch, 1894: 179; Hanitsch, 1895: 212; Heider, 1895: 281; Topsent, 1896: 114; Loisel, 1898: 38; Minchin, 1900: 20; Whitelegge, 1907: 503. Microciona seriata; Simpson, 1968a: 37, pls 9-10, text-fig.2 [Plymouth, England]. Not Rhaphidophlus seriatus Thiele, 1899: 14, p1.1, fig.6, p1.5, fig.7 [Celebes]. Ophlitaspongia papilla Bowerbank, 1866: 14, 378-380 [Vazon Bay, Guernsey]; Bowerbank, 1874, p1.70, figs 1-4; Bowerbank, 1882: 187 [Westport Bay, Guernsey]; Koehler, 1886a: 62 [English Channel]; Hallmann, 1912: 254 [note]. ? Clathria papilla; Schmidt, 1870: 77; Vosmaer, 1880: 155. Echinoclathria papilla; Hanitsch, 1894: 8-10, 16, 25, 26. Ophistospongia papilla; Gray, 1867: 515. MATERIAL. HOLOTYPE: BMNH1847.9.7.14, fragments BMNH1910.1.1.2368, 2369. NE Atlantic, S Africa, New Zealand, Mediterranean. This species is undoubtedly composite, consisting of at least two sibling species (Atlantic and Indo-Pacific populations). Clathria (Microciona) sigmoidea (Cuartas, 1992) Microciona sigmoidea Cuartas, 1992: 85-88, figs 53-57, 67 [Mar del Plata, Argentina]. MATERIAL. HOLOTYPE: MCNPC04-81-43. SW Atlantic. C1athria (Microciona) simae sp. nov. Axociella cylindrica; Sim & Byeon, 1989: 39-40, p1.5, figs 1-2 [S. Korea]. Not Esperiopsis cylindrica Ridley & Dendy, 1886: 340; Ridley & Dendy, 1887: 79-80, p1.19, figs 2a-b. Not Axociella cylindrica; Hamann, 1920: 780-783, p1.37, figs 2-4, text-fig.2. Not Rhaphidophlus cylindricus Kieschnick, 1900: 53, p1.44, fig. 10. MATERIAL. HOLOTYPE: Department of Biology, Han Nam University, Korea 18/vii/1987. S China Sea. C. (Axociella) cylindrica (Ridley & Dendy, 1886) has priority. ETYMOLOGY. Named for Dr C.J. Sim. Clathria (Microciona) similis (Thiele, 1903) Hymeraphia finials Thiele, 1903a: 957, fig.22 [Ternate, Moluccas]; Hentschel, 1912: 377 [Am I., Arafura Sea]. Eurypon sinidis; de Laubenfels, 1936a: 111. Not Microciona similis Stephens, 1915:441. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 225 Not Eurypon similis; Uriz, 1988a: 53-54, text-fig.29. MATERIAL. HOLOTYPE: ZMB7215. Indonesia. Clathria (Microciona) spinarcus (Carter & Hope, 1889) Microcionaspinarcus Carter & Hope, 1889:99-106, p1.6, figs 1-6 [Hastings, England]; Carter, 18896: 250; Hope, 1889: 339; Topsent, 1890c: 199, 202, 205; Topsent, 1892a: 113 [Banyuls]; Topsent, 1892c: 17; Topsent, 1894a: 8, 11; Topsent, 1896: 115; Ferrer Hernandez, 1914: 14; Topsent, 1928a: 62; Levi, 1960a: 74, 76, text-fig.18 [Atlantic]; Boury-Esnault, 1971: 324-325 [Banyuls]; Van Soest & Weinberg, 1980: 6-8, 10, text-fig.8 [Lough Inc, Ireland]; Boury-Esnault & Lopes, 1985: 193, fig.41 [Azores]; Uriz, 1988a: 88-89 [Namibia]; Uriz, 19886: 68 [Namibia]; Maldonado, 1992: 1152 [Aldoboran Sea]; Ackers, Moss & Picton, 1992: 146 [Ireland]. Microciona armata, in part; Carter, 1874c: 457; Carter & Hope, 1889: 101-106. Ligrota spinarcus; de Laubenfels, 1936a: 126 [transferred with hesitation]. Microciona acanthotoxa; Lilly et al., 1953 (Van Soest & Weinberg, 1980: 10). cf. Microciona prolifera; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: BMNH1910.1.1. 501, fragment BMNH1954.3.9.175. NE Atlantic, W Mediterranean, SW Africa. Not Euryponsimilis; Uriz, 1988:53 [see C. urizae, nom. nov.]. MATERIAL. HOLOTYPE: RSME1921.143. 1447, fragments INM.31.1914, BMNH1939. 3.20.11. South Africa. Clathria (Microciona)similis (Thiele, 1903a) has seniority. Clathria (Microciona) strepsitoxa (Hope, 1889) Microciona strepsitoxa Hope, 1889: 334-338, 342, p1.16, figs 1-10; Topsent, 1890c: 199, 205; Topsent, 1891a: 529 [Roscoff]; Topsent, 1892c: 17 [Banyuls]; Topsent, 1894a: 8; Topsent, I934a: 90-92 [Gulf of Gabes]; Alander, 1942: 62 [Sweden]; Levi, 1960a: 67, text-fig.10 [English Channel, Atlantic, Mediterranean]; Siribelli, 1960: 4, textfig. IA [Naples]; Sara & Siribelli, 1960: 67-69, text-fig.17 [Bay of Naples]; Poggiano, 1965: 3, table 1; Boury-Esnault, 1971: 326 [Banyuls]; Pulitzer-Finali, 1977:62 [Bay of Naples]; Rodriguez SolOrzano et al., 1979: 44, 58-59, text-fig. 14 [Galacia, Spain]; Van Soest & Weinberg, 1980: 6, text-fig. 7 [Lough Ine, Ireland]; Pulitzer-Finali, 1983: 573, 610 [Mediterranean]; Wintermann-Kilian & Kilian, 1984: 134 [Colombia]; Boury-Esnault & Lopes, 1985: 192-193, fig.40 [Azores]; Ackers, Moss & Picton, 1992: 145 [Ireland]. cf. Microciona prolifera; Vosmaer, 1935a: 608, 640. MATERIAL. HOLOTYPE: BMNH1910.1.1. 500. NE Atlantic, Mediterranean, Caribbean. Clathria (Microciona) tenuis (Stephens, 1915) Clathria (Microciona) spinatoxa (Hoshino, 1981) Microciona spinatoxa Hoshino, 1981: 155, text-fig.67, p1.7, fig.! [Sasajima]. MATERIAL. HOLOTYPE: MMBSSIS-033. Japan. Clathria (Microciona) spinosa (Wilson, 1902) Microciona spinosa Wilson, 1902: 396-397 [St.Thomas, Puerto Rico]; Simpson, I968a: 37, pls 9-10, text-fig.2 [Bahamas]; Simpson, 19686: 26, pls 6-8, tables 5-7; Wiedenmayer, 1977: 141-142, text-fig. 145 [Bimini]. Axociella spinosa; de Laubenfels, 1936a: 113 [Florida]; de Laubenfels, 1949a: 16 [Bimini]; Storr, 1964: 42 [W coast Florida]; Hechtel, 1965: 43. Clathria (Microciona) spinosa; Van Soest, 19846: 95-96, table 4, text-fig.37 [Curacao]. Clathria spinosa; Zea, 1987: 167, text-fig. 57 [Colombian Caribbean]. cf. Microciona prolifera; Vosmaer, 1935a: 608, 645. MATERIAL. HOLOTYPE: USNM7680. Caribbean. Clathria (Microciona) spongigartina (de Laubenfels, 1930) Aaata spongigartina de Laubenfels, 1930: 27 [Carmel, California]. Anaata spongigartina; de Laubenfels, 1932: 89-91, text-fig. 52; Sim & Bakus, 1986: 12 [California]. Clathria spongigartina; Van Soest, 19846: 7 [generic synonymy]. MATERIAL. HOLOTYPE: USNM21428, fragment BMNH1929.8.22.13. NE Pacific. Microciona tenuis Stephens, 1915:443, p1.40, fig.5 [Saldanha Bay]; Levi, 1963: 67. Hymantho tenuis; de Laubenfels, 1936a: Ill. MATERIAL. HOLOTYPE: RSME missing. South Africa. Clathria (Microciona) tenuissima (Stephens, 1916) Eurypon tenuissima Stephens, 1916: 240 [W coast Ireland; 780m depth]; Stephens, 1921. Dictyociona tenuissima; de Laubenfels, 1936a: 110 [note]; de Laubenfels, 1953a: 528. Microciona tenuissima; Levi, 1960a: 69 [W coast Ireland]; Biblioni, 1993: 312-3, fig.8 [W. Mediterranean]. Not Leptolabis tenuissima Hentschel, 1911: 360-361, textfig.41 [Shark Bay, Western Australia]. MATERIAL. HOLOTYPE: INMSR151.11. 1916. NE Atlantic. Clathria (Microciona) tetrastyla (Hentschel, 1912) Hymeraphia tetrastyla Hentschel, 1912: 379-380, p1.20, fig.36 [Am I., Arafura Sea]. Eurypon tetrastyla; de Laubenfels, 1936a: 111. MATERIAL. HOLOTYPE: SMF 954. Indonesia. Clathria (Microciona) thielei (Hentschel, 1912) Hymeraphia thieleiHentschel, 1912:377-378 [Aru I., Arafura Seat Not Ophlitaspongia thielei Burton, 1932a: 322, p1.55, fig.8, text-fig.32; Koltun, 1964a: 70. MATERIAL. HOLOTYPE: SMF 1708. Indonesia. Clathria (Microciona) stephensae sp. nov. Clathria (Microciona) toximajor Topsent, 1925 Microciona similis Stephens, 1915: 441, p1.40, fig.6 [Saldanha Bay]; Levi, 1963: 58-59, text-fig.67, p1.9A [St. Helena Bay]; Hechtel, 1965: 43-44 [note]. Axocielita sitnilis; de Laubenfels, 1936a: 118. Not Hyrneraphia similis Thiele, 1903a: 957. Clathria toximajorTopsent, 1925: 653-655, text-fig.11 [Gulf of Naples]; Sara, 1960a: 461 [Ischia]. Microciona toximajor, Levi, 1960a: 67 [Naples]; Siribelli, 1960: 6, text-fig.1B [Naples]; Sara & Siribelli, 1962: 48 [Gulf of Naples]; Poggiano, 1965: 3, table 1; Riitzler, 1966 226zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM [Banyuls]; Boury-Esnault, 1971: 325 [Banyuls]; PulitzerFinali, 1983: 610 [list]. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT326. Mediterranean Clathria (Microciona) toxirecta (Sara & Siribelli, 1960) Microciona toxirecta Sara & Siribelli, 1960: 75-77, textfig.21 [Bay of Naples]; Siribelli, 1960: 14, text-fig.5A [Naples]; Sara & Siribelli, 1962: 48 [Gulf of Naples]; Pulitzer-Finali, 1983: 610 [list]. MATERIAL. HOLOTYPE: IMZUN382. Mediterranean. Clathria (Microciona) toxitenuis Topsent, 1925 Clathria toxitenuis Topsent, 1925:655 [Gulf of Naples]; Sara, 1958: 261-262, text-fig.23 [Gulf of Naples]; Sara, 1960a: 462 [Ischia]. Pseudanchinoe toxitenuis; de Laubenfels, 1936a: 109 [note]. Microciona toxitenuis; Levi, 1960a: 69, text-fig.11 [Marseilles, Naples]; Sara & Siribelli, 1960:69-71 [Bay of Naples]; Siribelli, 1960: 10-12, text-fig.4 [Naples]; Sara & Siribelli, 1962: 48 [Gulf of Naples]; Labate, 1964: 334-335, p1.2, fig.8 [Adriatic Sea]; Sara, 1964: 230 [Ligurian Sea]; Poggiano, 1965: 3, table 1; Cabioch, 1968a: 244 [Roscoff, France]; Pulitzer-Finali, 1977: 62 [Bay of Naples]; BouryEsnault, 1971: 325 [Banyuls]; Pulitzer-Finali, 1983: 573, 610 [Mediterranean]; Wintermann-Kilian & Kilian, 1984: 134 [Colombia; possible misidentification]. MATERIAL. HOLOTYPE: MOM, fragment MNHNDT325. Mediterranean, NE Atlantic. Clathria (Microciona) tumulosa (Bowerbank, 1882) Microciona tumulosa Bowerbank, 1882: 7, 18, 50-52, p1.11, figs 1-4 [Westport Bay, Ireland]; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: BMNH1910.1.1.82. NE Atlantic. Clathria (Microciona) tunisiae sp. nov. Microciona chelifera Levi, 1960a: 70, fig. 12 [Sicily-Tunisian Strait]; Pulitzer-Finali, 1983: 610. MATERIAL. HOLOTYPE: MNHN missing. W Mediterranean. Spanioplon (= Clathria) cheliferum Hentschel, 1911 has seniority. Clathria (Microciona) urizae sp. nov. Eurypon similis; Uriz, 1988a: 53-55, text-fig. 29 [Namibia]. Not Hymeraphia similis Thiele, 1903a: 957. Not Microciona sitnilis Stephens, 1915: 441 [see C.(M) stephensae sp. nov.]. MATERIAL. SPECIMENS: ABIP7B-58, 7B-59, 7B-61. SW Africa. Note: E. similis of Uriz (1988a) is conspecific with neither Clathria (Microciona) similis (Thiele, 1903a), nor Microciona similis Stephens, 1915 (= C. (M.) stephensae sp. nov.), and requires a new name. ETYMOLOGY. Named for Dr. M.J. Uriz. Clathria (Microciona) vacelettia nom. nov. Microciona curvichela Vacelet & Vasseur, 1965: 106-108, p1.9, fig.31 [Madagascar]. Not Wilsonella curvichela Hallmann, 1912: 247. MATERIAL. HOLOTYPE: MNHN missing. W Indian Ocean. C. (D.) curvichela (Hallmann, 1912) has priority. TRANSFERS Other species described in Clathria (Microciona), or a synonym, but now transferred to other genera.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP Microciona ambigua Bowerbank, 1862b: 1110; Bowerbank, 1864: 188; Bowerbank, 1866: 124, 136-138 [Britain]; Gray, 1867: 535; Norman, 1869: 330; Schmidt, 1870: 76; Bowerbank, 1874: 65, p1.25, figs 1-9; Marenzeller, 1878: 2, 4, 5, 14-15, 370, p1.1, fig.3, p1.2, figs 3-3a; Urban, 1880: 257; Bowerbank, 1882:7,18,53; Topsent, 1891a: 528,543, 554 [Roscoff, France]; Norman, 1892:6,11; Vosmaer, 1935a: 607. Hastatusambiguus; Fristedt, 1885: 31-32, p1.3, fig. la-h; Fristedt, 1887: 443, 465. Plocamia ambigua; Topsent, 1894a: 8, 14, 21-22, 23, 26; Topsent, 1895: 214, 216; Topsent, 1896: 115, 127; Topsent, 1898: 226; Topsent, 1900: 112-113; Topsent, 1904a: 10, 24, 26, 154-155, 201 [et var. elegans; Azores]; Arnesen, 1903: 22-23, p1.3, fig.5 [Norway]; Thiele, 1903b: 389, 395, 397, p1.21, fig.21; Arndt, 1913: 119; Topsent, 1913b: 6, 7,32, 63, p1.5, fig.15; Hentschel, 1914: 120; Burton, 1930a: 494 [Norway]; Hentschel, 1929: 895, 973 [White Sea]. Plocamionida ambigua; Topsent, 1927: 1-19; Alander, 1942: 53 [Sweden]; Lilly et al., 1953 [Lough Ine, Ireland]; Burton, 1959b: 39 [Iceland]; Vacelet, 1969: 208 [Mediterranean]; Van Soest & Weinberg, 1980: 10 [Lough Inc. Ireland]; Uriz & Rosell, 1990:387-388, figs 4g-k [Mediterranean]; Ackers, Moss & Picton, 1992: 137 [Ireland]. Scopalina ambigua; Schmidt, 1866a: 149; Schmidt, 1866b: 15; Schmidt, 1868: 26, 40. Amphilectus ambiguus; Vosmaer, 1880: 116. Stylostichon ambiguum; Hanitsch, 1894: 176, 195. Hymedesmia indistincta Bowerbank, 1874: 303-306, p1.87, figs 1-10. Myxilla indistincta; Vosmaer, 1880: 129. Hymeraphia indistincta; Hanitsch, 1894: 181, 196. Plocamia microcionides; Carter, 1876: 390 [Cape St. Vincent]; Topsent, 1891a: 529, 544-545 [Roscoff]; Topsent, 1892a: 117. MATERIAL. HOLOTYPE: BMNH1930.7.3.227, fragment BMNH1910.1.1.65. Referred to Anchinoidae, Plocamionida. Microciona bihamigera Waller, 1877: 261 [Torbay,Britain; nomen nudum]; Waller, 1878: 1, pis 1-2 [new, cf. Zool. Rec. 1877]; Ridley & Dendy, 1887:139; Vosmaer, I935a: 608. Stylostichon bihamigera; Lilly et al., 1953 [Lough 1ne, Ire.]. Pronaxbihamigera; Van Soest & Weinberg, 1980: 10 [Lough Ine,lreland]. MATERIAL. HOLOTYPE: unknown. Referred to Anchinoidae, Pronax. Microciona dives Topsent, 1891a: 529, 543-544, 554, p1.22, figs 2-3 [Roscoff, France]. Stylostichon dives; Pulitzer-Finali, 1983: 567 [Mediterranean]; Ackers, Moss & Picton, 1992: 136 [Ireland]. cf. Hymedesmia zetlandica; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: MOM, fragments BMNH1910.1.1.498-9. Referred to Anchinoidae, probably Plumohalichondria. Microciona fictitia Bowerbank, 1866: 124-126 [Guernsey, Britain]; Vosmaer, 1935a: 607. Hytnedestnia fictitia; Alander, 1942: 36 [ Sweden]. Anchinoe fictitius; Stephens, 1916: 242 [W coast of Ireland]. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 227 Phorbas fictitius; Lilly et al., 1953 [Lough the, Ireland]; Van Soest & Weinberg, 1980: 10 [Lough Inc. Ireland]. MATERIAL. HOLOTYPE: BMNH1930.7.3. 199, fragment USNM5043. Referred to Anchinoidae, Phorbas. Microciona intexta Carter, 1876: 238-239, p1.15, fig.43a-c [Cape St. Vincent, Hebrides]; Vosmaer, 1935a: 607. Rhabderemia intexta; Topsent, 1892a: 116; Topsent, 1904a: 152 [transferred with hesitation]; Hooper, 1990: 72; Van Soest & Hooper, 1993: 337. Rhabdosigma intexta; Hallmann, 1916b: 520; Hallmann, 1917: 398-399. MATERIAL. HOLOTYPE: BMNH1890.4. 10.12. Referred to Rhabderemiidae, Rhabderemia. Microciona minutula Carter, 1876: 479, p1.16, fig.5 I [justified emendation]; Carter, 1880a: 44; Vosmaer, 1935a: 608. Rhabderemiaminutula; Topsent, 1904a: 152-153, p1.1, fig. 10, p1.13, fig. 13 [Banyuls]; Levi, 1956b: 393, fig.2; Boury-Esnault, 1971: 306 [Banyuls]; Biblioni & Gili, 1982: 231; Pulitzer-Finali, 1983: 533-534, text-fig.51 [Mediterranean]. Microciona pusilla Carter, 1876: 239, p1.16; Carter, 1880c: 437; Topsent, 1889: 41, text-fig. 7. Rhabderemia pusilla; Topsent, 1892a: 116; Hallmann, 1917: 399; Dendy, 1922: 85; Van Soest, 1984b: 534; Hooper, 1990: 72 [note]; Van Soest & Hooper, 1993: 323. Rhabderemia indica, in part; Sara, 1961: 44, text-fig. 8; Pulitzer-Finali, 1983: 534. MATERIAL. HOLOTYPE: BMNHI902.11. 16.32, fragment BMNH1954.3.9.178. Referred to RhabderemiicLae, Rhabderemia. Axinella monticularis Ridley & Dendy, 1886: 481; Ridley & Dendy, 1887: 185, p1.38, fig.5 [Cape Verde Is]. Aulospongus monticularis; Hallmann, 1917: 373 [footnote]; Hooper, 1991: 1307 [note]. Microciona monticularis; Burton, 1956: 132 [Sao Vincente, W. Africa]. Aulospongiella monticularis; Burton, 1956: 141. MATERIAL. HOLOTYPE: BMNH1887.5.2.20, paratype BMNH1887.5.2.273. Referred to Raspailiidae, A ulospongus. Spongia plumosa Montagu, 1818: 116 [Devon, UK]. Hymeniacidon plumosa; Bowerbank, 1866:195, figs 141-143. Pronax plutnosa; Gray, 1867: 536. Microciona plumosa; Bowerbank, 1874: 61-63, p1.24, figs 7-13; Topsent, 1891b: 128 [France]. Myxilla (?) plumosa; Ridley & Dendy, 1887: 145-146 [Bahia, Brazil]. Stylostichon plumosa; Topsent, 189Ia: 529; Lilly et al., 1953 [Lough Ine, Ireland]. Plumohalichondria plumosa; Kerville, 1901: 175 [Normandy]. Hymedesmia plumosa; Vosmaer, 1935a: 607. Pronax plumosus; Van Soest & Weinberg, 1980: 10 [Lough [tie, Ireland]. MATERIAL. HOLOTYPE: unknown, fragments BMNH1930.7.3.216, 224. Referred to Anchinoidae, Pronax. MATERIAL. HOLOTYPE: BMNH1902.11. 16.32. Referred to Rhabderemiidae, synonym of Rhabderemia minutula (Dendy, 1905: 180). Microciona quinqueradiata Carter, 1880a: 43, 153, p1.4, figs 5a-e [Gulf of Manaar]. cf.Microciona prolifera; Vosmaer, 1935a: 608. MATERIAL. HOLOTYPE: LFM destroyed, no fragment in BMNH. Referred to Raspailiidae, Cyamon. Microciona simplicissima Norman, 1869: 330; Bowerbank, 1874: 198, 204-205, p1.73, figs 16-19 [Shetland]; Vosmaer, 1935a: 607. Tedania simplicissima; Hanitsch, 1890: 192. Bubaris simplicissima; de Laubenfels, I936a: 131. Not Leptosia simplicissima Hentschel, 1911: 359-360, textfig.40 [Shark Bay, Western Australia]. Not Protoclathria simplicissima Burton, 1932a: 321, p1.56, fig.2, text-fig.31 [Falkland Is]. MATERIAL. HOLOTYPE: BMNH1930.7. 30.212. Referred to Axinellidae, Bubaris. Microciona spinulenta Bowerbank, 1866: 124, 132, 133 [Britain]; Gray, 1867: 534; Vosmaer, 1935a: 607. Pocillon spinulenta; Topsent, 1893b: 34 [plus Isodictya implicita Bowerbank]. MATERIAL. HOLOTYPE: BMNH1930.7. 3.213. Referred to Myxillidae, lophon. Microciona virgula Sara & Siribelli, 1960: 77-79, text-fig.22 [Bay of Naples]; Siribelli, 1960: 16-17, text-fig.7A [Naples]. MATERIAL. HOLOTYPE: IMZUN237: 0.5. Synonym of Antho involvens (Sara, 1964: 228-229). Clathria (Dendrocia) Hallmann, 1920 Dendrocia Hallmann, 1920: 767. Paradoryx Hallmann, 1920: 767. Wilsonella in part; sensu Hallmann, 1912: 242 (not Carter, 1885f: 366). DEFINITION. Single undifferentiated category of smooth auxiliary spicule (style, subtylostyle or modified style) forming plumose or plumoreticulate choanosomal tracts, ectosomal brushes and dispersed between skeletal tracts; echinating acanthostyles usually heavily spined and distributed evenly over skeletal tracts; microscleres include isochelae ranging from typical palmate form (straight shaft, lateral alae fused to shaft), modified palmate forms (thickened, curved shaft, partially detached lateral alae) to anchorate-like forms (alae detached from shaft, shaft with lateral ridge); toxas if present include oxhorns. Microciona pusilla Carter, 1876: 239, p1.16, figs 51a-d [? TYPE SPECIES. Claihria pyramida Lendenfeld, tropical]; Carter, 1880c: 437 [name emended to M. minutula]; Topsent, 1889: 41, text-fig. 7 [Banc de Campeche]. Rhabderemia pusilla; Topsent, 1892a: 116; Hallmann, 1917a: 399; Dendy, 1922: 85; Pulitzer-Finali, 1983: 533-534, fig.51 [Mediterranean]; Van Soest, 1984b: 108. Van Soest & Hooper, 1993: 323. REMARKS. Seven species are included in Clathria (Dendrocia), all of which are endemic to temperate Australian waters, with an hypothesised Gondwanan origin. 1888: 222 (by original designation). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 228zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Dendrocia) curvichela (Hallmann, 1912) (Figs 104-105) Wilsonella curvichela Hallmann, 1912: 247-249, p1.34, fig.4, text-fig.51. Paradoryx curvichela; Hallmann, 1920: 768. Clathria curvichela; Hooper & Wiedenmayer, 1994: 263. Not Microciona curvichela Vacelet & Vasseur, 1965: 106. MATERIAL. HOLOTYPE: AMZ59 (part) + E926a (part): 21km S. of St. Francis I., SA, 32°44'S, 133 ° 18'E, 60m depth, coll. FIV `Endeavour' (dredge). HABITAT DISTRIBUTION. 60m depth; substrate unknown; S Aust (Fig. 104E). DESCRIPTION. Shape. Branching digitate, 130mm high, 35mm maximum width, with short, cylindrical, bifurcate, anastomosing, tapering branches up to 14mm diameter, and short subcylindrical stalk, 15mm long, 8mm diameter. Colour Live colouration unknown, yellowishbrown in ethanol. Oscules. Minute oscules, up to 1 mm diameter, dispersed on lateral margins of branches Texture and surface characteristics. Smooth, even, compressible. Ectosome and subectosome. Ectosome not hispid, although peripheral spongin fibres produce small surface projections; subectosomal auxiliary subtylostyles tangential to surface, or protruding through ectosome only a short way. Choanosome. Choanosomal skeletal architecture almost regularly reticulate, with heavy, thick spongin fibres incompletely divided into primary and secondary components, differentiated only by presence or absence of coring auxiliary megascleres, respectively; spicule tracts in primary fibres vaguely ascend to surface in multispicular bundles; secondary fibres pauci- or aspicular, usually running parallel to surface; all fibres very heavily echinated by acanthostyles, dispersed evenly over fibres; mesohyl matrix heavy but only lightly pigmented, with scattered microscleres. Megascleres. Choanosomal principal styles absent or undifferentiated from auxiliary spicules. Subectosomal auxiliary subtylostyles (coring fibres and scattered below membraneous ectosome) straight or slightly curved, hastate, with subtylote, slightly pointed, smooth bases. Length 132-(149.5)-186p.,m, width 1.543.5)-411m. Acanthostyles subtylote, with large and even spination. Length 64474.1)-8911m, width 4(6.8)-811m. Microscleres. Isochelae palmate, large, with greatly curved shaft; lateral alae fused with shaft about 3/4 way along ala; front ala ranges from well developed to vestigial; poorly silicified sigma-like forms also present. Length 22423.8)32p,m. Toxas absent. REMARKS. Hallmann (1912) originally assigned this species to Wilsonella because it had only one type of auxiliary spicule coring the fibres and scattered in the subectosomal skeleton. However, it lacks detritus in fibres, and principal spicules, and has modified palmate isochelae (curved, thickened with partially detached lateral alae) and heavily echinated fibres indicates that it belongs with Derzdrocia. Hallmann (1920) erected Paradoryx for this and several other species (C. oxyphila, C. piniformis, C. dura and C. elegantula), of which the present species has the most heavily echinated fibres and the most regular fibre reticulation, although slightly plumose near the periphery. Other species of Dendrocia have predominantly plumose (or plumo-reticulate) choanosomal skeletons. Clathria (Dendrocia) dura Whitelegge, 1901 (Figs 106-107, Table 23) Clathria dura Whitelegge, 1901: 83, 84, 117, p1.11, fig. 11; Hooper & Wiedenmayer, 1994: 263. ? Wilsonella dura; Hallmann, 1912: 242, 244, 245, 298; Shaw, 1927: 426; Guiler, 1950: 9. Paradoryx dura; Hallmann, 1920: 768. cf. Microciona prolifera; sensu Vosmaer, 1935a: 611, 644. Not Antherochalina dura Lendenfeld, 1887b: 788. Not Clathria dura var. mo//is Hentschel, 1911: 370. MATERIAL. LECTOTYPE: AMG3046: (dry) Tuggerah Beach, NSW, 3318'S, 151 °30'E, coll. NSW Fish Commission (trawl). PARALECTOTYPE: AMG3046a: (dry) same locality.OTHER MATERIAL: NSW- AMZ1052. HABITAT DISTRIBUTION. Ecology unknown; Tuggerah Beach, Balmoral Beach and Port Jackson (NSW) (Whitelegge, 1901), Maria I. (Tas.) (Shaw, 1927; Guiler, 1950) (Fig. 106E). DESCRIPTION. Shape. Flabellate or flabellate- digitate, lobate, planar or multiplanar growth form, 95-110mm high, 85-210mm wide, composed of fused anastomosing or free digits and flattened lobes, on small compressed basal stalk, 8-15rrim diameter, and branches bifurcate and taper towards apex. Colour Sandy yellow preserved. Oscules. Oscules mostly confined to lateral margins of branches. ^ 229 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS FIG. 104. Claihria (Dendrocia) curvichela (Hallmann) (portion of holotype AMZ59). A, Subectosomal auxiliary subtylostyle. B, Echinating acanthosty le. C, Modified palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. MEMOIRS OF THE QUEENSLAND MUSEUM 230^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 105. Clathria (Dendrocia) curvichela (Hallmann) (portions of holotype A-B, AME926a, AMZ59). A, Choanosomal skeleton. B, Fibre characteristics (x744). C, Subectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Acanthostyle spines. F, Base of auxiliary subtylostyle. G-I, Modified palmate isochelae with full (G) or vestigial front alae (I). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 231 FIG. 106. Clathria (Dendrocia) dura Whitelegge (holotype AMG3046). A, Subectosomal auxiliary style/ subtylostyles. B, Echinating acanthostyles. C, Modified palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. Texture and surface characteristics. Firm, flexible; even, microscopically reticulate and porous. Ectosome and subectosome. Ectosome with nearly continuous palisade of acanthostyles standing erect on peripheral fibres, with plumose brushes of subectosomal auxiliary styles projecting and forming bundles; spicule bundles correspond in position with ascending choanosomal primary spongin fibres; subdermal region also with scattered subectosomal megascleres lying tangential to surface, interdispersed with projecting acanthostyles. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 232zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 107. Clathria (Dendrocia) dura Whitelegge (paratype AMG3046a). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Modified palmate isochelae. E, Echinating acanthostyle. F, Acanthostyle spination. G, Subectosomal auxiliary style. H-I, Bases and apex of auxiliary styles. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 233 TABLE 23. Comparison between present and published records of Clathria (Dendrocia) dura (Whitelegge). Measurements in pm, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Choanosomal principal styles Lectotype Paralectotype Specimen (AMG3046) (AMG3046a) (N=1) absent absent absent 78-(88.2)-102 x 3 - (4.1) - 5.5 82496.5)-115 x 3 - (3.6) - 5 56-(64.6)-75 x 4-(5.6)-9 14-(17.4)-19 61-(66.3)-75 x 4 - (5.1) - 8.5 16-(18.8)-22 absent absent Subectosomal 58487.4)-105 auxiliary x 2 - (3.9) - 6 styles 48-(57.0)-66 x Echinating acanthostyles 4-(5.5)-8 17-(18.6)-22 Chelae Toxas absent Choanosome. Choanosomal skeletal architecture irregularly reticulate, with sinuous, relatively thick spongin fibres; fibres incompletely divided into primary ascending, and secondary transverse components; primary fibres with a multispicular core of subectosomal auxiliary styles, identical to those in ectosomal skeleton, together with heavy tracts of acanthostyles secondarily incorporated into fibres; secondary fibres with multi- or paucispicular core of both subectosomal and acanthostyle megascleres; acanthostyles echinate fibres in moderate quantities, at right angles to skeleton, also lying inside the spicule tracts; mesohyl matrix heavy, darkly pigmented, granular with scattered megascleres and microscleres of all kinds. Megascleres (Table 23). Choanosomal principal megascleres absent or undifferentiated from auxiliary spicules. Subectosomal auxiliary styles quasidiactinal, curved or straight, hastate, with tapering, rounded, styloid, or pointed, smooth bases. Acanthostyles rounded or slightly subtylote, with large, evenly dispersed spines. Microscleres (Table 23). Isochelae abundant, palmate, with lateral alae fused to shaft for only about 1/2 length of ala; shaft greatly curved, thickened ('hunchback'), front ala well developed. Toxas absent. Larvae. Incubated parenchymella, up to 2701Lm in diameter, with heavy mesohyl matrix. REMARKS. In possessing a single category of structural megasclere this species is assigned to C. (Dendrocia). Its quasidiactinal megascleres are reminiscent of those found in C. (D.) pyramida; the modified palmate isochelae (curved, thickened, partially detached lateral alae) are similar to those found in other species assigned to Paradoryx by Hallmann (1920) (e.g., C. (D.) curvichela); and skeletal architecture and spongin fibre characteristics are reminiscent of those in C. (C.) sartaginula. The presence of acanthostyles incorporated into skeletal spicule tracts is now known to occur in several species, collectively termed here the `phorbasiformis' group (including C. (Thalysias) phorbasiformis, C. (T) orientalis, C. (Dendrocia) dura, C. (D.) impelfecta, C. (D.) myxilloides and C. (Clathria) squalorum), and this feature is analogous to (but not homologous with) that seen in the Crellidae. The presence of acanthostyles in the ectosomal skeleton is also seen in the Crellidae. Whereas species of Crella have a thick tangential ectosomal crust of acanthose megascleres, in C. (D.) dura these acanthostyles stand erect on peripheral skeletal fibres, pierced by plumose tracts of smooth subdermal (auxiliary) spicules. The synonymy cited above for C. (D.) dura requires further comment. Lendenfeld (1887a) described Antherochalina dura from Port Jackson, but the species is unrecognisable from his brief description. Lendenfeld's type material was not located in the AM collections, and only a slide of the holotype was found in the BMNH collections (BMNH1886.8.27.684). Apparently a dry specimen of the species was also deposited in the BMNH collections (BMNH1886.8.27.608, possibly the holotype), but this has not yet been rediscovered. Re-examination of the holotype slide confirms that A. dura is a synonym of Phakellia flabellata, as supposed by Burton (1934a). Clathria (Dendrocia) elegantula Ridley & Dendy, 1886 (Figs 108-109) Clathria elegantula Ridley & Dendy, 1886: 474; Rid- ley & Dendy, 1887: 149, p1.28, figs 3,3a, p1.29, figs 14a-b; Guiler, 1950: 7; Carpay, 1986: 26; Hooper & Wiedenmayer, 1994: 263. Clathria elegantula var. occidentalis Hentschel, 1911: 372-374, text-fig.46. Wilsonella elegantula; Hallmann, 1912: 241. Paradoryx elegantula; Hallmann, 1920: 768. MATERIAL. HOLOTYPE: BMNH1887.5.2.91: Moncoeur I., Bass Strait, Tas, 39 ° 14'S, 146 ° 30'E, coll. HMS 'Challenger' (dredge). HOLOTYPE of var. occidentalis: ZMB4445: Top of inner bar, Shark Bay, WA, 25°30'S, 113 ° 03'E, 6-9m depth, coll. W. Michael sen & R. Hartmeyer (dredge). HABITAT DISTRIBUTION. 6-76m depth; on sand and shell grit substrate; Bass Strait and E coast (Tas) MEMOIRS OF THE QUEENSLAND MUSEUM 234zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Clat hria ( Dendrocia) elegant ula Ridley & Dendy (holotype BMNH1887.5.2.91). A, Subectosomal FIG. 108. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA auxiliary style/ subtylostyle. B, Echinating acanthostyles. C, Modified sigmoid palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. G, Section of peripheral skeleton of variety occident alis (holotype ZMB4445). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 235 FIG. 109. Clathria (Dendrocia) elegantula Ridley & Dendy (holotype BMNH1887.5.2.91). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Modified sigmoid palmate isochelae. E, Echinating acanthostyle and modified form. F, Acanthostyle spination. G-H, Bases and apex of subectosomal auxiliary subtylostyles. MEMOIRS OF THE QUEENSLAND MUSEUM 236zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ (Ridley & Dendy, 1886; Carpay, 1986), central W coast (WA) (Hentschel, 1911) (Fig. 108E). DESCRIPTION. Shape. Massive, lobate fan, 73mm long, 75mm wide, with erect, flattened, digitate lobes, 22-31mm long, 8-20mm wide, lobes with uneven margins, arising from sprawling encrusting base. Colour Live colouration unknown, pale beige preserved. Oscules. Differentiated inhalant and exhalant surfaces of lobes; ostia scattered, 1-2mm diameter, oscules confined to sieve-plates. Texture and surface characteristics. Harsh; uneven, prominently microconulose, translucent dermal membrane, covering reticulate fibrous surface, stretched between microconules. Ectosome and subectosome. Fibrous, reticulate ectosomal skeleton, with sparse tangential subectosomal auxiliary subtylostyles lying directly on surface; sinuous plumose tracts of these spicules below, ascending from choanosomal skeleton. Choanosome. Choanosomal skeleton plumoreticulate, cavernous, with well developed primary and secondary spongin fibres; primary ascending fibres sinuous, almost dendritic, cored by several discrete multi- or paucispicular tracts of subectosomal auxiliary subtylostyles, occupying only small portion of fibre diameter; primary fibres interconnected by reticulate secondary fibres, and terminating in plumose spicule bundles at periphery; secondary transverse fibres pauci- or aspicular, thin; echinating acanthostyles most abundant in axial region, sparse or absent in peripheral skeleton; mesohyl matrix light, with abundant scattered chelae. Megascleres. Choanosomal principal megascleres absent or undifferentiated from auxiliary spicules. Subectosomal auxiliary subtylostyles straight, rarely slightly curved, quasidiactinal, hastate points, with tapering, rounded or slightly pointed, smooth bases. Holotype: Length 145-(160.6)168Rm, width 1.5 -(2.6)-4p,m. (Hentschel's specimen: 133-(142.3)-152Rm, width 243.5)4p,m). Acanthostyles slender, evenly spined, with rounded or slightly subtylote bases, sharp points. Length 53-(64.1)-74,m, width 2-(3.6)-5p,m. (Length 52468.2)-84pm, width 3-(6.4)-81.1.,m). Microscleres. Isochelae large, palmate, sigmoid curved, with lateral alae nearly completely fused to shaft, sometimes well developed, sometimes vestigial and reduced to small ridge on shaft; front ala usually reduced. Length 13415.4)-1711m. (Length 13417.2)-2011m). Toxas absent. REMARKS. The nearly dendri tic, predominantly plumose skeletal architecture and the concentration of acanthostyles in the axial region are quite distinctive features for this species. These characters, together with the possession of a single category of structural spicule in both fibres and the ectosomal skeleton indicate that the species is best placed in C. (Dendrocia). In having quasidiactinal modified, vestigial structural megascleres with principal and auxiliary spicules having similar geometry, and vague similarities in skeletal structure, this species is included in the `oxyphila' species group (also containing C. (C.) rap hana, C. (C.) oxyphila and C. (C.) piniformis). Spongin fibres are also characteristic (whereby each fibre may have several discrete spicule tracts), but in most other respects this species is very similar to C. (D.) dura Whitelegge (both having sinuous spongin fibre systems and heavily echinated fibres). These two species differ in the size of their auxiliary styles (notably straight, longer, more slender, with subtylote bases in C. (D.) elegantula, whereas in C. (D.) dura these spicules are curved, short, thick, with tapering hastate (or sometimes pointed) bases)), and the incorporation of acanthostyles into skeletal fibre tracts in C. (D.) dura. There is a remarkably close concordance in these features between Bass Strait and Shark Bay populations, despite their widely separated geographical distribution. These samples differ only slightly in the more robust acanthostyles seen in Shark Bay material, but no other morphological differences were seen to justify subspecies separation (Hentschel, 1911). Clathria (Dendrocia) imperfecta Dendy, 1896 (Figs 110-111) Clathria imperfecta Dendy, 1896: 35; Ayling et al., 1982: 103; Hooper & Wiedenmayer, 1994: 264. Wilsonella imperfecta; Hallmann, 1912: 242. MATERIAL. HOLOTYPE: NMVG2369 (RN376) (fragment BMNH1902.10.18.335): Port Phillip Bay, Vic, 38°09'S, 144°52'E, 36m depth, coll. J.B. Wilson (dredge). HABITAT DISTRIBUTION. 36m depth; substrate unknown; Port Phillip Bay (Vic) (Fig. 110E). DESCRIPTION. Shape. Erect, bulbous-lobate, with small cylindrical lobate digits. Colour Colour in life recorded as dull brownorange exterior, yellow interior. ^ 237 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR Oscules. Minute, less than 2mm diameter, mainly on apex of lobes. Texture and surface characteristics. Surface irregular, uneven, micropapillose, with subdermal ridges and grooves. Texture is crumb-of-bread, easily torn. Ectosome and subectosome. Microscopically hispid, with thick brushes of subectosomal auxiliary styles from peripheral skeleton protruding through dermal crust, composed of same megascleres, lying paratangential to or standing erect on surface; subectosomal architecture thickly plumose, arising from ultimate ascending choanosomal fibres. Choanosome. Choanosomal skeleton irregularly plumo-reticulate, with prominent primary ascending spicule tracts interconnected by irregularly dispersed secondary tracts; fibres weakly developed in spongin but fully cored by subectosomal auxiliary styles and also acanthostyles secondarily incorporated into tracts; fibres moderately echinated; fibre anastomoses form irregular oval-elongate meshes; mesohyl matrix heavy but only lightly pigmented, with auxiliary styles and sparse detritus scattered thrroughout. Megascleres. Choanosomal principal styles absent or undifferentiated from auxiliary megascleres. Subectosomal auxiliary spicules fusiform, mostly straight, occasionally slightly curved towards bases, with tapering, or rounded, or slightly subtylote smooth bases. Length 189(213.3)-234p.m, width 1.544.4)-7.5p.m. Acanthostyles long, rounded or faintly subtylote, with large and evenly distributed spines. Length 102-(114.8)-122p,m, width 5-(5.8)-7p.m. Microscleres absent. Larvae. Oval-elongate parenchymella, 345412x275-370p.m, without larval spicules but with heavy mesohyl and differentiated cells clearly visible. REMARKS. This species was assigned to Wilsonella by Hallmann (1912) because it has only a single category of smooth megasclere, but was tacitly returned to Clathria by Hallmann (1920) when he restricted Wilsonella to the type. Clathria (Dendrocia) impeifecta is more appropriately placed in Dendrocia because it lacks detritus incorporated into fibres and has only one undifferentiated category of smooth auxiliary style, whereas Wilsonella (sensu Hallmann, 1912) and Paradoryx (sensu Hallmann, 1920) have two categories of auxiliary styles, one found in fibres and the other on the ectosomal skeleton. All three nominal genera lack true principal megascleres. This species is similar to C. (Thalysias) orientalis, C. (T) phorbasifonnis, C. (D.) myxilloides and C. (D.) dura in having acanthostyles incorporated into fibres, intermingled amongst the smooth coring megascleres (termed the `phorbasifonnis' species complex). Clathria (Dendrocia) myxilloides Dendy, 1896 (Figs 112-113, Plate 3A) Clathria myxilloides Dendy, 1896: 35; Hallmann, 1920: 768; Ayling et al., 1982: 104; Hooper & Wiedenmayer, 1994: 264. Wilsonella myxdloides; Hallmann, 1912: 242. MATERIAL. HOLOTYPE: NMVG2376 (RN729) (fragment BMNH1902.10.18.334): Port Phillip Bay, Vic, 38 ° 09'S, 144 ° 52'E, coll. J.B. Wilson (dredge). OTHER MATERIAL: S AUST- QM G300613 (NCIQ66C-2202-K, fragment NTMZ3535). HABITAT DISTRIBUTION. 18m depth; on rock substrate; Port Phillip Bay (Vic), Kangaroo I. (SA) (Fig. 112E). DESCRIPTION. Shape. Massive, subspherical, lobate digitate growth form. Colour. Orange-red alive (Munsell 5R 5/10), grey-brown in ethanol. Oscules. Small, less than 3mm diameter, scattered evenly over surface. Texture and surface characteristics. Harsh, compressible; shaggy, uneven, microconulose, appearing pock-marked alive due to scattering of small oscules. Ectosome and subectosome. Relatively dense discrete bundles of subectosomal auxiliary styles, identical to those coring the fibres, forming erect palisade on surface. Tracts of isochelae also found in peripheral skeleton, scattered throughout heavily pigmented peripheral spongin; subectosomal region relatively disorganised, paratangential, merging into peripheral choanosomal fibres lying almost immediately subectosomal. Choanosome. Choanosomal skeletal architecture irregularly plumo-reticulate, without clearly differentiated primary or secondary components although fibre diameter varies considerably and fibres become sinuous towards surface; fibres composed of only light spongin, heavily cored by multispicular tracts of subectosomal auxiliary styles and heavily, irregularly echinated by acanthostyles lying parallel with, or at acute angles to spicule tracts but also secondarily incorporated into fibres; mesohyl matrix light, with smaller, MEMOIRS OF THE QUEENSLAND MUSEUM 238zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 110. Clathria (Dendrocia) impelfecta Dendy (holotype NMVG2369). A, Subectosomal auxiliary styles. B, Echinating acanthostyle. C, Section through peripheral skeleton. D, Parenchymella larva in situ (diameter 400um). E, Australian distribution. F, Holotype. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 239 FIG. 111. Clathria (Dendrocia) impetfecta Dendy (holotype NMVG2369). A, Choanosomal skeleton. B, Fibre characteristics (x419). C, Echinating acanthostyle. D, Acanthostyle spination. E, Bases of subectosomal auxiliary styles. MEMOIRS OF THE QUEENSLAND MUSEUM 240^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 112. Clathria (Dendrocia) myxilloides Dendy (holotype NMVG2376). A, Subectosomal auxiliary subtylostyle. B, Echinating acanthostyle. C, Anchorate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, QMG300613 alive. G, Holotype. thinner auxiliary spicules, and numerous isochelae organised into tracts surrounding large choanocyte chambers. Megascleres. ^Principal^choanosomal megascleres absent, or undifferentiated from auxiliary spicules. Subectosomal auxiliary styles differ slightly in size according to location within skeleton: those coring fibres and in ectosomal skeleton relatively homogenous, straight or slightly curved, hastate, with smooth, tapering or slightly subtylote bases, some appearing quasi-diactinal. Length 287(311.4)-330p,m, width 3.5-(4.9)-6Rm; auxiliary styles also scattered between fibres probably younger forms of main structural megascleres, being thin, whispy, hastate, usually curved or ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 241 FIG. 113. Clathria (Dendrocia) myxilloides Dendy (holotype NMVG2376). A, Choanosomal skeleton. B, Fibre characteristics. C, Acanthostyle spination. D, Echinating acanthostyles. E, Anchorate, unguiferous and sigmoid isochelae. 242zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM sinuous, with rounded or tapering bases. Length 212-(233.0)-284Rm, width 2-(2.2)-31.1.m. Acanthostyles long, slender, straight or slightly curved near basal end, evenly and heavily spined. Length 84-(112.6)-133t.tm, width 6-(7.4)-10iLm. Microscieres. Isochelae relatively large, abundant, anchorate, with well formed lateral alae detached from shaft nearly completely, lateral ridge on shaft (possibly vestigial point of attachment of lateral alae); some modified unguiferous forms with pointed alae, or sigmoid forms with greatly reduced alae also present. Length 21(24.4)-29Rm. Toxas absent. REMARKS. This species is similar to C. (D.) impetfecta in growth form and incorporation of some acanthostyles into fibres (see remarks for C. (Thalysias) phorbastfonnis). It also shows similarities to C. (D.) pyratnida in growth form and megasclere morphology, although all three species differ in isochelae geometry. Clathria (Dendrocia) pyramida Lendenfeld, 1888 (Figs 114-115, Table 24, Plate 3B) Clathria pyramida Lendenfeld, 1888: 222; Capon & MacLeod, 1987: 1200; Chernoff, 1987: 160; Hooper et al., 1992: 263; Hooper & Wiedenmayer, 1994: 264. Wilsonella pyramida; Hallmann, 1912: 240, 244; Shaw, 1927: 426; Guiler, 1950: 9. Dendrocia pyramida; Hallmann, 1920: 767. Clathriaalata Dendy, 1896: 34; Hentschel, 1911: 375377, text-fig.48; Hentschel, 1923: 387, fig.356; Ayling et al., 1982: 100; Van Soest, 1984b: 120. Wilsonella alata; Hallmann, 1912: 241. Dendrocia alata; Hallmann, 1920: 767. MATERIAL. LECTOTYPE: AMG9047: Port Jackson, NSW, 33°51'E, 151 ° 16'E, no other details known. PARALECTOTYPE: BMNH1887.1. 24.62 (fragment AMG3579): same locality. LECTOTYPE of C. alata: NMVG2280 (fragment BMNH1902.10.18.330): Port Phillip Bay, Vic, 38°09'S, 144°52'E, coll. J.B. Wilson (dredge, RN 752). PARALECTOTYPES of C. alata: NMVG2281 (fragment AMG2686): same locality (RN 763). NMVG2282 (fragment BMNH1902.10.18.31): Sorrento Jetty, Port Phillip Bay, 38°33'S, 145 ° 21'E (dredge, RN 792). NMVG2283 (fragment BMNH1902.10.18.49): same locality (RN 801). NMVG2284 (fragment BMNH1902.10.18.332): same locality (RN 842). NMVG2285 (fragment BMNH1902.10.18.333): same locality (RN 843). OTHER MATERIAL: VIC- AME298, AMZ1145, AM unreg. (small vial, label 'donated by A Dendy'), AM unreg. (fragment BMNH1887.4.27.103) (small vial, label 'Clathria tethyopsis, donated by A Dendy, Lendenfeld or Burton MS name'). NSW- AMZ3216, NTMZ2667, QMG304507, QMG304522, QMG304574. S AUST- SAMTS4095 (fragment NTMZ1629), SAMTS 6290, QMG3005 03 (NCIQ66C-2119-T) (fragment NTMZ3520), QMG301354, QMG301358, QMG301374, QMG301382, QMG304041, QMG304049, QMG304051, QMG304053, QMG304054, QMG304063, QMG304064, QMG304069, QMG304070. WA- QM G300604 (NCIQ66C-4271J), QM G300191 (NCIQ66C-4651-V). (Not AM unreg. (label 'Clathria alba. Port Jackson, NSW', = Lendenfeld or Burton MS name; = Crella incrustans)). HABITAT DISTRIBUTION. 0.8-25m depth; on sand, rock reef or muddy bottoms; associated with oysters and algal beds, Spondylus, Pinna and Chalamus asperrimus bivalves, rock or wood jetty substrates; Illawarra, Port Jackson, N. Sydney, Port Hacking and Durras I. (NSW); Maria I. (Tas); Port Phillip Bay (Vic); St. Vincent Gulf, Yorke Peninsula and Kangaroo I. (SA); Albany and Pelsart Is, Houtman Abrolhos (WA) (Fig. 114E). DESCRIPTION. Shape. Massive, lobate, lobo- digitate, irregular growth form with small lobate surface projections. Colour. Dark brown or orange-brown live exterior (Munsell 5YR 4/6-5/10), paler choanosome, brown in ethanol. Oscules. Few, relatively large oscules (up to 8mm diameter) on apex of lobes, each with slightly raised membraneous lip, and with membraneous, divided exhalant canals visible inside oscule; oscules collapse on preservation. Texture and surface characteristics. Firm, compressible; smooth, relatively even, bulbous, fleshy surface in live state; optically smooth, membraneous surface in preserved state, with irregularly rugose microscopic conulose and lobate projections. Ectosome and subectosome. Microscopically hispid, with dense crust of subectosomal auxiliary styles forming continuous, erect, regular (straight) or irregular (stellate, paratangential), plumose palisade. Choanosome. Skeletal architecture irregularly plumo-reticulate, with sinuous spongin fibres ascending to surface in meandering tracts; fibres anastomose more frequently at axis than in peripheral skeleton; peripheral fibres and skeletal tracts often diverge becoming plumose in subectosomal region, or forming a paratangential layer immediately below erect ectosomal skeleton; fibres contain moderately light to very heavy spongin, cored by multispicular, sinuous tracts of subectosomal auxiliary styles; fibres not obviously divided into primary or secondary components, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 243 although fibre diameter varies within same sections of skeleton; acanthostyles heavily echinating most fibres, occasionally rare or absent from some; mesohyl matrix very heavy but only lightly pigmented, with abundant isochelae and auxiliary megascleres; microalgae present in mesohyl of some specimens. Megascleres (Table 24). Choanosomal principal megascleres absent, or at least undifferentiated from auxiliary spicules. Subectosomal auxiliary styles are thin, hastate, straight or slightly curved towards basal end, usually with smooth, slightly subtylote bases that taper towards end into a small point (partially mucronate), points hastate. Acanthostyles vary considerably in length, usually club-shaped, subtylote, with large spines mostly confined on base and more-or-less aspinose towards point; spines usually robust. Microscleres (Table 24). Isochelae large, very abundant, primarily palmate, heavily silicified, usually with thickened and slightly curved shaft, large lateral alae completely fused to shaft resembling swings', with front ala free, but sometimes with reduced alae and sigmoid curvature. Toxas absent. Larvae. Larvae oval-elongate parenchymella, up to 518x34011m, with larval raphidiform spicules scattered throughout the axis and forming stellate tufts at the posterior pole. REMARKS. Type material of Clathria pyramida and of C. alata are virtually identical in their growth form, surface features, spicule geometry, spicule distribution and fibre characteristics, differing only slightly in choanosomal skeletal construction (predominantly plumo-reticulate versus predominantly plumose to slightly plumo-reticulate), and ectosomal structure (either producing a continuous erect palisade (C. alata s.s.) or punctuated palisade resulting in stellate plumose brushes (C. pyramida s.s.), respectively). But there are no consistent or significant differences between these species in the spongin content of fibres, density of spicule tracts, presence or absence of conecting secondary fibres, or spicule dimensions (Table 24), as supposed by Hallmann (1912: 241). On this basis the two species are merged here. For all other specimens examined the main variability involves the degree to which fibres were compacted (mesh size), the spongin content of skeletal tracts ('fibre' diameter), and spicule dimensions between different geographic populations (Table 24). In this latter regard populations from NSW and WA had relatively larger, more robust acanthostyles than other populations (VIC, SA). In particular, spicules from a small population in the Gulf of St Vincent, SA, were noticeably more poorly silicified and less robust than 'typical' specimens found in all other localities. In spiculation and skeletal architecture C. (D.) pyramida is remarkably similar to Crella incrustans var. digitata ('cotype' AME503), although upon careful examination of both species there are obvious differences in skeletal structure (Crella with a tangential ectosomal layer), composition of the ectosomal skeleton (styles versus acanthostyles or acanthoxeas), and spicule geometry (quasi-diactinal auxiliary styles versus anisoxeas or quasi-monactinal megascleres, respectively). Nevertheless, these species demonstrate remarkably close convergences in several prominent features making them easily confused in the field and laboratory. Specimens from the south coast of NSW have peculiar biochemistry and significant biological activity against both gram positive and gram negative bacteria (Hooper et al., 1992). Unlike many other bioactive sponges, however, the activity in C. (D.) pyramida was found to be related to a unique, modified free sugar, 5-Thio-D-mannose (Capon & MacLeod, 1987). This discovery represents the only known occurrence to date of that class of thiosugar in nature, although the compound has a glucose-based synthesised analogue which has been known for several years. Thio sugars have considerable pharmaceutical and medical potential: they inhibit the release of insulin and transport of glucose, and are able to cause reversible inhibition of sperm-cell development without displaying acute toxicity (R. Capon, pers. comm.). The taxonomic significance of those compounds is not clear, but it is possible to speculate on the biological role of those chemicals. Recent evidence (A. Butler, pers.comm.) suggests that C. (D.) pyramida together with a few other species (e.g., Crella incrustans) occur in association with (are epizootic on) scallops (Chlamys asperrima) from South Australian waters. Chemoff (1987) found that the presence of those epizootics increased the survival of the scallop in both field and caging experiments and it is possible that the sponges provide some sort of chemical defence of the host: thio-mannose sugar may play some sort of role in that defence. 244zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 114. Clathria (Dendrocia) pyramida Lendenfeld (holotype AMG9047). A, Subectosomal auxiliary subtylostyle. B, Echinating acanthostyle. C, Modified palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. G, Paralectotype of C. alata NMVG2283. H, QMG300238 in situ. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 245 FIG. 115. Clathria (Dendrocia) pyramida Lendenfeld (QMG301358). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spination. F-G, Modified thickened and sigmoid palmate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM 246^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Dendrocia) pyram ida TABLE 24. Comparison between published records and different populations of zyxwvutsrqponmlkjihgfedcbaZYXWVUTS Lendenfeld. All measurements are given in pm, denoted as range (and mean) of spicule length x spicule width (N=25) (bracket = number of specimens per locality examined). -^ - - Lectotype (AMG9047) SPICULE Lectotype of C. Alata (NMVG2280) Specimens NSW (N=7) Specimens VIC (N=10) Specimens SA (N=16) Specimens WA (N=2) Choanosomal principal styles absent absent absent absent absent absent Subectosomal auxiliary styles 208-(220.6)-231 x 4-(6.4)-8 211-(229.4)-243 x 3-(4.8)-7 190-(209.5)-225 x 4-(5.1)-7 2024222.6)-247 x 4-(5.9)-9 195-(213.4)-228 x 4-(4.9)-7 82-(103.9)-145 x 5-(8.4)-12 68-(89.6)-I25 x 4(7.2)-12 64-(78.6)-90x 4(7.4)-11 214(227.4)-242 x 4-(5.3)-7 82-(103.5)-155 x 6-(9.0)-11 20423.6)-27 20-(22.5)-25 17-(22.1)-30 22-(24.3)-27 Echinating acanthostyles Chelae 87-(99.0)-135 x 8- 75-(85.2)- 118 x 7(10.2)-12 (9.2)-11 21-(23.8)-27 23-(24.5)-27 Clathria (Dendrocia) scabida (Carter, 1885) (Figs 116-117, Table 25) Carter, 1885b: 112, p1.4, figs 4-5; Carter, 1886g: 449. Micr ociona scabida; Dendy, 1896: 31; Hal'mann, 1912: 150; Vosmaer, 1935a: 608. ? St ylot ellopsis or Clat hrissa scabida; Hallmann, 1912: 151. Anaat a scabit a Papsusl; de Laubenfels, 1936a: 109. Clat hria scabida; Hooper & Wiedenmayer, 1994: 264 Halichondria pust ulosa, in part; Carter, 1886g: 450. Not Halichondria pust ulosa Carter, 1882a: 285, p1.11, fig. 1. Halichondr ia scabida MATERIAL. HOLOTYPE: BMNH1887.7.11.9 (fragment AMG2760): Port Phillip, Vic, 38°09'S, 144 ° 52'E, coll. J.B. Wilson (dredge). OTHER MATERIAL: VIC - NMVRN413, NMVRN1025, NMVRN1038, BMNH1887.7.11.26 (slide AMG2916). HABITAT DISTRIBUTION. 38m depth; substrate unknown; Port Phillip and Westernport Bays (Vic) (Fig. 116F). DESCRIPTION. Shape. Low, globular, erect, arborescent, digitate sponge, 80-95mm high, 4570mm wide, with subspherical, lobate branches, 28-42mm long, 12-28mm diameter, on a short stalk, 22mm long, 12-15mm diameter. Colour Live colouration orange-brown, brown exterior and yellow-brown interior preserved. Oscules. Large oscules, up to 5mm diameter, situated in groups (seive-plates) on all sides of branches; seive-plates sunken, resembling porocalyses of the Tethyidae. Texture and surface characteristics. Surface uneven, fibrous, with numerous large pores, seiveplates and subdermal canals covered with prominent membraneous skin, raised into small papillae and ridges. Ectosome and subectosome. Ectosome hispid, with subectosomal auxiliary subtylostyles form- ing sparse but continuous dermal palisade of erect brushes, sometimes tangential, protruding from surface, piercing heavy crust of isochelae lying on ectosomal membrane; subectosomal region plumose, with sinuous tracts of subectosomal subtylostyles diverging from ends of choanosomal fibres, and with numerous, relatively large subdermal cavities lined by isochelae. Choanosome. Choanosomal skeleton irregularly plumo-reticulate, with sinuous, heavy, spongin fibres cored by subectosomal auxiliary styles and fewer larger acanthostyles, and echinated by two sizes of acanthostyles; smaller acanthostyles echinate fibres in relatively heavy, even concentrations, whereas larger acanthostyles most common at fibre nodes and echinate fibre terminations in radiating brushes; spongin fibres not divided into primary or secondary components, and fibre anastomoses occur irregularly throughout skeleton; mesohyl matrix very heavy, with abundant isochelae scattered and also congregated around fibres and choanocyte chambers. Megascleres (Table 25). Choanosomal principal megascleres absent or at least completely undifferentiated from auxiliary spicules. Subectosomal auxiliary subtylostyles, coring fibres and in dermal membrane, long, thin, mostly straight, hastate, with elongated, subtylote, smooth bases. Larger acanthostyles subtylote, slightly curved, with sparse but large spines along shaft and base, and usually with an aspinose apex. Smaller acanthostyles subtylote, evenly and heavily spined. Microscleres (Table 25). Isochelae extremely abundant, anchorate-like, with curved shaft, lateral alae nearly completely detached from shaft, completely free front ala, and lateral ridge on shaft (possibly vestigial point of alae attach- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 247 TABLE 25. Comparison between present and published records of Clathria (Dendrocia) scabida (Carter). Measurements in i.tm, denoted as range (and mean) of spicule length x spicule width (N=25). SPICULE Holotype (BMNH1887.7.11.9) Specimens (N.3) absent absent Choanosomal principal styles 201-(237.2)-273 x 3- 1964226.5)-278 x 2(4.4)-5.5 (3.7)-5 153-(191.6)-219 x 193-(201.1)-221 x6Echinating acanthostyles I 5.5-(7.1)-9 (7.3)-9 78490.4)-98 x 382-(95.3)-105 x 5Echinating acanthostyles II (6.8)-10 (6.4)-8 16-(18.2)-21 19-(20.9)-24 Chelae I (sigmoid) Subectosomal auxiliary styles Chelae II (arcuatelike) 24 - (29.2) - 33 28-(31.4)-34 Chelae III (bipocilla-like) 14416.2)-18 18-(19.3)-22 35-(72.4)-120x 1(1.8) 3 uncommon 70-(121.1)-156x 1- Toxas - (2.2)-3 common known species of Clathria (Dendrocia) with toxa microscleres. Clathria (Axociella) Hallmann, 1920 Axociella Hallmann, 1920: 779; Bergquist & Fromont, 1988: 116. Axosuberites Topsent, 1893a: 179. Tenaciella Hallmann, 1920: 772. DEFINITION. Well differentiated axial and extra-axial skeletal architecture (reminiscent of Raspailiidae); axial skeleton markedly compressed, reticulate; extra-axial skeleton radial, plumose or plumo-reticulate composed of large subectosomal auxiliary styles-subtylostyles; specialised ectosomal skeleton present composed of smaller auxiliary spicules; echinating megascleres absent although principal spicules may protrude through fibres at acute angles. TYPE SPECIES. Esperiopsis cylindrica Ridley & Dendy, 1886: 340 (by original designation). ment); reduced chelae also present, unguiferous, with pointed alae. Toxas oxhorn, with wide central curvature and slightly reflexed arms. REMARKS. Carter's (1885a) holotype (BMNH1887.7.11.9), and his (1886g) specimen of H. pustulosa, BMNH1887.7.11.26 are identical, as suspected from their respective descriptions, whereas H. pustulosa from the Falkland Is (Carter, 1882a) is different. Clathria scabida has skeletal architecture, fibre structure, ectosomal characteristics and spicule geometry characteristic of Dendrocia, whereas the presence of a second, larger size class of acanthostyle (which may be incorporated into skeletal spicule tracts as well as echinating the fibre endings and fibre nodes) is unusual to the genus. Possession of hymedesmoid-plumose or plumose skeletal architecture links several microcionids termed the scabida' group (C. (D.) scabida, C. (Microciona) si,nilis, C. (M.) hentscheli, C. (M.) tetrastyla, C. (M.) thielei, C. (Thalysias) tingens sp. nov. and C. (T) distincta). Earlier records of this species (Carter, 1885a, 1886g; Hallmann, 1912) differentiate isochelae microscleres into 3 or 4 morphs, but all isochelae in both specimens are not as vastly different as supposed by Carter (1885a). They are not arcuate, as supposed by previous authors, but anchorate with completely detached lateral alae and ridges on the lateral sides of shaft. This is the only REMARKS. Axociella is represented in Australasian waters by six species, one new, all of which have very well differentiated skeletal structures closely resembling the compressed skeletons common in Raspailidae. The existence of microcionids with compressed axial skeletons and differentiated axial and extra-axial skeletons supports the proposal for a closer relationship between Raspailiidae and Microcionidae, as proposed by Hooper (1990a, 1991), and formalised further by Hajdu et al. (1994) in their subordinal classification of Poecilosclerida. Crucial characters differentiating the Axociella group from typical Raspailiidae are the possession of chelae and absence any true echinating spicules in the former, versus possession of a specialised ectosomal skeleton (composed of small styles or oxeas in brushes surrounding larger protruding spicules) in the latter (see Hooper, 1991). Clathria parva Levi (from S and SW Africa) and Axosuberites fauroti Topsent (from the Gulf of Aden), are also referred here to C. (Axociella) because they have well differentiated axial and extra-axial regions, whereas 3 NZ species (Bergquist & Fromont, 1988) are retained in this group only on a tentative basis since their skeletal structures are not typical of Axociella. Other species referred to Axociella e.g., de Laubenfels, 1936a) are simply encrusting and/or lack echinating acanthostyles (a secondary loss common amongst Microcionidae and Raspailiidae), and do not belong to Axociella as defined here. 248zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 116. Clathria (Dendrocia) scabida (Carter) (NMVRN1038). A, Subectosomal auxiliary style/ subtylostyle. B, 2 sizes echinating acanthostyle. C, Anchorate-like isochelae. D, Oxhom toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype BMNH1887.7.11.9. H, NMVRN1025. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 249 'FIG. 117. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hr ia ( Dendr ocia) scabida (Carter) (NMVRN1025). A, Choanosomal skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spination. F, Oxhorn toxas. G, Anchorate isochelae. H, Uniguiferous anchorate isochelae. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 250zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Axociella) canaliculata (Whitelegge, 1906) (Figs 118-119) Esperiopsis canaliculata Whitelegge, 1906: 471, p1.43, fig.7 Tenaciella canaliculata; Hallmann, 1920: 773, p1.36, figs 1-2, p1.37, fig.1, text-fig.1; Ristau, 1978: 585586. Artemisina canaliculata; Ristau, 1978: 585-586. Rhaphidophlus canaliculata; Van Soest, 1984b: 115. Clathriacanaliculata; Hooper & Wiedenmayer, 1994: 262. MATERIAL. LECTOTYPE: AMG4325: Off Wata Mooli, Bulgo, Woolongong, NSW, 34 °30'S, 151 ° 10'E, 104-126m depth, 01.iii.1900, coll. FIV 'Thetis' (dredge). PARALECTOTYPE: AMZ988: Off Woolongong, NSW, 34°25'S, 151 ° 10'E, 220-224m depth, 01.iii.1900., coll. FIV 'Thetis' (dredge). OTHER MATERIAL: QLD- QMG300460 (fragment NTMZ1562). HABITAT DISTRIBUTION. Growing in soft sediments, associated with gorgonian beds in deeper waters of the continental shelf; 94-126m depth; Wollongong (NSW), S. Stradbroke I. (SEQ) (Fig. 118G). DESCRIPTION. Shape. Erect, whip-like, cylindrical branching digits, 150-190mm long, with short cylindrical stalk 30-50mm long, 1015mm diameter, and few thick subcylindrical branches, mostly in 1 plane, bifurcating and occasionally anastomosing. Colour Drab grey in ethanol. Oscules. Small, less than 2mm diameter, dispersed over lateral margins of branches. Texture and surface characteristics. Firm, flexible; surface smooth, fleshy, velvetty, feltlike, slightly porous, without obvious ornamentation. Ectosome and subectosome. Thick, erect, plumose brushes of ectosomal auxiliary subtylostyles form dense, continuous palisade on exterior surface; discrete tracts of larger subectosomal auxiliary styles form thick, multispicular, ascending or paratangential brushes supporting ectosomal skeleton. Choanosome. Partially compressed, openmeshed reticulate axial skeleton and radial extraaxial skeleton, becoming plumose in periphery; skeleton composed of very heavy spongin fibres of large diameter, up to 250p,m diameter, forming ovoid meshes, and clearly divided into primary ascending and secondary connecting elements, differing substantially in diameter; fibres cored by multispicular tracts of choanosomal principal styles occupying most of fibre diameter; spicule content of fibres increases towards periphery; secondary fibres less heavily cored; echinating megascleres absent; mesohyl matrix moderately heavywith scattered microscleres, but megascleres largely confined within fibres except at periphery. Megascleres. Choanosomal principal styles thick, slightly curved or rarely straight, fusiform, with smooth, rounded or very slightly subtylote bases. Length 130-(318.7)-465p,m, width 4(21.2)-26pm. Subectosomal auxiliary styles long, thick, straight, fiisiform, with tapering rounded or very slightly subtylote bases, usually smooth, occasionally microspined. Length 2404465.0)590p,m, width 10-(12.2)-16p.m. Ectosomal auxiliary subtylostyles straight or slightly curved, with microspined subtylote bases, fusiform points. Length 80-(114.6)165m, width 5-(6.1)-8p,m. Microscleres. Palmate isochelae with long lateral alae completely fused to shaft and fused front ala; chelae clearly differentiated into two size classes, the smaller often twisted. Length I: 4-(4.6)-8jun; length II: 14-(17.5)-22p.m. Toxas accolada, usually long, thin, slight central curvature and straight or only slightly reflexed points. Length 184220.8 )-55011m, width 0.541.9)-3.311m. REMARKS. Hallmann (1920) erected Tenaciella for this species on the basis that it lacked echinating spicules and had a Thalysias-like ectosomal skeleton. Simpson (1968a) found no correlation between these 2 features and histological evidence to support recognition of this genus distinct from Clathria. Spicule geometry and the ectosomal skeleton in C. (A.) canaliculata indicate relationship with Thalysias, whereas skeletal structure (particularly the radial choanosomal skeleton), is reminiscent of Raspaila nuda (Hooper, 1991). The shared skeletal structure in Axociella and Raspailiidae support the contention that they are monophyletic, belonging to the suborder Microcionina (Hajdu et al., 1994). The well developed felt-like ectosomal spiculation and radial subectosomal spicule tracts in the peripheral skeleton of C. (A.) canaliculata is virtually at the opposite end of a continuum from encrusting Microciona-like species (with membraneous dermal skeletons). These differences contrast so greatly with species such as C. (M.) aceratoobtusa that it is tempting to maintain generic separation between typical Microciona and Thalysias-like species, but as demonstrated REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 251 elsewhere in this work there are too many other microcionid species with intermediate conditions making it impossible to clearly define a generic boundary within this continuum. Clathria(Axociella)canaliculata is most closely related to C. (A.) cylindrica, with similar growth form, surface characteristics and skeletal structure. However, this species has slightly compressed open-reticulate axis, radial extra-axis, and continuous plumose ectosome, whereas C. (A.) cylindrica has a greatly compressed closemeshed axis, a radial extra-axis and a sparse, discontinuous, plumose ectosome. Toxa morphology and spicule size also differ substantially between these two species. CIathria (Axociella) cylindrica (Ridley & Dendy, 1886) (Fig. 120-121) Esperiopsis cylindrica Ridley & Dendy, 1886: 340; Ridley & Dendy, 1887: 79-80, p1.19, figs 2a-b. Axociella cylindrica; Hallmann, 1920: 780-783, p1.37, figs 2-4, text-fig.2; Not Axociella cylindrica; Sim & Byeon, 1989: 39-40, p1.5, figs 1-2. Clathria cylindrica; Hooper & Wiedenmayer, 1994: 262. Not Rhaphidophlus cylindricus Kieschnick, 1900: 53, p1.44, fig.10. MATERIAL. HOLOTYPE: BMNH1887.5.2.96: Off Port Jackson, NSW, 33°51'S, 151 ° 16'E, 60-70m depth, coll. HMS 'Challenger' (dredge). OTHER MATERIAL: NSW- AMZ1527. HABITAT DISTRIBUTION. Soft sediments; 40-70m depth; Port Jackson, Botany Bay (NSW) (Fig. 1200). DESCRIPTION. Shape. Thin, cylindrical digitate, branching, whip-like, 130-190mm long, 10-20mm diameter, with thin, dichotomously branched, cylindrical or slightly flattened, distally tapering branches, up to 130mm long, 16mm diameter, short stalk and expanded basal attachment. Colour Grey-brown in ethanol. Oscules. Not seen. Texture and surface characteristics. Firm, flexible, tough consistency; surface even, feltlike, unornamented, prominently hispid. Ectosome and subectosome. Sparse, plumose brushes of small ectosomal auxiliary subtylostyles form discrete, discontinuous bundles on surface, arising from ends of radial skeletal columns, scattered around the larger, protruding subectosomal auxiliary styles which project some distance through surface. Choanosome. Skeletal structure with clearly differentiated axial and extra-axial components, markedly compressed in axis and radial in extraaxis; compressed axial skeleton with longitudinal fibres cored by short choanosomal principal subtylostyles, interconnected by thinner pauci- or aspicular fibres; axial spongin fibres heavy, and fibre anastomoses very close-meshed forming elongate reticulation; mesohyl matrix in axis light with abundant loose auxiliary styles dispersed between and congregated around fibres; echinating megascleres absent; radial extra-axial skeleton with large auxiliary styles perpendicular to axis, forming pauci- or multispicular radial (non-plumose) tracts associated with very light, ascending spongin fibres but very few transverse uni- or aspicular connecting fibres; mesohyl matrix in extra-axial region moderately heavy, with few scattered auxiliary megascleres. Megascleres. Choanosomal principal subtylostyles coring axial fibres relatively short, slightly curved at centre, with smooth subtylote or sometimes evenly rounded bases and fusiform points. Length 215-(311.2)-395pm, width 5-( Subectosomal auxiliary styles forming extraaxial bundles much longer and thicker than principal spicules, with smooth rounded or tapering (hastate) bases, and fusiform points. Length 424(559.6)-725p.m, width 15-(21.0)-29p,m. Ectosomal auxiliary subtylostyles straight, variable length, with smooth rounded or slightly subtylote bases and fusiform points. Length 208(361.8)-575p.m, width 6-(8.2)-10iim. Microscleres. Palmate isochelae unmodified, differentiated into two size classes, with long lateral alae entirely fused to shaft and completely fused front ala. Length I: 6-(9.3)-13p.m; length ^19(22.3)-25p.m. Toxas oxhorn, thick, with wide central curvature and slightly reflexed points. Length 45(86.2)-130pm, width 2.5-(4.3)-6p.m. REMARKS. The chelae and toxa microscleres show that this species belongs to the Microcionidae, whereas skeletal architecture is most closely related to the Raspailiidae (cf Raspailia (Syringe/la) and Ectyoplasia). Like C. (A.) canaliculata, this species lacks echinating megascleres, and this was the primary reason why Hallmann (1920) created Axociella. Despite contrary arguments by Van Soest (1984b), Axociella is considered sufficiently different from Thalysias in skeletal construction to be differentiated at the supraspecific level (although not for 252zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 118. Clathria (Axociella) canaliculata (Whitelegge) (lectotype AMG4325). A, Choanosomal principal style. B, Subectosomal auxiliary style. C, Ectosomal auxiliary subtylostyle. D, Accolada toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. FI, Lectotype. I, QMG300460. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 253 FIG. 119. Clathria (Axociella) canaliculata (Whitelegge) (A, lectotype AMG4325; B-F, specimen QMG300460). A, Choanosomal skeleton. B, Extra-axial fibre characteristics (x144). C, Base of ectosomal auxiliary subtylostyle. D-E, Palmate isochelae. F, Accolada toxas. 254zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM A a FIG. 120. Clathria (Axociella) cylindrica (Ridley & Dendy) (AMZ1527). A, Choanosomal principal subtylostyle (coring axial fibres). B, Subectosomal auxiliary style (coring extra-axial skeleton). C, Ectosomal auxiliary subtylostyles. D, Oxhorn toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype BMNH1887.5.2.96. the reason that it lacks echinating acanthostyles; Simpson, 1968a). Supporting this interpretation is the fact that a specialised Thalysias ectosomal skeleton is absent in C. (A.) macropora and C. (A.) thetidis. AMZ1133 from the F1V 'Thetis' expedition, although not described by Hallmann (1912), is very close to C. (A.) cylindrica differing only in having echinating acanthostyles and long thin, almost straight rhaphidiform toxas. This material almost certainly represents a new species but its antiquated condition is too poor to serve as the holotype of a new taxon. Clathria (Axociella) cylindrica also shows some similarities to C. (Thalysias) procera in growth form and having well differentiated axial and extra-axial skeletons, but these species differ in most other respects. Axociella cylindrica of Sim & Byeon (1989) from Korea does not compare to the Australian population, differing markedly in most features, and is referred here to Clathria (Microciona) simae sp. nov. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 255 FIG. 121. Clathria (Axociella) cylindrica (Ridley & Dendy) (holotype BMNH1887.5.2.96). A, Choanosomal skeleton. B, Fibre characteristics. C, Oxhorn toxas. D-E, Palmate isochelae. F, Cross-section through branch of AMZ1527. MEMOIRS OF THE QUEENSLAND MUSEUM 256^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Axociella) nidificata (Kirkpatrick, 1907) (Figs 122-123) Ophlitaspongia nidificata Kirkpatrick, 1907: 274; Kirkpatrick, 1908b: 25, p1.22, fig.6, p1.24, fig.5; Hallmann, 1912: 254; Burton, 1929a: 433; Burton, 1932a: 324; de Laubenfels, 1936a: 120; de Laubenfels, 1954: 162. Axociella nidificata; Burton, 1940: 116; Koltun, 1964a: 70, p1.12,figs 7-11; Koltun, 1976: 155, 190; Desqueyroux, 1975: 67-68, p1.4, figs 47-50; Hoshino, 1977a: 45-46, table 1. Clathria (Axociella) nidificata; Hooper & Wiedenmayer, 1994:263; Mothes & Lerner, 1995: 159-160, figs 22-27, 55. Ophlitaspongia flabellata Topsent, 1916: 167; Topsent, 1917: 41, p1.1, fig.4, p1.6, fig.2; Burton, 1929: 433; Burton, 1932: 325; Burton, 1934b: 34. Axociella flabellata; Koltun, 1964a: 70-71, p1.12, figs 12-14; Hoshino, 1977a: 45-46, table 1. Axociella rameus Koltun,1964a: 71, text-fig.17; Hoshino, 1977a: 45-46, table 1. MATERIAL. HOLOTYPE: BMNH1908.2.5. 131: Coulman I., Victoria Land, Ross Dependency, Antarctica, 73°30'S, 17°00'E, 200m depth, coll. HMS 'Discovery' (dredge). HABITAT DISTRIBUTION. On hard substrates; 93540m depth; Antarctica: BANZARE coast; Budd Coast, Wilkes Land; Kemp Land, Enderby Land; Mawson Peninsula, Oates Land; Coulman I., Victoria Land; Prydz Bay, MacKenzie Bay and Mawson Coast, MacRobertson Land; King Edward Ice Shelf, Enderby Land; Joinville and Elefante Is; Palmer Archipelago; Graham Land; also South Georgia, S. Shetland Is (Fig. 122E). DESCRIPTION. Shape. Erect, claviform-flabellate, narrow stalk. Colour Yellow-brown in ethanol. Oscules. Oscules on upper surface of club, I 2mm diameter. Texture and surface characteristics. Texture compressible, tough, harsh surface; surface hispid, conulose, membraneous between conules. Ectosome and subectosome. Ectosomal skeleton plumose, composed of discrete bundles of subectosomal auxiliary subtylostyles perched on ends of protruding principal spicules, corresponding to surface conules. Choanosome. Skeleton compressed in axis and plumose in extra-axis; axial region slightly compressed containing heavy spongin fibres cored by plumo-reticulate pauci- or multispicular tracts of choanosomal principal styles; skeletal tracts form oval meshes, up to 250p.m diameter; extra-axial tracts plumose, diverging towards periphery, with pauci- or multispicular tracts of choanosomal principal styles coring heavy spongin fibres; styles in peripheral regions of fibres often protrude from fibres at acute angles, but these are identical to principal megascleres and cannot be considered as true echinating spicules; mesohyl matrix moderately heavy, granular, containing abundant toxas, often in bundles, and occasional detritus. Megascleres. Choanosomal principal styles long, thick, entirely smooth, slightly curved towards base, with slightly subtylote evenly rounded or slightly constricted hastate bases and fusiform points. Length 595-(914.7)-1140p-m, width 21(43.7)-61p.m. Subectosomal auxiliary subtylostyles short, slender, straight, with slightly subtylote microspined bases, fusiform pointed. Length 331-(405.4)-447p.m, width 6-(9.4)-12p.m. No echinating spicules. Microscleres. Chelae absent. Toxas accolada, thick, variable in length, with prominent but narrow angular central curvature, straight arms and straight or slightly reflexed tips; larger toxas with spined tips, small toxas entirely smooth. Length 98-(286.2)-546p.m, width 1.5(3.2)-5p.m. REMARKS. The synonymy given above follows Koltun (1976). The species appears to vary in growth form, from bushy, branching, reticulate, digitate to flabellate, but skeletal architecture and spicule geometry are relatively consistent. Several echinating acanthostyles were seen in spicule preparations made from the holotype (evenly spined, club-shaped, 94-123x5-9Rm), but none were seen in section preparations and it is presumed that these are contaminants, although it is possible that they are present but rare. The specialised ectosomal skeleton of discrete auxiliary spicule brushes perched over larger principal spicules, and compressed `axinellid' skeletal construction, are reminiscent of Raspailiidae (e.g., Aulospongus), and were it not for the toxas in this species it could be included in Raspailiidae. This species fits best in Clathria (Axociella) although it lacks differentiated ectosomal and subectosomal spicules (i.e., it has 1 size class of auxiliary styles). Its toxa and spination should be compared with C. (A.) georgiaensis sp. nov. (see below). Clathria (Axociella) patula sp. nov. (Figs 124-125) MATERIAL. HOLOTYPE: NTMZ2909: Between North and East Wallabi Is, Houtman Abrolhos, WA, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 257 FIG. 122. Clathria (Axociella) nidificata (Kirkpatrick) (holotype BMNH1908.2.5.131). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Accolada toxas. D, Section through peripheral skeleton. E, Antarctic distribution. 28°17.9'S, 113°47.8'E, 11.vii.1987, 39m depth, coll. J.N.A. Hooper (beam trawl). PARATYPE: NTMZ2978 (fragment QMG300209): W. of Carnarvon, WA, 24 ° 55.6'S, 112°50.8'E, 14.vii.1987, 85m depth, coll. J.N.A. Hooper (beam trawl). OTHER MATERIAL. WA- QMG304633. HABITAT DISTRIBUTION. Broken limestone reef, usually in sand sediments; 39-85m depth; Camarvon and Wallabi Is, Houtman Abrolhos (WA) (Fig. 124G). DESCRIPTION. Shape. Thickly flabellate, palmate-digitate fan, 140-475mm long, 180-310mm ^ MEMOIRS OF THE QUEENSLAND MUSEUM 258zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Ectosome and subectosome. Thick plumose or paratangential layer of erect auxiliary styles, usually in thick bundles, forming more-or-less continuous palisade on ectosomal skeleton; subectosomal skeleton with plumose brushes of both subectosomal auxiliary styles and choanosomal principal styles supporting ectosomal palisade, arising directly from choanosomal fibres in peripheral skeleton. Choanosome. Skeletal structure wide-meshed heavily reticulate, slightly compressed axial region and differentiated axial and extra-axial skeletons; axial skeleton composed of heavy spongin fibres, 80145p,m diameter, thickest and bulbous at fibre nodes, 130200p,rn diameter, forming relatively wide reticulation of oval or rectangular meshes, 180360Rm diameter; axial fibres cored by both principal styles and subectosomal styles in multispicular brushes of 2-5 spicules; principal spicules also protrude through fibres in FIG. 123. Clathria (Axociella) nidificata (Kirkpatrick) (holotype plumose bundles, at acute BMNH1908.2.5.131). A, Plumose extra-axial skeletal column. B, Plumo- angles, although these cannot reticulate axial skeleton. C, Bundles of toxas. be considered as echinating spicules; extra-axial skeleton with wider fibre meshes than wide, 6-15mm thick, with even apical margin or axial region, with heavy spongin fibres, 60long, erect flattened digits arising from apex of 12011m diameter, bulbous fibre nodes, 80-140p.m fan; digits single or planar branching, 40-70mm diameter, forming elongate or rectangular wide, bifurcated and tapering, with short, thick cylindrical stalk, 35-63mm long, 15-32mm meshes, 300-470Rm diameter, cored by ascenddiameter; lateral margins of fans prominently ing multispicular tracts of intermingled crenellated, apical margins range from slightly choanosomal principal and subectosomal styles, up to 10 spicules per fibre, and thinner transverse crenellated to prominently digitate. fibres cored by only principal spicules in Colour Dark red, red-brown or orange-brown paucispicular tracts; spicules usually protrude alive (Munsell 5R 5/10 - 1OR 7/8), grey-brown in through fibres in plumose bundles, particularly in ethanol. peripheral skeleton, but true echinating spicules Oscules. Small oscules, less than 2mm diameter, absent; mesohyl matrix heavy but only lightly scattered over surface. pigmented, containing abundant microscleres; Texture and surface characteristics. Firm, com- choanocyte chambers large, oval, 50-1301.Lm pressible, flexible; surface uneven, rippled, diameter. slightly conulose, with few large digitate projec- Megascleres. Choanosomal principal styles varitions arising at oblique angles from surface in fan able in length, predominantly long, thick, slightly specimens. curved near basal end, evenly rounded smooth ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 259 FIG. 124. Clathria (Axociella) patula sp.nov. (holotype NTMZ2909). A, Choanosomal principal style. B, Subectosomal auxiliary styles. C, Ectosomal auxiliary styles. D, Wing-shaped - oxhorn toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Known Australian distribution. H, Holotype. I, Paratype. MEMOIRS OF THE QUEENSLAND MUSEUM 260^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 125. Clathria (Axociella) panda sp.nov. (holotype NTMZ2978). A, Choanosomal skeleton. B, Fibre characteristics. C-E, Bases of choanosomal principal, subectosomal auxiliary and ectosomal auxiliary spicules. F, Palmate isochelae. G, Oxhorn - wing-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 261 bases and fusiform points. Length 2254383.6)370p,m, width 10413.2)-1*m. Subectosomal auxiliary styles long, thinner than principal spicules, straight, with rounded microspined bases, slightly subtylote spined bases, or subtylote smooth bases, and fusiform points. Length 1934265.2)-30311m, width 5(7.1)-8p.m. Ectosomal auxiliary styles short, straight or slightly curved near base, relatively thick, with rounded smooth or microspined bases and fusiform points. Length 104-(147.2)-185Rm, width 1.5-(3.8)-6Rm. Echinating spicules absent. Microscleres. Palmate isochelae small, unmodified, with lateral and front alae of approximately equal length; lateral alae fused completely to shaft, front ala entire. Length 14(15.8)-19pm. Toxas basically oxhorn, with some intermediate wing-shaped, short, moderately thick, rounded or slightly angular central curvature and slightly reflexed points 57-(69.6)-84Rm, width 141.9)-2.511m. Woolongong, NSW, 34 ° 30'S, 151 ° 12'E, 98-138m depth, coll. FIV 'Thetis' (dredge).0THER MATERIAL: NSW- QMG303752. S. AUSTAMZ4624. HABITAT DISTRIBUTION. Deeper offshore reefs; 50-142m depth; Sydney and Woolongong (NSW), SW. Great Australian Bight (SA) (Fig. 126H). DESCRIPTION. Shape. Erect, arborescent or digitate, whip-like growth forms, 190-480mm long, with long thin cylindrical stalk and thinly cylindrical branches, 15-25mm diameter; branching planar or in more than one plane, either irregularly bifurcate or pinnate, with evenly dichotomous bifurcations arising from larger, laterally disposed branches; apical margins of digits taper to fine points. Colour. Deep red alive (Munsell 2.5R 5/10), greybrown when dry. Oscules. Very small, less than 1.5mm diameter, scattered over lateral sides of branches. Texture and surface characteristics. Firm, flexible; surface optically even, velvetty, microscopically prominently furry, hispid. Ectosome and subectosome. Ectosome very ETYMOLOGY. Latin zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA pat ulus, spread out; for the flat hi spid, with discrete, multispicular plumose growth form. REMARKS. This species could be included in C. brushes of larger (extra-axial) principal styles (Thalysias), having a thick tangential ectosomal protruding through surface, with sparsely disskeleton composed of two discrete size classes of persed, tangential or paratangential tracts of subauxiliary styles, but it is most appropriately ectosomal auxiliary styles in between principal placed in C. (Axociella) given that it has differen- spicules. tiated axial and extra-axial skeletons and lacks Choanosome. Skeletal architecture reticulate, any echinating spicules. C. (Axociella)patula has with compressed axis and plumose extra-axial a flabellate, planar growth form similar to C. (T) regions; strong axial compression composed of cancellaria, although these species differ sub- irregularly reticulate, heavy, large spongin fibres stantially in their spicule geometry (the latter producing oval to elongate meshes, 90-220p.,m having echinating acanthostyles and accolada diameter; axial fibres relatively heavy, thick, irtoxas), different spicule sizes of most categories, regularly anastomosing, with bulbous fibre and skeletal structures (the latter species with a nodes, cored by comparatively small choanoheavy, evenly reticulate skeleton and abundant somal styles; axial fibres incompletely differentiated into primary, uni- or paucispicular, echinating acanthostyles). ascending, long, radial fibres, 70-1801.im Clathria (Axociella) thetidis (Hallmann, 1920) diameter, and secondary, connecting, mostly unispicular, sometimes bi- or aspicular, short (Figs 126-127, Plate 4B) fibres, 58-102p,m diameter. Axial and extra-axial skeletons strongly differentiated (cf. Hallmann, Ophlit aspongia t het idis Hal!mann, 1920: 779. 1920), with fibres becoming plumose and fibre Axociella t het idis; de Laubenfels, 1936a: 113. Clat hria t het idis; Hooper & Wiedenmayer, 1994: 263 reticulation more regular towards periphery; Esperiopsis cylindrica, in part; Whitelegge, 1906: 470, extra-axial fibres form elongate-eliptical, wider p1.43, fig.6. meshes than axis, 155-510Rm diameter; extraNot Esperiopsis cylindrica Ridley & Dendy, 1887: 79. axial fibres diminish in thickness towards peripheral skeleton, whereas coring spicules inMATERIAL. LECTOTYPE: AMG9199: Off Woolongong, NSW, 34 ° 25'S, 151 ° 10'E, 104-142m crease in size and density towards surface; principal styles in both primary and secondary fibres depth, coll. FIV 'Thetis' (dredge). PARALECprotrude through fibres at oblique angles, in both TOTYPE: AMG9191: Off Wata Mooli, Bulgo, ^ MEMOIRS OF THE QUEENSLAND MUSEUM 262zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 126. Clathria (Axociella) thetidis (Hal'mann) (holotype AMG9199). A, Choanosomal principal style (axial skeleton). B, Choanosomal principal style (extra-axial skeleton). C, Subectosomal auxiliary subtylostyle. D, Oxeote toxa. E, Wing-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. the axial and extra-axial regions, but true echinat- 11-(14.2)-20p,m), and larger in extra-axial ing spicules absent; mesohyl matrix heavy, rela- skeleton (length 377-(608.0)-825iim, width 22tively darkly pigmented; choanocyte chambers (26.4)-35p.m. paired, oval-elongate, 60-1541,m diameter; Subectosomal auxiliary styles long, thin, numerous microscleres and subectosomal fusiform, straight, slightly curved or flexuous, with subtylote, smooth or lightly rnicrospined auxiliary subtylostyles scattered between fibres.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGF bases. Length 158-(379.1)-585p,m, width 3Megascleres. Choanosomal principal styles divided into two size categories, both fusiform, (5.8)-811m. thick, long or short, slightly curved, with evenly Microscleres. Palmate isochelae abundant, with rounded, smooth bases; smaller size found in lateral alae completely attached to shaft and front axial skeleton (length 1804246.5)-365 pm, width ala entirely fused. Length 9413.4)-1611m. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 263 FIG. 127. Clathria (Axociella) thetidis (Hallmann) (QMG303752). A, Choanosomal skeleton. B, Extra-axial fibre and spicule skeleton (x63). C, Palmate isochelae. D, Wing-shaped toxas. E, Oxeote toxas. 264zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Toxas divided into two geometrically different forms: oxeote toxas common, usually found in dragmata dispersed between fibres, long, thin, symmetrical or asymmetrical-sinuous, with slight angular central curvature or completely straight, fusiform points. Length 175-(774.5)1280p,m, width 1.542.4)-5 p.m; wing-shaped toxas common, thin, with large rounded central curvature, reflexed points. Length 224104.4)168p.,m, width 142.1)-3.41,m. REMARKS. This species is not conspecific with Esperiopsis cylindrica Ridley & Dendy as supposed by Whitelegge (1906), having different ectosomal skeletal architecture, spicule geometries and spicule sizes; Hallmann (1920) recognised a new species for Whitelegge's material referring it to Ophlitaspongia (= Echinoclathria) on the basis that it lacked echinating megascleres; de Laubenfels (1936a) assigned it to Axociella for the same reason. De Laubenfels was correct in this transfer, although not for his stated reason (that it lacked acanthostyles) but because of its compressed skeletal construction, spicule localisation and spicule geometry. Hallmann's tentative placement of C. (A.) thetidis in Echinoclathria was probably also based on comparisons with species such as E. nodosa (Carter) and to a lesser extent E. subhispida (Carter) which, unlike most species of Echinoclathria, have a slightly compressed axial skeleton and plumose, plumo-reticulate or radial extra-axial fibres. However, spiculation and spicule localisation within the skeleton of C. (A.) thetidis is different from all those species. Similarly, in C. (A.) thetidis there are two distinctive size categories of principal megascleres, restricted to either axial or extra-axial fibre skeletons, whereas Echinoclathria have homogeneous principal spicules dispersed throughout all skeletal tracts. Clathria (Axociella) georgiaensis sp.nov. (Figs 128-129) Ophlitaspongia thielei Burton, 1932a: 322, p1.55, fig.8, text-fig.32; Koltun, 1964a: 70. Axociella thielei; de Laubenfels, 1936a: 113 [note]. Not Hymeraphia thielei Hentschel, 1912: 377-378. MATERIAL. HOLOTYPE: BMNH1928.2.15. 219 (fragment AMZ2198): 6.3nm N 89 ° E to 4nm N 39 ° E off Jason Light, Cumberland Bay, South Georgia, S. Atlantic, 120-204m depth, RRS 'Discovery', 1928 (otter trawl). HABITAT DISTRIBUTION. On rocks; 18-236m depth range; Wilkes Land, Australian Antarctic Territory (Fig. 128F); also South Georgia, SW. Atlantic. DESCRIPTION. Shape. Subspherical, massive sponge. Colour Grey-brown in ethanol. Oscules. Up to 2mm diameter, scattered over surface, with slightly raised membraneous lip. Texture and surface characteristics. Firm, compressible; surface conulose, with meandering ridges producing a clathrous, convoluted mass. Ectosome and subectosome. Erect plumose brushes of sparse choanosomal principal styles protruding from peripheral skeletal tracts, and paratangential bundles of subectosomal auxiliary subtylostyles in variable abundance, heavier on ends of surface conules, lighter between conules. Choanosome. Skeletal architecture reticulate, slightly plumo-reticulate near surface, vaguely renieroid reticulate at core; skeleton with differentiated primary ascending spongin fibres cored by multispicular tracts of choanosomal principal styles, with 3-6 spicules per tract, and lighter transverse connecting fibres containing 1-2 spicules per tract, together producing a slightly renieroid skeleton; no marked differentiation between axial or extra-axial regions; fibre reticulation produces cavernous rectangular or triangular meshes, up to 550p..m diameter; true echinating spicules absent although principal spicules protrude through fibres at obtuse angles; mesohyl matrix heavy, slightly granular, with abundant chelae and toxas. Megascleres. Choanosomal principal styles moderately long, slender, straight or slightly curved at centre, rounded smooth bases, fusiform points. Length 3904446.6)-51811m, width 14(17.4)-2211m. Subectosomal auxiliary subtylostyles short, slender, straight, slightly subtylote bases, heavily microspined with large spines, slightly hastate or rounded points. Length 216-(285.2)-348itm, width 5-(7.4)-10Rm. Echinating spicules absent. Microscleres. Palmate isochelae abundant, relatively small, with lateral and front alae approximately same length, lateral alae fused to shaft, front alae nearly completely detached, virtually no curvature of shaft. Length 9-(l3.2)17 p.m. Toxas wing-shaped, thick, variable in length, the thicker ones with wide angular central curvature, curved arms, slightly reflexed points, and terminal spines, thinner ones more sharply curved ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 265 FIG. 128. Clathria (Axociella) georgiensis sp. nov. (holotype BMNH1928.2.15.219). A, Choanosomal principal styles. B, Subectosomal auxiliary subty losty les. C, Wing-shaped toxas. D, Palmate isochelae. E, Section through peripheral skeleton. F, Antarctic distribution. G, Peripheral spicule tracts. H, Ectosomal skeleton. at centre, sharply pointed. Length 284107.7)258p.m, width 0.541.7)-3.511m. REMARKS. This species requires a new name since C. (Microciona) thielei (Hentschel, 1912) has seniority. Previous authors could not agree on its generic assignment: Burton (1932a, 1938b) compared C. (A.) georgiaensis with Artemisina, but the possession of a well-structured, slightly compressed choanosomal skeleton indicates that 266zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 129. Clathria (Axociella) georgiaensis sp. nov. (AMZ2198). A, Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal principal style. D, Base and apex of principal spicule. E, Base and apex of subectosomal auxiliary subtylostyle. F, Palmate isochela. G, Wing-shaped toxas. H, Spined toxa point. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 267 placement in Clathria is more appropriate. Based on Burton's (1932a) description this species was included in Echinoclathria by Hooper & Wiedenmayer (1994), but re-examination of the holotype showed that skeletal structure was not markedly renieroid, there was no difference between principal spicule sizes in the axis and those in the peripheral skeletons, and spicule sizes were magnitudes larger than those found in other Ecinoclathria (but much closer to those seen in Axociella). De Laubenfels (1936a) also suggested this species should be included in Axociella although his reasons for doing so are different (i.e., lack of echinating spicules). Koltun (1964a) also made a general comparison between C. (A.) georgiaensis and Ophlitaspongia membranacea Thiele (the latter referred here to Clathria (Thalysias)), but this comparison is simply based on similarities in toxa morphology, whereas C. (T) membranacea has two size classes of auxiliary spicules and true echinating spicules (albiet smooth). Toxa measurements recorded by Burton (1932a) for this species differ substantially from those actually seen in the holotype, but in most other characters his description is an accurate representation of the species. OTHER SPECIES OF CLATHRIA (AXOCIELLA) Clathria (Axociella) fauroti (Topsent, 1893) Axosuberites fauroti Topsent, 1893a: 179-181, fig.3 [Tadjoura. Gulf of Aden]; de Laubenfels, 1936a: 118. ? Rhaphidophlus fauroti; Van Soest, 1984b: 130 [possible generic synonymy]. MATERIAL. HOLOTYPE: MNHNDT1859. Arabian GulfRed Sea. Clathria (Axociella) fromontae sp. nov. Axociella toxitenuis Bergquist & Fromont, 1988, 117-118, p1.55, figs a-c; Dawson, 1993: 36 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPOR120. New Zealand. Note: C. toxitenuis Topsent, 1925 has seniority. Clathria (Axociella) lambei (Koltun, 1955) Microciona lambei Koltun, 1955a: 49, 67, p1.4, fig.5 [Japan and Okhotsk Seas]; Koltun, 1958: 66-67, text-fig.22 [Kuriles]; Koltun, 1959: 183, p1.29, fig.2, text-fig.143 [USSR]. MATERIAL. HOLOTYPE: ZIL, fragment BMNH1932.11.17.67. NW. Pacific, Japan. Note: Koltun (1955a) attributes this species to Burton (1935c), but it does not appear in that publication. Clathria (Axociella) macrotoxa Bergquist & Clathria (Axociella) multitoxaformis Bergquist & Fromont, 1988 Axociella multitoxaformis Bergquist & Fromont, 1988: 118119, p1.55, figs d-f, p1.56. fig.a; Dawson, 1993: 36 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPOR121. New Zealand. Clathria (Axociella) parva (Levi, 1963) Clathria parva Levi, 1963: 56-57, text-fig. 64, p1.10D [Cape of Good Hope, South Africa]; Uriz, 1988a: 84-85, p1.21b, text-fig.60 [Namibia]. Not Clathria parva; Sim & Byeon, 1989: 39, pI4, figs 3-4 [Korea; dubious conspecificity]. MATERIAL. HOLOTYPE: UCT (fragment MNHNDCL612). S and SE Africa. Clathria (Isociella) Hallmann, 1920 Isociella Hallmann, 1920: 784; Bergquist & Fromont, 1988: 114. DEFINITION. Relatively homogeneous isodictyal (triangular meshes) and/or renieroid (rectangular meshes), wide-meshed, main skeleton with primary plumose ascending, multispicular tracts cored by smooth choanosomal styles, interconnected by secondary, uni- or paucispicular tracts cored by ,same spicules, and choanosomal spicules sometimes diverging and forming plumose brushes at surface; echinating megascleres absent; ectosomal skeleton with single category of auxiliary spicule, tangential, paratangential. Microscleres palmate-derived isochelae and toxas. TYPE SPECIES. Clathria macropora, Lendenfeld, 1888: 221 (=Phakelliajacksoniana Dendy, 1897:236) (by monotypy). REMARKS. Four species of Isociella are known for the Australian fauna, three from tropical WA, NT and Qld., and one temperate species from NSW. Only one other species is known from New Zealand. Clathria (Isociella) eccentrica (Burton, 1934) (Figs 130-131, Table 26, Plate 4A) Ophlitaspongia eccentrica Burton, 1934a: 560, pls 1,8,9, text-fig.12a. Axociella eccentrica; de Laubenfels, 1936a: 113. Isociella eccentrica; Bergquist & Tizard, 1967: 186187, p1.5, fig. 1. Clathria eccentrica; Hooper & Wiedenmayer, 1994: 265. Fromont, 1988 MATERIAL. HOLOTYPE: BMNH1930.8.13. 109: Axociella macrotoxa Bergquist & Fromont, 1988: 117, p1.54, figs c-f; Dawson, 1993: 36 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPOR119. New Zealand. Crab Spit, Low Isles, Great Barrier Reef, Qld, 16 °23'S, 145°34'E, intertidal, 5.iv.1929, coll. Great Barrier Reef Expedition (dredge). OTHER MATERIAL: GREAT ^ MEMOIRS OF THE QUEENSLAND MUSEUM 268zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A 0 FIG. 130. Clathria (Isociella) eccentrica (Burton) (NTMZ2170). A, Choanosomal principal styles. B-C, Subectosomal auxiliary styles. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype BMNH1930.8.13.109. I, NTMZ2139. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 269 FIG. 131. Clathria (Isociella) eccentrica (Burton) (QMG303266). A, Choanosomal skeleton. B, Fibre characteristics (x135). C-D, Bases of larger and smaller subectosomal auxiliary styles. E, Wing-shaped toxas. F, Palmate isochelae. 270zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 26. Comparison between present and published descriptions of Texture and surface characClaihria (Isociella) eccentrica.Measurements in p.m (N=25). teristics. Rubbery, easily com- pressible, mucusy, but tough, difficult to tear; surface SPICULE porous, membraneous, (GBR,Queensland) minutely microconulose with 2324361.7)-540 276-(343.1)-450 Choanosomal 318-464 x 8-22 288-440 x 9.1-18 protruding fibre nodes from styles x7-(17.4)-30 x 11-(19.2)-29 peripheral skeleton, with 2574335.6)-440 284-(373.3)-455 Subectosomal 298-457 x 3-9.5 300-470 x 5-8 small ridges and depressions styles I x 3-(6.8)-13 x 4-(6.3)-9 connecting adjacent conules; 141-(191.8)-255 85-(185.0)-255 Subectosomal 85-286 x 1.5-5.2 208-220 x 4.0 surface usually silt covered. styles II x 1-(3.5)-7 x 2-(3.7)-6 Ectosome and subectosome. 15-19 16-17.5 8-(17.4)-23 13-(17.2)-21 Chelae Ectosomal skeleton micros64-(161.2)-292 16-(148.0)-283 10-334 x 1.5-10 76-270x 2.5-6.5 Toxas copically hispid, with x 1-(5.4)-11 x 2-(6.0)-11 choanosomal principal styles protruding up to 300p,m from BARRIER REEF, QLD - QMG304398, QMG304400, surface, at regular intervals 150-400p.m apart, QMG304401. DARWIN REGION, NT - AMZ3109, NTMZ0268, NTMZ0288, NTMZ0158, NTMZ0159, singly or in paucispicular bundles of 2-4 spicules, with peripheral spongin fibres forming projecNTMZ2116, NTMZ2139, NTMZ2205, NTMZ2210, QMG300147 (fragment NTMZ2224), QMG300509 tions and enclosing at least basal portion of (fragment NTMZ2235), NTMZ2416, NTMZ2540, protruding spicules; subectosomal auxiliary QMG303315, NTMZ2549, NTMZ2557, NTMZ1100, styles form tangential or paratangential tracts at NTMZ0386, NTMZ2170, QMG303266, base of peripheral skeleton, interdispersed beNTMZ0045, NTMZ1396, NTMZ3274. AMG4291. tween choanosomal principal styles of subrenieroid skeleton; auxiliary spicules sparsely HABITAT DISTRIBUTION. Usually on dead or pardispersed, rarely protruding through surface, tially dead faviid coral heads, coral rubble, sand and composed of 2 sizes of auxiliary styles without Halimeda substrates; most specimens partially sheltered under coral rubble or in crevices; shallow sublit- regional localisation; peripheral skeleton undiftoral distribution; 0-18m depth range; Lizard I. (FNQ), ferentiated from choanosomal skeleton, and Darwin Harbour, Bynoe Harbour, Trepang Bay, Port choanosomal fibres immediately subdermal. Essington, Cobourg Peninsula (NT) (Fig. 1306); also Choanosome. Irregular subisodictyal, or in places SE. Indonesia (unpublished data). more regular isodictyal, sometimes renieroid reticulation of moderately light spongin fibres, DESCRIPTION. Shape. Bulbous-digitate, mas- 30-110v1,m diameter, without any obvious or consive or semi-encrusting, with irregularly anas- sistent differentiation between primary or secontomosing, erect or stoloniferous branches; dary elements; fibres cored by 1-10 rows of branches irregularly cylindrical, flattened or bul- choanosomal principal styles entirely enclosed bous; specimens range from thick encrustations within fibres, occupying entire fibre diameter; 45mm high, 20-30mm diameter, with few bul- spicules diverge only slightly towards periphery bous branches on surface, to massive branching whereas at core of skeleton spicules more evenly growth forms up to 130mm high, 300mm wide, dispersed within fibres; echinating megascleres with branch diameter between 15-40mm. absent; fibre anastomoses form irregular, elonColour Dark red (Munsell 5R 3/8), orange-red gate-oval, triangular or sometimes regularly rec(5R 6/10), or less commonly bright orange (10R tangular meshes, 250-850Rm diameter; mesohyl 6/10); pigmentation water miscible, associated matrix light, granular, with abundant with mucous, confined to the ectosomal and sub- microscleres and auxiliary spicules; collagen ectosomal regions; pigment washed from sponge; heaviest at fibre nodes but sparse elswhere; subectosomal colouration light brown or beige; choanocyte chambers oval or elongate, 60150p,m diameter. grey-brown in ethanol. Megascleres (Table 26). Choanosomal principal Oscules. Large, 4-15mm in diameter, on apex of bulbous digits, usually at extremities of branches; styles straight or slightly curved near base, with oscules with slightly raised membraneous lips, smooth rounded or very slightly subtylote bases, collapsing upon dessication and preservation; fusiform points. numerous inhalant pores, 0.5-1.0mm diameter, Subectosomal auxiliary subtylostyles thin, scattered over entire surface. usually straight, either with slightly subtylote or Holotype (BMNH 1930.8.13.109) Bergquist & Tizard (1967) (Danvin,NT) (19=19) (Darwin,NT) (N=3) (Cobourg Pen.,NT) REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 271 rounded bases, and microspined or smooth bases in approximately equal proportions, and with fusiform points; 2 size categories recognised, although undifferentiated in distribution. Echinating spicules absent. Microscleres (Table 26). Palmate isochelae abundant, relatively large, unmodified, with long lateral alae completely fused to shaft and entirely fused front ala. Wing-shaped toxas variable length, relatively thick, with moderate, rounded central curvature, slightly reflexed or straight points; some oxeote toxas also present but rare; toxas frequently occur in dragmata within mesohyl. Associations. Sometimes smothering live faviid coral heads, produced localised bleaching and necrosis of coral tissues at the point of contact with sponge; probable that this species involved in chemical bioerosion of coralline substrate; most specimens (81%) contained scyllid polychaete worms. Morphological variation. In growth form, live colouration, surface features, skeletal construction and spiculate geometry this species shows very little apparent variation. Some variation observed for: Detritus incorporated into ectosomal skeleton: abundant (33%), lightly dispersed particles (24%), entirely clear of detritus (43% of specimens). Ectosomal skeleton: strictly tangential (43%), disorganised paratangential (33%), or with both structures on different parts of the surface (24% of specimens). Choanosomal skeleton: characteristically subisodictyal (82%), regularly isodictyal (9%), or regular renieroid construction (9% of specimens). Fibre meshes: cavernous (86%), or close-meshed skeletal reticulation (14% of specimens). Mesohyl matrix: relatively heavy, darkly pigmented (19%), heavy, lightly pigmented (57%), or very light, unpigmented (24% of specimens). Detritus incorporated into mesohyl: seen in only 14% of specimens. Abundance of microscleres: very abundant in tracts or irregularly dispersed throughout the mesohyl (67%), or uncommon (33% of specimens). Auxiliary styles dispersed between fibres within mesohyl: numerous (43%), moderate (19%) or sparse extra-fibre tracts (38% of specimens). Spicule dimensions: spicule dimensions varied considerably between specimens, but this variation was not explained by either geographic distribution or seasonality of collections. REMARKS. This species is a distinctive component of the tropical Australian intertidal fauna, although it is only known from two disjunct loca- tions (the 'Top End' of the NT and Cairns region, Qld.). No intermediate populations have yet been discovered despite major collections undertaken recently in that region. This species is well characterised by its haplosclerid-like, predominantly isodictyal skeletal construction, relatively poor development of the extra-fibre skeleton (including the cavernous fibre meshes and relatively light mesohyl matrix), and spicule geometries. The species is also distinctive in the field, mainly by its predominantly bright red colouration and abundant mucus. It has the ability to survive extensive periods of exposure to direct sunlight and air, for up to six hours duration (several times each week during ELWS tides in the Darwin region), and to water temperatures in rockpools which may exceed 40°C. Under such conditions C. (I.) eccentrica produces copious quantities of mucus, literally dripping its red pigmentation. Nevertheless, individuals appear to survive these harsh conditions, and necrotic areas of the surface are relatively quickly regenerated or recolonised within several weeks. Despite extensive monitoring of individuals in the Darwin region over many seasons, reproductive products have not yet been recorded, and it is possible that its propogation in the tropics is predominantly clonal. Clathria (Isociella) macropora Lendenfeld, 1886 (Figs 132-133, Table 27) Phakelhaflabellata Ridley & Dendy, 1886: 478; Rid- ley & Dendy, 1887: 171, p1.34, figs 2-3, p1.40, fig.6 [preocc.1. Not Phakelliaflabellata Carter, 1885f: 363. Clathria macropora, in part, Lendenfeld, 1888: 221; Hallmann, 1920: 768; Bergquist & Fromont, 1988: 110; Hooper & Wiedenmayer, 1994: 265. Not Plectispa macropora Lendenfeld, 1888: 226; Hallmann, 1912: 203, 205, 242. Not Clathria macropora; Whitelegge, 1901: 91. Not Wilsonellamacropora; Hallmann, 1912: 203, 240, 242, 205. Not Plumohalichondria australis Whitelegge, 1901: 90, p1.11, fig.14. Isociella flabellata; Hallmann, 1920: 784-789, p1.39, figs 1-2, p1.40, fig.1, text-fig.3. Phakelliajacksoniana Dendy, 1897: 236; Whitelegge, 1907: 507. Isociella jacksoniana; Bergquist & Tizard, 1967: 187. MATERIAL. HOLOTYPE: AMZ466: Port Stephens, NSW, 32° 42'S, 152 °06'E, no other details known (label 'Clathria macropora Lend., type'). LECTOTYPE of P. jacksoniana: BMNH1887.5.2.9: Port Jackson, NSW, 33 ° 51'S, 151 ° 16'E, 54-90m depth, coll. HMS 'Challenger' (dredge). PARALECTOTYPE of MEMOIRS OF THE QUEENSLAND MUSEUM 272^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA cD 0 FIG. 132. Clathria (Isociella)macropora Lendenfeld (holotype AMZ466). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Modified palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. G, Paralectotype of Phakellia jacksoniana BMNH1887.5.2.8. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 273 FIG. 133. Clathria (lsociella) macropora Lendenfeld (holotype AMZ466). A, Choanosomal skeleton. B, Semi-renieroid fibres. C, Bases of subectosomal auxiliary styles. D, Modified palmate isochelae. 274zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM P. jacksoniana: BMNH1887.5.2.8: same locality. OTHER MATERIAL: VIC - AMZ771. HABITAT DISTRIBUTION. Subtidal rock reef to deeper offshore reefs, in soft sediments; 12-90m depth known only from Australia: Port Stephens, Port Jackson (NSW), Port Phillip (Vic) (Fig. 132E). TABLE 27. Comparison between present and published records of Clathria (lsociella) macropora (Lendenfeld). Measurements in p,m, denoted as range (and mean) of spicule length x spicule width (N=25). . DESCRIPTION. Shape. Irregularly flabellatedigitate or flabellate, planar, 85-170nrun long, 55-105mm maximum breadth, with short cylindrical stalk 15-25mm long, 7-15mm diameter, one or more thinly lobate, bifurcated branches, up to 8mm thick, either free or fused to adjacent branches, with rounded, digitate, uneven or shaggy margins. Colour Yellow-grey or brown in ethanol. Oscules. Oscules small, 1-3mm diameter, in special areas (sieve-plates) scattered over surface of branches, with series of stellate subdermal drainage canals surrounding each osculum. Texture and surface characteristics. Firm, flexible; branches with separate inhalant and exhalant faces; one surface porous, rugose, with irregular longitudinal ridges, microconules or irregular striations; other surface relatively smooth, membraneous. Ectosome and subectosome. Ectosome membraneous, hispid, with points of choanosomal principal styles protruding through surface, individually or in multispicular plumose bundles; surface skeleton with relatively sparse tangential, paratangential or sometimes plumose erect skeleton of small subectosomal auxiliary styles projecting between principal spicules, sometimes surrounding (in proximity to) principal spicules reminiscent of Raspailiidae. Choanosome. Choanosomal skeleton with very slightly compressed axis and plumose sub-isodictyal, sometimes renieroid extra-axis. Axial region with moderately heavy spongin fibres, forming tight irregularly reticulate meshes cored by paucispicular tracts of choanosomal principal styles; axis (corresponding to central lamellae and basal stalk) has few ascending, primary tracts, forming multispicular, halichondroid structures, 250-400p,m diameter, producing few multispicular, dendritic tracts running from basal stalk to periphery, 50-150Rm diameter; extraaxial skeleton sub-isodictyal with plumose spicule tracts bound by collagen (without fibre component), ascending to surface, cored by uni-, pauci- or less frequently multispicular tracts of choanosomal principal styles; primary ascending extra-axial spicule tracts (up to 5 spicules sideby-side) arise perpendicular to axis, intercon- Para1ectotype of SPICULE Choanosomal principal styles P. jacksoniana Holotype (AMZ466) AMZ77I (BMNH 3694446.4y 552x 21 - (26.6) - 35 1887.5.2.8) 441-(494.7)558 x 19 - (28.8) - 38 350(462.3)548 x 11 - (24.5) - 36 155(216.7) 384 x 2.5 (5.9)-9 Subectosomal 191 (287.0) 162 (197.1) auxiliary 424 x 4 - (6.4) - 9 226 x 4 - (6.7) -9 styles - Echinating acanthostyles - - absent absent Chelae 8-(12.9)-16 9412.0)-14 Toxas absent absent - - - absent 9-(12.5)-15.5 — absent nected by more or less transversely orientated, - - smaller secondary uni- or paucispicular tracts producing predominantly subisodictyal structure; peripheral spicule tracts more plumose than deeper choanosomal tracts; meshes produced by spicule-fibre anastomoses in extra-axial region triangular or rectangular in shape, 280-560Rm diameter; echinating acanthostyles absent; mesohyl matrix contains abundant relatively heavily pigmented spongin, with moderate quantities of auxiliary spicules, and choanocyte chambers oval, 50-19811m diameter. Megascleres (Table 27). Choanosomal principal styles long or short, thick, slightly curved at centre, less often straight, with rounded or slightly tapering, smooth bases, varying from fusiform to hastate points. Subectosomal auxiliary styles variable in length, thin, straight or very slightly curved, with basal terminations varying from evenly rounded, tapering hastate, quasi-diactinal mucronate or slightly subtylote, and with hastate points. Echinating spicules absent. Microscleres (Table 27). Palmate isochelae with highly modified, relatively small alae bearing wing-shaped fluted processes; lateral alae entirely fused to shaft; front ala complete or bifurcated with medial tooth; chelae frequently twisted or occasionally anisochelate. Toxas absent. REMARKS. The synonymy of C. macropora and P. jacksoniana is obvious on type material, but virtually impossible to tell from published descriptions, so the synonymy was overlooked by Hooper & Wiedenmayer (1994). In nearly all REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 275 features type material of both species is identical although auxiliary spicules in macropora are relatively larger. The name macropora is the senior name. Ridley & Dendy's (1886) flabellata is a junior homonym of Carter's (18850 species, and Dendy's (1897) replacement jacksoniana now considered to be redundant. Unfortunately macropora requires further clarification given that it has been misused and misinterpreted. My interpretation of macropora is based solely on the holotype because despite Hallmann's (1912: 166) arguments in defense of Lendenfeld's systematics, there is no doubt that Lendenfeld was dealing mainly with specimens which did not belong to the Microcionidae. Clathria macropora is nothing like Plectispa macropora (sensu Lendenfeld, 1888; holotype AMG9159), which has smooth echinating spicules, identical to those coring fibres, and is referred here to Holopsamma. Clathria macropora Lendenfeld (1888) (AMZ959) from Port Jackson, NSW and C. macropora from Port Chalmers, Qld (BMNH1950.2.12.60) are specimens of Crella incrustans (Crellidae). Three other specimens in the AM collections bearing the name macropora': one specimen from Nelson Bay, New Zealand collected by Arthur Dendy (AM unreg.) is a Crella with an ectosomal crust similar to C. levis var. digitata (AMZ454) (= C. incrustans); another (AMZ4035) collected from Dee Why, Sydney (RRIMPFN1338) and the third (AMZ4187 (RRIMPFN1428)) from 'Tumbledown', Jibbon Head, NSW, both from the Roche Collection, are haplosclerid sponges with three dimensional ectosomal skeletons, probably related to Amphimedon (Niphatidae). This taxon was not among several hundreds of specimens from subtidal and deeper water in Port Jackson, Port Stephens and the adjacent coastline (collected by the NSW Environmental Protection Authority and Sydney Water Board benthic monitoring surveys). It is also doubtful that it occurs in New Zealand, as Bergquist & Fromont (1988) questioned Lendenfeld's (1888) record given that it was not subsequently rediscovered during their substantial contemporary collections of the NZ fauna. This is confirmed here from re-examination of Lendenfeld's voucher specimen from Nelson Bay, NZ (mentioned above), which belongs to Crella. Clathria (Isociella) macropora has distinctive fluting on the teeth isochelae. This feature is barely visible under light microscopy, and therefore it is possible that it may also occur in other species of Clathria, which have not yet been studied using SEM, but apparently it is unique to the family. The species also has a slightly compressed axial skeleton partially offset from the diverging, plumose, subisodictyal reticulate extra-axial skeleton, showing vague structural similarities to Ceratopsion and Raspailia (Syringella) (Raspailiidae). This skeletal structure could also justify its inclusion in C. (Axociella), but it is considered here that the subisodictyal reticulation dominates the skeleton and is more characteristic of lsociella than Axociella. Clathria (Isociella) selachia sp. nov. (Figs 134-135, Plate 3F) MATERIAL. HOLOTYPE: NTMZ2946: E. side of Steep Point Lighthouse, South Passage, Dirk Hartog I., Shark Bay, WA, 26°08.5'S, 113 ° 10.3'E, 13.vii.1987, 7m depth, coll. J.N.A. Hooper (SCUBA). PARATYPE: QMG300562: same data. HABITAT DISTRIBUTION. Encrusting on excavated limestone plates, in surge zone; 7m depth; central W coast (WA) (Fig. 134E). DESCRIPTION. Shape. Massively encrusting, bulbous lobate-digitate, holotype 155mm wide, 105mm high, paratype 65mm wide, 60mm high, with thickly flabellate, slightly flattened lobes, up to 55mm thick, 110mm long, mostly fused to adjacent lobes, together forming a bulbous mass. Colour Bright red-orange alive (Munsell lOR 6/10), grey-brown in ethanol. Oscules. Large, up to 8mm diameter, clustered on tops of bulbous lobes or on margins of flattened lobes, slightly raised above surface with membraneous lip. Texture and surface characteristics. Firm, compressible, moderately difficult to tear; surface fleshy, bulbous, slightly microconulose, membraneous in situ, with membrane collapsing upon preservation producing fibrous, reticulate, porous, prominently conulose, shaggy surface. Ectosome and subectosome. Ectosome membraneous, slightly hispid from protruding choanosomal spicule tracts, with relatively thick paratangential or tangential skeleton of subectosomal auxiliary subtylostyles in multispicular tracts; ectosomal skeleton thickest at apex of surface microconules, perched on ends of ascending choanosomal skeletal tracts. Choanosome. Choanosomal skeleton plumoreticulate, subisodictyal and subrenieroid, with differentiated primary ascending and secondary transverse spongin fibres and spicule tracts; primary ascending fibres well developed, 65- MEMOIRS OF THE QUEENSLAND MUSEUM 276zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ A 0 c FIG. 134. Clathria (Isociella) selachia sp.nov. (holotype NTMZ2946). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Anchorate-like isochela. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. 1201J,m diameter, cored by multispicular tracts of larger choanosomal principal subtylostyles, 3-15 spicules per tracts; spicules not occupying entire fibre diameter; primary fibres bifurcate repeatedly, but anastomose only occasionally, producing prominent plumose structure most noticeable at periphery; principal subtylostyles mainly form axial core of spicules but sometimes they protrude through primary fibres producing plumose brushes; primary fibres interconnected at more-or-less regular intervals by well developed uni- or paucispicular spongin fibres, 20-451im diameter, cored by principal subtylostyles, forming triangular, rectangular or oval meshes, 90-180p.m diameter; skeleton slightly more cavernous near periphery, more plumose in structure; fibres heaviest in axis, more subisodictyal in structure; echinating megascleres absent; mesohyl matrix light, nearly unpigmented, with many smaller auxiliary subtylostyles and isochelae dispersed between fibres; generally spicules dispersed between fibres more slender, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 277 Microscleres. Isochelae anchorate -like, with slightly to greatly curved shaft, variable alae development from vestigial, tooth-like producing unguiferous chelae, to spatulate, fused forming lobate alae; lateral alae detached from shaft; front alae incompletely fused to adjacent ala; distal portions of shaft with lateral ridge. Length 26-(31.2)-34p.m. Toxas absent. ETYMOLOGY. Greek selachos, shark, for Shark Bay, WA. REMARKS. This species belongs to Clathria, based on its spiculation, choanosomal skeletal structure, ectosomal skeleton and fibre characteristics. It is assigned here to C. (Isociella) given its prominent subisodictyal skeleton, most obvious in the axial region, and lack of echinating spicules. It differs from other Isociella in having a predominently plumo-reticulate skeleton in the peripheral region (although subisodictyal in the axis) and in its spicule dimensions. When this species was first examined it was considered that the anchorate-like isochelae described above might be contaminants from another FIG. 135. Clathria (Isociella) selachia sp.nov. (holotype NTMZ2946). A, sponge or from the substrate upon which it grew. But Choanosomal skeleton. B, Fibre characteristics. C, Anchorate isochelae. numerous histological preparations made from the holotype, from various regions within the sinuous (probably juvenile) than in ectosomal sponge, repeatedly turned up these chelae. Furskeleton; choanocyte chambers large, oval, 220thermore, examination of chelae in situ found 27011m diameter. Megascleres. Choanosomal principal subtylos- them to be scattered both within the surface tyles long, thick, straight or slightly curved skeleton, intermingled with the paratangential towards base, with slightly subtylote or bundles of auxiliary spicules, and also surroundprominently subtylote bases, tapering fusiform or ing choanocyte chambers within the choanotelescoped points. Length 231-(260.6)-303Rm, somal mesohyl. They are native to this species, although this is at odds with the current hypothesis width 8-(11.6)-16vim. Subectosomal auxiliary subtylostyles relative- concerning the derivation and evolutionary sigly long, slender, usually straight, occasionally nificance of these spicules (Hajdu et al., 1994). The holotype and paratype were found growing sinuous, with prominently subtylote bases, tapering fusiform points. Length 125-(213.9)-294Rm, side by side and are probably clones of the same individual. width 1-(3.4)-6Rm. MEMOIRS OF THE QUEENSLAND MUSEUM 278zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 136. Clathria (lsociella) skia sp.nov. (holotype QMG300449). A, Auxiliary subtylostyle (coring fibres). B, Ectosomal auxiliary subtylostyle. C, Palmate isochela. D, Raphidiform toxas. E, Section through peripheral skeleton. F, Known Australian distribution. G, Holotype. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 279 35mm diameter, with irregularly and closely anastomosing digits, rounded and irregular margins, branches 8-18rrim diameter. Colour. Live colouration unknown, dark brown in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, compressible; surface macroscopically even, microscopically conulose, hispid, with subectosomal striations. Ectosome and subectosome. Ectosome with specialised skeleton composed of smaller auxiliary subtylostyles, forming sparse multispicular bundles on surface, arising directly from heavy, darkly pigmented peripheral fibres; tips of primary spongin fibres in peripheral region protrude through surface, producing surface microconules up to 300pLm long; majority of peripheral fibres nearly tangential, forming irregular reticulate meshes, cored by multispicular tracts of larger auxiliary spicules (6-10 spicules per tract). Choanosome. Skeletal architecture is irregularly isodictyal and renieroid reticulate, with clearly differentiated primary, ascending fibres, 60-125Rm diameter, cored by pauci- or multispicular fibres of larger auxiliary subtylostyles, up FIG. 137. Clathria (Isociella) skia sp.nov. (holotype QMG300449). A, to 8 spicules abreast, interconEctosomal skeleton. B, Fibre characteristics. C, Skeletal structure. D, nected by secondary, transverse, Palmate isochela. E, Raphidiform toxas. regular or irregular, uni- or paucispicular fibres (22-48p.m Clathria (Isociella) skia sp. nov. diameter); spongin fibres heavy; echinating (Figs 136-137, Table 28) spicules absent, although coring megascleres may protrude through fibres at oblique angles, MATERIAL. HOLOTYPE: QMG300449 (fragment becoming more plumose towards periphery; fibre NTMZ1522): W. of Sudbury Reef, Cairns region, anastomoses produce triangular, rectangular or Great Barrier Reef, Qld, 17 °03'S, 146 °07.8'E, 33-36m depth, 28.i.1981, coll. A. Kay (trawl). PARATYPE: oval meshes, 190-425Rm in diameter, becoming AMG5043: Masthead I. lagoon, Capricorn-Bunker more regular near periphery; mesohyl matrix Group, Great Barrier Reef, Qld, 23 ° 32'S, 151 °43'E, heavy, darkly pigmented, and many scattered auxiliary subtylostyles dispersed between fibres; 40m depth, no other details known. choanocyte chambers small, oval, 40-85 p.m HABITAT DISTRIBUTION. Attached to shell frag- diameter. ments or coral rubble, in soft sediments; 33-40m depth; Me gascleres (Table 28). Principal spicules abCairns region (NEQ), Gladstone region (MEQ) (Fig. sent. 136F). Auxiliary subtylostyles (coring fibres) long, DESCRIPTION. Shape. Erect, bulbous-lobate slender, invariably straight, with prominent, digitate, clathrous sponge, 45-95mm long, 18- smooth subtylote bases and fusiform points. 280zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Ectosomal auxiliary subtylostyles significantly smaller than coring spicules but otherwise of identical geometry. Echinating spicules absent. Microscleres (Table 28). Palmate isochelae uncommon, unmodified, with long lateral alae entirely fused to shaft and front ala completely fused, slightly spatulate. Toxas raphidiform, extremely thin, slightly curved at centre, sometimes nearly straight or bow shaped, rarely asymmetrical. ETYMOLOGY. Greek skias , dark, dim. REMARKS. This species is a greatly reduced Clathria, lacking true principal spicules, echinating spicules and having containing relatively sparse microscleres. Conversely its spongin fibre system is well developed and fibres are heavy. It is assigned to the Isociella group on the basis that its skeletal structure is essentially subisodictyal, and it lacks echinating spicules, but it is admitted that its reduced characteristics make its true affinities speculative. The sparsely developed ectosomal skeleton, composed of plumose bundles of smaller auxiliary styles, also indicates affinities to the Thalysias group, but in most respects (spicule geometry, skeletal structure, fibre development and growth form) it differs from all other species of either lsociella and Thalysias. Clathria (Isociella) skia was initially thought to be Amphilectus hispidulus Ridley, from Torres Strait (FNQ). From both published descriptions of A. hispidulus (Ridley, 1884a; Hentschel, 1911) it apparently lacked echinating megascleres, lacked principal spicules and had well developed subisodictyal skeletal structure, but re-examination of the holotype showed that it does have echinating acanthostyles, and differentiated principal and auxiliary megascleres (thus more appropriately included in Clathria (Clathria); see above), and it also has substantially different fibre characteristics from C. (I.) skia indicating that they are not conspecific. OTHER SPECIES OF CLATHRIA (ISOCIELLA) TABLE 28. Comparison between of Clathria (lsociella) skia sp.nov. Measurements in p.m, denoted as range (and mean) of spicule length x spicule width (N=25). Paratype (AMG5043) Holotype (QMG300449) SPI CULE Choanosomal principal styles absent absent zyxwvutsrqponmlkjihgfedcbaZYXWVUTS Auxiliary (coring) styles I 62-(248.4)-368x 2.5- ( 4.5) - 8 1544242.1)-318 x 244.1) - 6 Auxiliary (ectosomal) styles 97-(111.9)-129 x l.542.4)-4 87-(110.2)-132 x 1.5- ( 2.4) - 4 absent absent Echinating acanthostyles Chelae Toxas 9-(15.3)-18 14-(16.3)-I 9 724104.4)-145 x 65-(81.1)-108 x 0.2-(0.5)-0.8 0.240.4)-0.8 Clathria (Thalysias) Duchassaing & Michelotti, 1864 Thalysias Duchassaing & Michelotti, 1864: 82. Rhaphidophlus Ehlers, 1870: 19. Tenacia Schmidt, 1870: 56. Echinonema Carter, 1881a: 378. ? Thalassodendron Lendenfeld, 1888: 222. StylotellopsisThiele, 1905:456; de Laubenfels, 1936a: 112. Colloclathria Dendy, 1922: 74. Damoseni de Laubenfels, 1936a: 110. DEFINITION. Specialised ectosomal skeleton composed of two size classes of auxiliary (subtylo)styles, with smaller ectosomal spicules usually overlaying larger subectosomal ones forming a continuous palisade, or discrete bundles, mainly erect, sometimes paratangential, or rarely tangential to surface; choanosomal skeleton without any marked differentiation between axial and extra-axial regions; echinating acanthostyles usually present. TYPE SPECIES. Spongia juniperina Lamarck, 1814: 444 (by synonymy). REMARKS. Of 137 named species described in, or referred to Thalysias or one of its synonyms, 93 are thought to be valid of which 37 are recorded here from Australian waters including 10 new species. Clathria (Isociella) incrustans (Bergquist, 1961) Isociella incrustans Bergquist, 1961a: 42-43, text-figs Clathria (Thalysias) abietina (Lamarck, 1814) 15a-b [Ahipara Bay, N. New Zealand; originally (Figs 138-141, Tables 29-30, Plate 4C-D) assigned to the Suberitidae, Hadromerida]. Bergquist & Fromont, 1988: 114-116, p1.53, figs e-f, Spongia abietina Lamarck, 1814: 450, 377. p1.54, figs a-b; Dawson, 1993: 36.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathriaabietina; de Laubenfels, 1954: 141-142, textMATERI AL. HOLOTYPE: NMNZ unregistered. NZ. fig.90; Hooper & Wiedenmayer, 1994: 267. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 281 Clathria aculeata Ridley, 1884a: 443-444, p1.40, fig. 1, p1.42, fig.k; Ridley & Dendy, 1887: 147, 246, 254; Topsent, 1897b: 447; Burton, 1934a: 558; Burton, 1959a: 243; Levi & Levi, 1989: 80-81, p1.7, fig.3. Rhaphidophlus aculeatus; Topsent, 1932: 115, p1.4, fig. 10; Vacelet & Vasseur, 1977: 114; Vacelet et al., 1976: 73-74. Rhaphidophlus cervicornis, in part; Vacelet & Vasseur, 1971: 73. cf. Microciona prolifera; Vosmaer, 1935a: 610, 633, 664. MATERIAL. LECTOTYPE: MNHNDT634: precise locality unknown (suspected to be 'Australia'; Topsent (1932: 115)). PARALECTOTYPES: MNHNDT3342, 3343: same details. HOLOTYPE of C. aculeata: BMNH1882.2. 23.258: Torres Strait, Qld, 9°41'S, 142°17'E, 6-8m depth, coll. HMS 'Alert' (dredge). OTHER MATERIAL. MICRONESIA - USNM 22808, USNM23090. QLD - BMNH1887.5.2.110, QMG300791. DARWIN HARBOUR, NT NTMZ226, NTMZ426, NT MZ458, NTMZ468, NTMZ498, NTMZ512, NTMZ515, NTMZ886, NTMZ903, NTMZ928, NTMZ955, NTMZ2050, NTMZ2078, NTMZ 2079, NTMZ2085, NTMZ2258, NTMZ2390, NTMZ2391, NTMZ2395, NTMZ2399, NTMZ 2611, NTMZ2642, QMG300169, NTMZ2646, QMG304077, NTMZ I 943, NTMZ1958, NTMZ 1963, QMG303373, QMG303382, NTMZ2089, NTMZ2161, NTMZ2163, NTMZ2186, NTMZ2191, NTMZ2194, NTMZ2195, NTMZ820, NTMZ835, QMG300414. BYNOE HARBOUR, NT - NTMZ1073, NTMZ2106, QMG303447, QMG303534. SHOAL BAY, NT - QMG303539, QMG303571. TIMOR SEA, NT - NTMZ3090. PORT ESSINGTON, COBOURG PENINSULA, NT - NTMZ68, NTMZ69, NTMZ90, NTMZ1393, NTMZ3304, NTMZ577, NTMZ 1328, NTMZ1329, NTMZ1330, NTMZ1331, NTMZ1332, NTMZ1333, NTMZ1334, NTMZ 1343, NTMZ2500, NTMZ2501, NTMZ2509, NTMZ2510, NTMZ3245, NTMZ3255, NTMZ 3258, NTMZ3260, NTMZ3268, NTMZ3272, NTMZ3277, NTMZ3278, NTMZ3284, NTMZ 3289, NTMZ3295, QMG300386, NTMZ1352. ARAFURA SEA, NT - NTMZ2521, NTMZ 2522, NTMZ2523, NTMZ129, NTMZ130, NTMZ138. WESSEL ISLANDS, NT - NTMZ 3902, QMG300764 (NCIQ66C-4692-Q), NTMZ3921, NTMZ3930, QMG300757 (NCIQ 66C-4773-F), QMG300508 (NCIQ66C-4772-C), NTMZ3947, QMG300765 (NCIQ66C-4808-R). NORTHWEST SHELF REGION, WA - NTMZ1036, NTMZ1209, NTMZ1217, NTMZ 1244, NTMZ1272, NTMZ1314, NTMZ1411, NTMZ1423, WAM151-82, WAM15582 (fragments NTMZ1731, NTMZ1732), NTMZ1770, NTMZ1801, NTMZ1820, NTMZ1824, NTMZ 1852, NTMZ2272, NTMZ2329, NTMZ2349, NTMZ2486, NTMZ3017, NTMZ3030, NTMZ 3031, NTMZ3032, NTMZ3033, NTMZ3396, QMG300448 (NCIQ66C1517-P), QMG300117 (NCIQ66C1518-Q) (fragments NTMZ3488, NTMZ3489), PIB0004-595 (fragment QMG300051) HABITAT DISTRIBUTION. Shallow-water0-25m depth, predominantly on sides and tops of rock and dead coral heads, invariably exposed to currents. Deeper-water specimens (26-86m depth) mostly associated with exposed rock substrates in gravel, silt or shell-grit substrates; central SW Pacific Ocean (Low Isles, Great Barrier Reef (Burton, 1934a), Torres Strait (Ridley, 1884a; Ridley & Dendy, 1887)); E Indian Ocean (Arafura Sea, Timor Sea, mid-WA coast (present study)); also tropical Indo-west Pacific: central NW. Pacific Ocean (Marshall and Caroline Is (de Laubenfels, 1954), Philippines (Levi & Levi, 1989)); W. Indian Ocean (Madagascar (Vacelet et al., 1976, 1977), Red Sea (Burton, 1959a)). Within Australian waters this species extends across the N and NW coasts, from the Cairns region, Torres Strait and Gulf of Carpentaria, Qld, to the Exmouth Gulf region, WA (Fig. 138J). It is only rarely encountered on the E Qld. coast and must be considered a predominantly Indian Ocean species. DESCRIPTION. Shape. Arborescent; thickly cylindrical digitate branches on long or short, thick stalk (3-21mm basal diameter) with expanded basal attachment, never rhizomous; branches rarely anastomosing, branching mostly irregular, bushy, occasionally regular growing in I plane, sometimes flagelliform, occasionally expanded, club-shaped; total length and branch diameter variable (50-460mm; 2-22mm, respectively). Colour. Live colouration highly variable, ranging from maroon, red, orange, yellow, brown to grey; colour in ethanol ranges from black, brown to grey (Munsell values given below); pigments oxidise in air; maroon colouration is highly alcohol soluble; yellow pigmentation more stable in ethanol but easily scratched from surface of living sponge, leaving a maroon or mauve pigmentation beneath; pigmentation extends into periphery of choanosomal mesohyl. Oscules. Exhalent pores usually very small, 0.62.0mm diameter, barely visible optically, dispersed on distal ends of branches surrounded by surface conules, occasionally scattered along entire lateral margins of branches. Texture and surface characteristics. Firm, barely compressible; branches rubbery whereas stalk more rigid; surface dense, entirely opaque in life, with characteristic regularly scattered surface conules; conules tapering, distally rounded or blunt, usually absent from basal and distal extremities of branches, 2-6mm maximum basal diameter. Ectosome and subectosome. Very dense, continuous palisade of small ectosomal auxiliary subtylostyles forming discrete brushes overlay- 282zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 138. Clathria (Thalysias) abietina (Lamarck) (holotype MNHNDT634). A, Choanosomal principal subtylostyle. B, Ectosomal auxiliary subtylostyle. C, Subectosomal auxiliary subtylostyle. D-E, Echinating acanthostyles. F, Accolada toxa. G, Wing-shaped toxas. H, Palmate isochelae. I, Section through peripheral skeleton. J, Australian distribution. ing thickly plumose tracts of larger subectosomal secondary elements but forming irregular, closeauxiliary subtylostyles in peripheral skeleton (the meshed reticulation; fibre meshes oval to eliptilatter often obscuring ectosomal brushes); subec- cal, l00-350ttm diameter; fibres cored by tosomal spicule tracts arise from ultimate multispicular tracts of larger auxiliary subtyloschoanosomal fibres, intermingled with tufts of tyles occupying 60-80% of fibre diameter; principal choanosomal subtylostyles echinating choanosomal principal subtylostyle uncommon peripheral fibres; principal spicules not extending into ectosomal skeleton; mesohyl matrix heavy within fibres, mainly found at fibre nodes and pigmented in both ectosomal and subec- protruding through fibres individually or in bundles; fibres also echinated by acanthostyles tosomal regions.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA especially on peripheral fibres; mesohyl matrix Choanosom e. Skeleton irregularly reticulate; heavy spongin fibres, 50-125 t.i.m diameter, heavi- heavy but only lightly pigmented, slightly ly anastomosing, not divided into primary or granular; choanocyte chambers oval or eliptical, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 283 TABLE 29. Comparison in range of spicule dimensions between present and (more abundant in published records of Clathria (Thalysias) abietina. All measurements are given in smaller form). Length I: p.m and denoted as length x width (N=25). 7.9412.1)-19.9pm (lec- totype 11 -(12 .5 )14.5p,m), length II: 0.5Principal 172-254x 12.7 165-258 x 9-16 230 172-254 (5.8)-10Rm (lectotype 270-320 x 6-15 styles 4-(5)-6p,m). Subectosomal 157-343 x 5-10 350 x 8.5 149-276 x 2-5 120-250 x 2-3 127-387 x 1-20 Toxas relatively unstyles common with 2 Ectosomal 45-230x 1-10 62-132 x 2-5 81-117 x 2-4 geometric forms; (i) styles wing-shaped, short, 41-109 x 1-19 53-74 x 4-6 50-70 x 7-8 90 x 7.9 Acanthostyles 61-94 x 4-10 thin, generously curved 11-15 12.7 10-14 10-15 8-20 Chelae I at central with reflexed 4-6 1-10 Chelae II points; (ii) accolada 15-180 63 91-210 45-55 3-345 Toxas toxas long, thin almost Material: straight with small anholotype MNHN DT634. 1. Spongia abietina zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA gular central curvature 2. Clathria aculeata Ridley (1884a: 443). 3. De Laubenfels' (1954:141) material USNM 22808, 23090. and straight points. 4. Rhaphidophlus cervicornis, in part, (Vacelet & Vasseur, 1971:96). Length I: 5424.9)5. Present material (N=25). 75pm, width 1.8-(2.4)4.1pm (lectotype 40-170p.m diameter; larger auxiliary styles also 15-(30.8)-61 x 2.0-(2.9)-3.5p.,m). Length II: 58dispersed between fibres in disorganised tracts. (184.6)-345p,m, width 0.3-(0.8)-1.5pm (lecMegascleres. Choanosomal principal styles-sub- totype 65-(120.8)-180 x 0.540.9)-1.211m). tylostyles usually uncommon, short, stout, slightly curved at centre or near base, occasionally Larvae. Parenchymella larvae observed in straight, with slightly subtylote bases or evenly peripheral choanosomal skeleton of about 20 rounded bases; spicules usually completely specimens, predominantly during the tropical wet smooth, exceptionally with minutely season (February-April) and less so during the pre-dry season (May-July). Larval incubation microspined bases, tapering to abruptly (hastate) sharp points. Length 1214202 .7)-300.5p,m, was distinctly seasonal and occurred more-orwidth 5-(14.4)-24pm (lectotype 1654215.5)- less evenly throughout the population (i.e., irrespective of depth of collection; Fig. 141). 258pm x 9.5-(14.3)-15.5pm). Larvae were 600-950pm diameter, elongate-oval Subectosomal auxiliary subtylosytles straight in shape and many contained larval megascleres; or slightly curved near base, with prominent subcilia were not observed (preserved material). tylote, usually microspined bases, tapering to 3% of specimens had filamentous Associations. fusiform points. Length 127-(258.4)-386.9p,m, width 1.148.2)-20.1 p.m (lectotype 1574274.4)- algae coring fibres in addition to longer auxiliary megascleres. 343pm x 5-(6.7)-10p,m). Ectosomal auxiliary subtylostyles short, thin, Variation. Highly variable in live colouration, invariably straight, with prominently subtylote, non-fibre skeletal development, megasclere size, typically microspined bases, tapering to fusiform relatively consistent in growth form, surface feapoints. Length 44.8-(114.6)-230p,m, width 1.1- tures, fibre skeleton, spicule geometry. Gross (3.8)-10.1pm (lectotype 62-(85.5)-132 x 243.4)- morphology: stalked, bushy, branching in more than 1 plane, branches bifurcate, occasionally 5p,m). Echinating acanthostyles moderately long, anastomosing (46%), planar branching (33%), thick, straight, with prominently subtylote, dendritic planar branching (fans) (14%), or single spined base, virtually aspinose 'neck' (proximal digits with no or few bifurcations (7% of to base), and lightly spined shaft; spines relatively specimens). Atypical growth forms (few surface large. Length 40.6-(82.4)-109pm, width 1.1- conules, few branches, thin branching) found (7.5)-19pm (lectotype 61-(74.4)-94 x 446.5)- predominently in deeper offshore coastal waters (40m depth). Live colouration: highly variable, 1011,m). Microscleres. Palmate isochelae with long lateral no particular pigment considered to be typical, alae completely fused to shaft, shorter entire front ranging from (i) maroon, evenly pigmentation alae; two size categories present, both relatively (Munsell 5R4/10-2.5R4/8-10), (ii) red-maroon, abundant, both with examples of contort shafts even (5R518-10), (iii) bright red-orange, even SPICULE 1 - 2 3 4 5 MEMOIRS OF THE QUEENSLAND MUSEUM 284zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Clat hr ia ( Thaly sias) abiet ina (Lamarck) typical growth forms. A, Lectotype of C. aculeat a FIG. 139. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA BMNH1882.2.23.258. B, BMNH1887.5.2.110. C, Lectotype MNHNDT634. D, Paralectotype MNHNDT3343. E, Paralectotype MNHNDT3342. F-G, Deeper water specimens NTMZ1820, NTMZ1314. H, Shallow-water specimen NTMZ68. (5R5/12), (iv) bright orange, even (10R5112- 10R6/10-12-10R7/10), (v) orange-brown, even (10R418-10), (vi) orange-yellow, even (7.5YR7112), (vii) pale brown-pink, even (10R7/4), (viii) light brown, with pink conules (10R7/4, 5R8/4), (ix) light brown, with dark greybrown conules (2.5YR714-5YR712-6, 2.5YR6165 YR3-5/2), (x) pale brown, even (5YR616-5YR7/4-6-5YR8/4), (xi) muddy grey, even (7.5YR7/2-8/2) (xii) yellowish grey, even (2.5Y816), (xiii) yellow, even (2.5Y7/8), (xiv) yellow, with pink conules (2.5Y8/6, 10R7/6), (xv) lime-yellow, with dark brown conules (2.5Y7/10, 2.5Y5/2), (xvi) yellow-green mottle (5Y8/12). No obvious relationship between live colouration and depth or substrate type from present data; moreover, specimens with widely different pigmentation observed growing sideby-side (Plate 4D). Subectosomal skeletal development: correlation between branch diameter and extent of development of peripheral skeleton, with 8% of specimens (all thinly ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 285 TABLE 30. Summary of results from one-way ANOV's (Model I), testing for variability in spicule lengths and widths between locality, bathymetric and seasonal distributions of Clathria (Thalysias) abietina. LOCALITY SPICULE (N) DEPTH2 F Prob. F SEASON3 Prob. (N) F Prob. (775) 1.67 P>0.05 (775) 0.26 P>0.05 P<0.0005 (775) 2.46 P>0.05 4.04 P<0.01 (775) 2.08 P>0.05 5.48 P<0.001 (775) 4.15 P<0.01 Choanosomal styles L (1950) 3.31 P<0.05 2.11 P>0.05 W (1950) 0.54 P>0.05 4.44 P<0.005 Subectosomal (1950) 6.21 P<0.001 10.61 W (1950) 1.89 P>0.05 Ectosomal styles L (1950) 0.85 P>0.05 W (1950) 0.77 P>0.05 1.13 P>0.05 (775) 1.99 P>0.05 Acanthostyles L (1950) 4.74 P<0.01 5.95 P<0.0005 (775) 1.22 P>0.05 W (1950) 3.17 P<0.05 4.64 P<0.005 (775) 5.17 P<0.0025 Chelae I L (1925) 0.48 P>0.05 1.21 P>0.05 (775) 4.04 P<0.01 Chelae II L Toxas L (1925) 0.14 P>0.05 2.04 P>0.05 (775) 0.88 P>0.05 (1925) 0.39 P>0.05 2.72 P<0.05 (750) 1.17 P>0.05 W (1925) 0.42 P>0.05 5.49 P<0.001 (750) 1.33 P>0.05 styles L Number of groups: 1. 3 locality groups (NWS, DAR, CP localities). 2.4 depth groups (0-4m, 4-10m, I0-40m, 40m depth). 3. 4 seasonal groups (Darwin region only: wet (FMA), pre-dry (MJJ), dry (ASO), pre-wet (NDJ)). branching) having peripheral choanosomal fibres lying immediately below ectosomal crust, thin paratangential subectosomal region, and acanthostyles echinating peripheral fibres piercing ectosomal skeleton. Development of extra-fibre skeleton: 4% with very abundant juvenile auxiliary subtylostyles distributed throughout mesohyl, 90% with at least some interstitial auxiliary spicules, and 6% of specimens without any interstitial auxiliary spicules dispersed between fibres. Megasclere geometry: Principal spicules range from relatively common (21%), uncommon (61%), or very rare (18% of specimens). Bases of all principal spicules smooth (79%), or up to one-quarter of principal spicules with microspined bases (21% of specimens). Larger auxiliary spicules predominantly subtylote with minutely microspined bases, but 0-74% of spicules may be smooth in any particular specimen. Acanthostyle geometry relatively consistent, although in 5% of specimens two size categories were recognised (although smaller category probably juvenile form of larger and subsequently lumped together in analyses), in 4% they were significantly thinner, and 1% had significantly shorter and stouter acanthostyles than typical forms. Acanthostyle spination slightly variable, from scattered robust, recurved spines (65%), spines arranged in regular longitudinal rows (5%), or minutely microspined (12% of specimens). Microsclere geometry: Proportion of contort to normal morphs of palmate isochelae varied from 0-44% for smaller category, 0-20% for larger. Chelae typically abundant, 7% of specimens isochelae of both classes very rare, 3% smaller category rare but larger abundant, 4% larger category rare but smaller, 1% of specimens isochelae absent entirely. Toxas of both categories very abundant (12%), uncommon (70%), rare (17%), or absent entirely (1% of specimens). Variability in spicule dimensions: Some spicules (choanosomal principal styles, larger auxiliary subtylostyles, acanthostyles) showed significant variations in dimensions between samples collected from different localities (Tables 29-30), although statistical significance was never high, and no obvious patterns were apparent when groups of specimens from the same localities were compared with other groups. Some spicule categories varied between samples collected from different depths, in some cases with high levels of statistical significance (P<0.001) (e.g., larger auxiliary subtylostyles), but no obvious trends apparent. It is probable that effects of differential geographical and bathymetric distributions of specimens are linked due to the preponderance of deeper water samples from the Northwest Shelf region and shallow water samples from the Darwin region, making it im- 286zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 140. Clathria (Thalysias) abietina (Lamarck) (A-B, NTMZ2642; C-I, QMG303447). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E, Base of subectosomal auxiliary subtylostyle. F-G, Palmate isochelae. H, Wing-shaped toxas. I, Accolada toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 287 SEASON TOTAL SAMPLES SAMPLES WITH LARVAE WET 15 PREDRY 17 9 3 DRY PREWET 19 0 17 0 DEPTH 0-4m 5 0 4-10m 30 7 10-40m 23 5 >40m 11 0 FIG. 141. Clathria (Thalysias) abietina (Lamarck) Incidence of incubated larvae within sampled populations from NW Australia. possible to separate the influences of either factors on this observed variability. Variability of spicule size between samples collected during different seasons were analysed for Darwin samples only (Table 30), with significant differences observed for lengths of ectosomal auxiliary subtylostyles and larger isochelae, and widths of acanthostyles. Data also indicate a higher level of variability in sizes of larger isochelae during the wet season (February-April) than during other seasons, although this result is of uncertain biological significance. REMARKS. Notwithstanding its considerable morphological variability C. (T.) abietina is easily recognised in the field with distinctive stalked digitate growth form and prominent surface conules. The most similar species in growth form is C. (T) cervicornis but this has a much thinner, stoloniferous branching morphology, lacks principal megascleres completely, fibres have less spongin, toxas are different in morphology and size, and spicule dimensions are different. Skeletal structure is unusual where larger auxiliary subtylostyles are found in three locations in the skeleton: 1) subectosomal skeletal tracts forming organised plumose tracts supporting the ectosomal skeleton; 2) dispersed between fibres in disorganised tracts; 3) and coring all spongin fibres to the virtual exclusion of principal spicules. Principal styles mainly form plumose brushes protruding from fibres in plumose bundles, functionally representing a second category of echinating spicule. There are some minor differences between type material and other specimens examined, including a higher proportion of choanosomal principal spicules found echinating fibres, the absence of contort isochelae, and specific details in some spicule dimensions (Table 29). But given the large range of variation in some characters, the relatively large sample sizes from widely dispersed populations, and the antiquity of the dried holotype this variability is insignificant. Re-examination of Ridley's (1884a) holotype of C. aculeata confirmed that it is a synonym of C. (T) abietina, with shape, texture, colour in spirit, spiculation and skeletal architecture virtually identical. Conversely, C. (T) coralliophila (see below) has different spicule geometry and skeletal architecture, and Burton's (1959a: 243) proposed merger of C. coralliophila into C. aculeata is rejected. Specimens described by de Laubenfels (1954) from the central west Pacific differ from Australian populations in that they have a more restricted size range of isochelae (i.e., one size category), relatively small acanthostyles (Table 29), and an ectosomal (peripheral) skeleton almost completely covered by subectosomal spicule brushes (as opposed to ectosomal spicule brushes). In fact de Laubenfels (1954) completely overlooked the presence of ectosomal megascleres, not differentiating between spicules coring fibres from those forming the peripheral skeleton. Clathria (Thalysias) aphylla sp. nov. (Figs 142-143, Plate 5D) MATERIAL. HOLOTYPE - QMG300477 (NCIQ66C-4640-K): NW. of E. Passage, Easter Group, Houtman Abrolhos, WA, 28°40'S, 113°50'E, 20m depth, 17.ix.1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION. Staghom and plate coral fringing reef; 20m depth; known only from the type locality, Houtman Abrolhos (WA) (Fig. 142H). DESCRIPTION. Shape. Thin, leaf-like, foliaceous, convoluted, basically frondose bundles of lamellae covering coral substrate; individual fronds attached directly to substrate, sometimes completely enveloping staghorn corals, or attached via small basal stalk, up to 24mm long, 5mm diameter, or attached to adjacent lamellae; individual lamellae usually flat, elongate, oval or eliptical, up to 65mm long, 2nun thick, with rounded or sinuous, convoluted margins, superficially resembling a Padina algae, or palmate-digitate margins, or sometimes curled in vasiform growth forms. Colour. Dull yellow alive (Munsell 2.5Y 8/8), pale brown in ethanol. Oscules. Small, on upper surface, less than 2rnm diameter, flush with surface. 288zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Texture and surface characteristics. Soft, compressible but difficult to tear; upper osculiferous surface slightly concave, smooth, usually even, occasionally concentrically striated, or producing short fronds of folds on surface; lower porous surface slightly convex, even or slightly striated, with parasitic zoanthids covering parts of surface. Ectosome and subectosome. Erect or paratangential brushes of ectosomal auxiliary styles forming thin surface layer, with choanosomal principal styles protruding through ectosome perpendicular to surface, singly or several together, extending up to 150p.m through surface; choanosomal fibres immediately below ectosome, with reduced subectosomal skeleton; subectosomal auxiliary styles tangential or paratangential, lying immediately below surface underlying ectosomal spicule brushes; mesohyl matrix in peripheral skeleton heavy, darkly pigmented. Choanosome. Regularly reticulate, slightly renieroid skeleton; without compression or major differences between peripheral skeleton and core; spongin fibres heavy, 20-70p,m diameter, not obviously differentiated into primary or secondary elements, cored by multispicular (up to 15 spicules abreast) or paucispicular (2 or more spicules abreast) tracts of principal choanosomal styles; at core of skeleton principal styles confined entirely within fibres whereas in peripheral fibres, principal spicules erect, perpendicular to fibres, protruding through surface in plumose bundles; fibres moderately lightly echinated by acanthostyles, mainly at fibre nodes; fibre meshes mainly rectangular (fibres oval), 90-170iim diameter, with some triangular meshes, relatively even throughout skeleton; mesohyl matrix light, with few scattered megascleres; choanocyte chambers oval to eliptical, 25-40Rm diameter, lined by abundant, minute isochelae. Megascleres. Choanosomal principal styles long, thick, straight or slightly curved towards basal end, rounded or slightly tapering bases, invariably smooth bases, fusiform points. Length 152(205.1)-252pm, width 4.5-(8.4)-11p.m. Subectosomal auxiliary styles long, slender, straight, with rounded or faintly subtylote smooth bases, fusiform points. Length 208-(258.l)297 p.m, width 2-(2.7)-4.5 p.m. Ectosomal auxiliary styles short, slender, straight, rounded or slightly subtylote bases, smooth or microspined (or mucronate) bases, fusiform points. Length 162-(178.2)-204Rm, width 1-(1.6)-2.5p.m. Echinating acanthostyles relatively long, slender, subtylote, sharply pointed, evenly spined although fewer spines in 'neck' region proximal to base (but not aspinose); spines small, sharp, recurved. Length 45-(86.3)-102Rm, width 3(5.5)-81.1.m. Microscleres. Palmate isochelae very abundant, minute, with lateral alae completely fused to shaft and partially fused to front ala. Length 445.1)6p.m. Toxas not abundant, predominantly v-shaped, variable in size, with angular central curvature and arms bent at approximately right angles to each other, non-reflexed arms, occasionally toxas forceps shaped with pinched central curve and nearly parallel arms. Length 34-(62.3)-106p.m, width 0.8-(1.3)-2.5pan. ETYMOLOGY. Greek phyllon , leaf; for the growth form. REMARKS. This species is a sister of C. (C.) angulifera (Vic. and S Qld), having similar live colour, slightly renieroid skeletal structure (mainly rectangular meshes), similar toxa morphology and chelae size. In particular both species have large v-shaped toxas, and a skeletal architecture that verges on isodictyal (some triangular meshes). However, they are clearly different species showing both obvious and subtle differences in a number of characters. Clathria (T) aphylla has a very thin, leaf-like growth form (whereas C. (C.) angulifera is thickly lamellate, lobate); very thick, well developed, evenly spaced fibres cored by multispicular tracts of large principal styles (versus widely spaced, vestigial fibres with light spongin, cored by much smaller principal spicules in uni- or paucispicular tracts); a specialised ectosomal skeleton composed of two size classes of auxiliary spicules forming brushes and principal spicules protruding through the surface forming plumose brushes and piercing the ectosome (versus a single category of auxiliary spicule forming a tangential or occasionally paratangential ectosomal skeleton, without participation of principal spicules in the ectosomal skeleton); clearly differentiated principal and auxiliary spicule geometry (versus more subtle differences, mainly in thickness and basal termination); rounded bases of principal and auxiliary megascleres with the smaller microspined (versus subtylote and completely smooth); and substantially larger dimensions for most megascleres. Another species with a similar, slightly renieroid skeleton reminiscent of C. (T.) aphylla is C. (C.) hispidula ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 289 FIG. 142. Clathria (Thalysias) aphylla sp.nov. (holotype QMG300477). A, Choanosomal principal styles. B, Subectosomal auxiliary style. C, Ectosomal auxiliary styles. D, Echinating acanthostyles. E, Palmate isochela. F, V-shaped toxas. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. J, Protruding choanosomal spicules in ectosomal skeleton. (from Torres Strait and Shark Bay), which has only one category of auxiliary spicule and different ectosomal structure, wing-shaped toxas, different acanthostyle geometry and specific differences in spicule sizes. All three species probably have sister species relationship based on skeletal architecture, here referred to the `anguliferd group. A few other Clathria species also have Vshaped toxas (C. (T) juniperina) and vaguely isodictyal skeletal structure (e.g., C. (T)hirsuta), but these differ from C. (T) aphylla in most other respects and are not considered here to be closely related. Clathria (Thalysias) arborescens (Ridley, 1884) (Figs 144-145) Rhaphidophlus arborescens Ridley, 1884a: 450-451, p1.40, fig.L, p1.42, figs n-n'; Thiele, 1903a: 958. MEMOIRS OF THE QUEENSLAND MUSEUM 290zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 143. Clathria (Thalysias) aphylla sp.nov. (holotype QMG300477). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Base of subectosomal and ectosomal auxiliary styles. G, Palmate isochela. H, V-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 291 Clathria arborescens; Hooper & Wiedenmayer, 1994: 268. cf. Microciona prolifera tropus spinosa; Vosmaer, 1935a: 610, 634. MATERIAL. HOLOTYPE - BMNH1881.10.21.272: Friday I., Torres Strait, Qld, 10°36'S, 142°10'E, 1.v.1881, coll. HMS 'Alert' (dredge). HABITAT DISTRIBUTION. Ecology unknown; Torres Strait, Qld (Fig. 14411). DESCRIPTION. Shape. Branching, arborescent, 125rrun long, 60mm maximum width of branching, with short stalk, 18mm long, 8mm diameter, no basal attachment (presumed not collected), proliferous bushy, slightly flattened cylindrical branches, up to 12mm diameter, with individual branches up to 55mm long, mostly anastomosing with adjacent branches; branching mostly planar. Colour Live colouration unknown, grey-brown in dry state. Oscules. Not observed. Texture and surface characteristics. Texture harsh in dry state; surface peel thick, darker than choanosomal skeleton, lightly hispid, with numerous irregular subectosomal striations, irregular conules and low ridges. Ectosome and subectosome. Ectosomal skeleton with very thick crust of discrete spicule brushes composed of ectosomal auxiliary subtylostyles, occasionally paratangential or tangential to surface, supported below by thick, vaguely plumose, slightly disorganised bundles of larger subectosomal auxiliary subtylostyles arising from terminal choanosomal fibres; subectosomal skeleton lacking any spongin fibres and only light mesohyl matrix; together ectosomal and subectosomal bundles extend up to 800Rm from peripheral fibre skeleton, and comprise up to 25% of branch diameter. Choanosome. Choanosomal skeleton irregularly close-meshed reticulate. Spongin fibres very large, heavy dark yellow-brown, 110-17011,m diameter, forming tight oval or elongate meshes, 130-350Rm diameter; fibre reticulation without any clear distinction between primary or secondary components, although ascending fibres marginally more abundantly echinated than transverse connecting fibres, and some smaller fibres between major meshes with uni- or paucispicular core; fibres generally all cored by multispicular tracts of choanosomal principal subtylostyles together with subectosomal auxiliary subtylostyles, forming a dense axial core within fibre but occupying only about 50% of fibre diameter; fibres abundantly echinated by relatively large acanthostyles, more-or-less evenly dispersed throughout skeleton (not congregated in periphery as reported by Ridley); choanocyte chambers not observed (dry material). Megascleres. Choanosomal principal subtylostyles with similar geometry to larger auxiliary spicules, long, straight, usually with subtylote, smooth or microspined bases, rarely rounded bases, with tapering fusiform points. Length 205(225.3)-262m, width 8-(9.7)-12p.m. Subectosomal auxiliary subtylostyles more slender but longer than principal spicules, straight or slightly curved towards base, with smooth, slightly constricted bases, fusiform points. Length 210-(251.4)-282p,m, width 4-(5.4)-7p.m. Ectosomal auxiliary subtylostyles relatively long, straight or slightly curved towards base, prominently subtylote, with smooth or microspined bases, fusiform points. Length 82(132.7)-147p.m, width 2-(3.3)-4pLm. Echinating acanthostyles long, robust, subtylote, large conical (not recurved) spines, scattered mainly over base and midsection of shaft, frequently with bare point and 'neck'. Length 68-(76.3)-88p.m, width 6-(8.2)-11p,m. Microscleres. Palmate isochelae small, frequently contort, single size category, with long lateral alae completely fused to shaft and entire front ala of equal length. Length 6-(10.8)-14p.m. Toxas short, robust, wing-shaped, with generous angular central curvature, recurved and slightly reflexed points Length 27-(41.8)-58p.,m, width 1.5-(1.8)-2.0p.m. REMARKS. This species is relatively poorly known, recorded so far only from the holotype. But several important characters were seen in the holotype using SEM (Fig. 145), not described by Ridley (1884a). These include: presence of differentiated principal and auxiliary spicules, presence of toxas, spination pattern on acanthostyles, and his spicule dimensions were innaccurate. These characters now provide better clues as to its affinities. Ridley (1884a) considered this species similar to C. (T) cactiformis in growth form and skeletal structure. They also both have principal and auxiliary megascleres very similar in geometry (i.e., on first appearances fibres seem to be cored only by auxiliary spicules), but they have many morphological differences. Clathria (T)arborescens has a substantially better developed ectosomal peel occupying a significant proportion of branch diameter, a close-meshed reticulate MEMOIRS OF THE QUEENSLAND MUSEUM 292^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 144. Clathria (Thalysias) arborescens (Ridley) (holotype BMNH1881.10.21.272). A, Choanosomal principal subtylostyles and bases. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyle. E, Wing-shaped toxa. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. choanosomal skeleton, very thick fibres all cored by megascleres (C. (T) cactifonnis has a relatively thin ectosomal skeleton, cavernous subectosomal region, wide-meshed choanosomal reticulation, substantially thinner fibres with only primary ones cored). Also, echinating acanthostyles are much larger than those of cactiformis, they have erect conical spines rather than hooklike recurved ones, and acanthostyles are found throughout the skeleton rather than mainly on peripheral fibres (although both species have spines absent from both "necks" and points of acanthostyles). Toxa morphology differs markedly between the two, with arborescens having only thicker, wing-shaped toxas, and cactiforniis with mainly very thin, raphidiform, sinuous or accolada toxas but occasionally also with small wing-shaped forms. Spicule dimensions also differ considerably between the two species. Both species belong to the juniperina' species complex (refer to discussion below under C. (T) cactifonnis and C. (T.) hirsuta). Dendy (1922), Burton & Rao (1932) and Burton (1938a) merged C. arborescens with C. (T) procera on the basis that choanosomal principal megascleres were supposedly excluded from the axis of fibres, but this is not entirely correct (in any case the character is also known to occur in ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 293 FIG. 145. Clathria (Thalysias) arborescens (Ridley) (holotype BMNH1881.10.21.272). A, Choanosomai skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of ectosomal auxiliary subtylostyle. F, Wing-shaped toxas. G, Palmate isochelae. 294zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM at least eight other species (see below under C. cactiformis)). Thiele (1903a) also compared C. (T) arborescens and C. (T) cervicornis, for the same reason (i.e., supposed absence of principal spicules), but C. procera does have principal spicules (exclusively echinating peripheral fibres), and those in C. (T) arborescens are intermingled with auxiliary subtylostyles within fibres. Skeletal structure and growth forms also differ considerably between these two taxa. Clathria (Thalysias) cactiformis (Lamarck, 1814) (Figs 146-148, Table 31, Plate 4E-F) Spongia cactifortnis Lamarck, 1814: 440, 370 Clathria cactiformis; Rudman & Avern, 1989: 335; Hooper & Wiedenmayer, 1994: 268. Spongia appendiculata, in part, Lamarck, 1814: 383, 362. Wilsonella appendiculata; Topsent, 1930: 46, p1.2, fig.3; Topsent, 1933: 22. Echinonema typicum Carter, 1875: 195 [nomen nudum]; Carter, 1881: 362, 377, 388, 384. Thalassodendron typica; Lendenfeld, 1888: 223; Hallmann, 1912: 242; Hal!mann, 1914a: 267. Not Thalassodendron typica; Whitelegge, 1901: 86. Clathria typica; Vosmaer, 1885b: 357; Ridley & Dendy, 1887: 153; Dendy, 1889c: 11;[in part] Dendy, 1896: 32; Whitelegge, 1901: 80, 117; Hentschel, 1911: 280, 364-367, text-fig. 43. Not Clathria ypica var. porrecta Hentschel, 1912: 298, 359-360. Not Clathria typica; Kirkpatrick, 1903: 248-249; Kirkpatrick, 1904: 148; Vacelet & Vasseur, 1971: 94-95, p1.3, fig.5, text-fig. 48. Wilsonella typica; Hallmann, 1912: 203, 242. Not Tenacia typica var. porrecta Hallmann, 1920: 771. Rhaphidophlus typicus; Hallmann, 1912: 176-177, 184-203, 234, 299, 300, p1.27, p1.28, figs 1-4, p1.29, text-figs 38-42 [et yard; Shaw, 1927:424-425; Topsent, 1932:88, p1.4, fig.4; Burton, 1938a: 12; Guiler, 1950: 8; Vacelet & Vasseur, 1977: 115; Vacelet et al., 1976: 73; Liaaen-Jensen et al., 1982: 170-172. Echinonema anchoratum Carter, 1881a: 362, 379, 380; Lendenfeld, 1888: 219-220. Echinonema anchoratum var. dura Whitelegge, 1901: 81. Echinonema anchoratum var. lamellosa; Lendenfeld, 1888: 219-220. Rhaphidophlus anchoratum; Bergquist & Fromont, 1988: 112; Dawson, 1993: 39. Wilsonella anchoratum var. lamellosa; Hall mann, 1912: 211 [imperfectly known]. Not Echinonema anchorata var. lamellosa; Whitelegge, 1901: 82. Echinonema flabelhformis Carter, 1885f: 352. Echinonema pectiniformis Carter, 1885f: 353. Phakelliaventilabrum var. australiensis Carter, 1886g: 379. Thalassodendron brevispina Lendenfeld, 1888: 225; Whitelegge, 1901: 87. Clathria favosa Whitelegge, 1907: 498-499. Clathria indurata Hallmann, 1912: 299. Clathria acanthodes Hentschel, 1911: 367-370, text- fig.44; Hentschel, 1923: 380, fig.349. Rhaphidophlus acanthodes; Hallmann, 1912: 177. Rhaphidophlus sp. 2; Vacelet & Vasseur, 1971: 97-98, text-fig. 52. cf. Microciona proltfera; Vosmaer, 1935a: 609, 628629, 647, 667. MATERIAL. LECTOTYPE: MNHNDT580: 'Southern Seas', Peron & Lesueur collection, no other details known. PARALECTOTYPE: MNHNDT3360: same details (= R. typicus var. cactiformis; Topsent, 1932: 88). LECTOTYPE of S. appendiculata: MNHNDT526 (fragment BMNH1953.4.9.83) (dry). PARALECTOTYPE of S. appendiculata: MNHNDT3394: King George Sound (Albany), WA. LECTOTYPE of T. brevispina: AMZ931: Port Jackson, NSW, 33°48'S, 151°20'E, coll. R. von Lendenfeld (dredge). SYNTYPE of E. typicum: BMNH1877.5.21.149: Fremantle, WA, 32°03'S, 115°38'E, coll. E. Clifton (dredge). HOLOTYPE of C. favosa: AMZ944: Off Port Jackson, NSW, 33°50'S, 151°30'E, 98-100m depth, coll. FIV 'Thetis' (dredge). LECTOTYPE of R. typicus var. proximus: AMZ930: Henley Beach, St. Vincent Gulf, SA, 34 455, 137°57'E, coll. T. Worsnop (dredge). PARATYPE of R. typicus var. obesus: AMZ937: Tuggerah Beach, NSW, 33°18'S, 151°30'E, coll. NSW Fish Commission (trawl). HOLOTYPE of R. typicus var. geminus: AMZ928: specific locality unknown, WA, no other details known. HOLOTYPE of R. typicus var. stellifer AME648: E. coast of Hinders I., Bass Strait, Tas, 40°01'S, 148°02'E, no other details known. SYNTYPE of E. anchoratum: BMNH1886.12.15.423: Port Phillip, Vic, 38°09'S, 144°52'E, no other details known. SYNTYPE of E. anchoratum var. dura and LECTOTYPE of C. indurata: AMG9113: specific locality unknown, WA, no other details known. PARALECTOTYPE of C. indurata: AM unreg.: specific locality unknown, WA, no other details known. HOLOTYPE of E. pectiniformis: BMNH1886.12.15.141 (fragment AMG2777): Port Phillip, Vic, 38°09'S, 144°52'E, 40m depth, no other details known. Fragment of HOLOTYPE C. acanthodes: ZMB4443: Surf Point, outer bar exit to South Passage, Shark Bay, WA, 26°08'S, 113°08.5'E, 0.5-3.5m depth, coll. W. Michaelsen & R. Hartmeyer (dredge). OTHER MATERIAL. NSW - QMG301403. TAS - AME624, AMZ1415, AMZ2203, QMG311412 (NCIQ66C3713-A) (fragment NTMZ3811). VICTORIA AMZ1430, AMZ1158, NMV RN359, NMVRN43 I, NMVRN436, NMVRN 438, NMVRN551, NMVRN677, NMVRN797, NMVRN840, NMVRN900, NM V RN959, AMZ3921 (RRIMPFN3527/000/02), AMZ4277 (RRIMPFN1906), NTMZ2832. S AUST AMZ41, SAM unreg. (fragment NTMZ1657), SAM unreg. (fragment NTMZ 1693), SAM unreg. (fragment NTMZI628), SW COAST, WA - MNHN unregistered 0 ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 295zyxwvutsrq A^BzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 146. Clathria (Thalysias) cactiformis (Lamarck) (lectotype MNHNDT580). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyles. E, Sinuous and wing-shaped accolada toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. (Peron & Lesueur collection), WAM656-81(1), WAM628-81(1), WAM631-81(1), WAM655-81(1). HOUTMAN ABROLHOS, WA - QMG300731 (NCIQ66C-4213-U), QMG3046 60, AMZ4331 (RRIMP-ZW62), WAM639-81(1), WAM661-81(1), W A M660-81, QMG300736 (NCIQ66C -4189-T), QMG300733 (NCIQ66C-4197-C), QMG300730 (NCIQ66C-4214-V). NORTHWEST SHELF, WA WAM133-82, NTMZ1128, NTMZ3018, NCIQ66C1460-C (fragment NTMZ3347). ERITREA, RED SEA - PIBOCB20-125 (fragment QMG300055). SOMALIA, W. INDIAN OCEAN - PIBOCB12-388 (fragment QMG300060). SEYCHELLES - PIB0001140 (fragment QMG300063). HABITAT DISTRIBUTION. Colonises a wide variety of substrates, including rock reefs, coral-rubble, soft sand, mud sediments, algal beds; shallow subtidal to 100m depth; widely distributed in temperate waters of eastern and western Australian coasts, with incursions into subtropical waters extending to the Port Hedland region on the west coast and Byron Bay on the east coast (Fig. 146H). This species is not yet recorded from the Indonesian archipelago, contrary to Hentschel (1912) — his records being misidentifications (see below) — but it does appear to range across the Indian Ocean, with confirmed records from Madagascar (Vacelet et al., 1971, 1976, 1977), East Africa and Seychelles (present study). DESCRIPTION. zyxwvutsrqponmlkjihgfedcbaZY Shape. Typically flabellate, planar, stalked, with long bifurcate, occasionally anastomosing digitate processes on margin of fan; gross morphology variable, with or without basal stalk and expanded basal attachment, varying from globular, cylindrical planar digitate, globular digitate, planar flabellate, lamellate branching in more than one plane, or thinly lamellate. Colour. Live colouration pale orange-brown (Munsell 5YR8/4) to bright red (5R5110); beige, ^ MEMOIRS OF THE QUEENSLAND MUSEUM 296zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 147. Clathria (Thalysias) cactiformis (Lamarck) typical growth forms. A, Lectotype MNHNDT580. B, Paralectotype MNHNDT3360. C, Lectotype of S. appendiculata MNHNDT526. D, Paralectotype of S. appendiculata MNHNDT3394. E-F, Variety cactiformis WAM656-81(1), WAM655-81(1). G, Variety stellifera NTMZ1708. H, Variety geminus WAM639-81(1). I, Somalia specimen PIBOCB12-388. J, Syntype of E. typicum BMNH1877.5.21.149. K, QMG300453. L,Variety obesus with Rostanga feeding (photo W. Rudman) NTMZ2832. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 297 FIG. 148. Clathria (Thalysias) cactiformis (Lamarck) (QMG300225). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Echinating acanthostyles. E, Acanthostyle spines. F-G, Bases of subectosomal and ectosomal auxiliary subtylostyles. H-I, Palmate isochelae. J, Accolada toxas. K, Wing-shaped toxas. 298zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM skeletal components; fibre anastomoses produce widely spaced rectangular meshes formed by ascending primary fibres, containing pauci- or multispicular bundles of choanosomal principal styles which occupy only the central portion of fibres, and uncored secondary fibres; occasionalSpecimens S.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Lectotype of zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Lectotype SPICULE ly secondary fibres uni- or paucispicular, usually (N=-46) appendiculat a with an optically diffuse pith reminiscent of dic103.6(199.2)Choanosomal 134(191.2)157-181 x tyoceratid sponges; spongin fibres charac314 x 3.2232 x 5-(7.6)principal 4.5-6 styles (9.7)-19.4 11 teristically thin, without marked differences between primary and secondary fibre diameter, 11534235.7)Subectos°mal 2 044229.8)191-297x 356.6 x 1.6auxiliary with pigmentation ranging from extremely light, 4-5.5 8 x 4-(5.4)-6 styles (6.2)-16.9 yellow (specimens from Port Phillip, Vic) to ex73.8(114.7)101(124.3)tremely heavy and dark brown (material from Ectosomal 85-104 x 2-3.5 162.1 x 1.6148x 1.5auxiliary Port Hedland, WA); echinating acanthostyles styles (4.4)-13.6 (2.5)-3 very heavy, especially on peripheral fibres imme38.2-(61.3)diately below subectosomal skeleton; larger 68-(74.4)-79 x Echinating 93.1 x 1.746-57 x 3-4.5 acanthostyles 5.5-(8.8)-11 auxiliary subtylostyles usually abundant between (6.7)-13.1 fibres, strewn without order within mesohyl; 8.5-(13.9)-22.2 7-8 12-(14.7)-18 Chelae I mesohyl matrix relatively light, choanocyte 2.5-(7.1)-11 15-17 6-(8.8)-I0 Chelae II chambers large, oval or elongate, up to 12011m 1134201.7)-296 82-146 x 0.5- 8-(127.8)-355 diameter. Toxas x 0.8-(1.3)-2.5 x 0.8-(1.2)-2 1.0 Megascleres (Table 31). Choanosomal principal styles morphologically similar to larger auxiliary brown, to dark brown in ethanol; ectosomal crust subtylostyles but relatively shorter, thicker, usually lighter colour than choanosomal regions. slightly curved near base, usually with tapering, Oscules. Oscules typically large (up to 3mm dia- evenly rounded smooth bases but sometimes meter), dispersed over entire sponge surface, or slightly subtylote, and fusiform points. sometimes clumped in special pore areas; oscules Subectosomal auxiliary subtylostyles nearly usually flush with surface, without raised lip. completely straight, occasionally slightly curved Texture and surface characteristics. Firm, com- or sinuous, usually with slightly subtylote smooth pressible, rubbery; surface characteristically bases, less frequently with microspined bases, fleshy, smooth but uneven, with bumps, ridges, tapering to fusiform points. depressions and subdermal striations commonly Ectosomal auxiliary subtylostyles more dispersed over surface; opaque skin-like covering prominently subtylote, slightly curved at centre, present stretched between adjacent conules in with microspined or smooth bases and fusiform live material; microscopic ridges and points. microconules abundant. Echinating acanthostyles relatively short, Ectosome and subectosome. Ectosomal skeleton stout, prominently subtylote, with large spines predominantly heavy, with a dense crust of mainly dispersed on base and midsection, leaving smaller auxiliary subtylostyles forming mainly a bare 'neck' (below basal swelling) and bare erect or less often paratangential brushes; oc- point. casionally ectosomal skeleton reduced to rela- Microscleres (Table 31). Larger palmate tively thin tangential or paratangential crust; isochelae variable in abundance, with lateral alae larger subectosomal auxiliary styles perched on completely fused to shaft, wide gap between ultimate choanosomal fibre endings, forming lateral and front alae, and fused front ala; usually plumose tracts underlaying brushes of smaller non-contort. Smaller isochelae common, freectosomal spicules, usually protruding through quently contort. and obscuring ectosomal bundles; subectosomal Toxas differentiated into two main morphs, region cavernous, with well differentiated, wide- larger often found in toxodragmata; accolada ly spaced plumose tracts of larger auxiliary sub- toxas most common, long, very thin, slight antylostyles. gular central curvature, straight arms, sometimes Choanosome. Skeleton irregularly reticulate, sinuous asymmetrical. Small toxas less abundant, vaguely radial, with clearly differentiated relatively thick, widely curved at centre, slightly primary ascending and secondary transverse reflexed points. TABLE 31. Comparison between present and published records of Clathria (Thalysias) cactiformis (Lamarck). All measurements are given in p.m, denoted as range, or as range (and mean) of spicule length x spicule width (N=25). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 299 Larvae. Reproductive products observed in only four specimens. Incubated parenchymella larvae large, oval to eliptical in shape, 220-420Rm diameter, containing well developed centrally aggregated larval styles. Associations. Heavy infestations of zoophytic organisms observed on surfaces of several specimens (both compound ascidians and zooanthid polyps); numerous microalgal cells seen in sponge mesohyl of few samples; sponge apparently serves as a food source to Rostanga sp. nudibranchs (W. Rudman, pers. comm.). Variation. Extremely variable in some features (e.g., gross morphology, spicule size) but not in others (e.g., surface features, spicule geometry, skeletal structure). Gross morphology: Victorian, Queensland and Indonesian morphs showed greatest differences in growth form, but all specimens retained most other characteristics common to other populations. Ectosomal skeletal structure: typically heavy, erect plumose brushes, well differentiated from subectosomal plumose tracts (52%), moderately developed (34%), or thick tangential ectosomal peel reminiscent of Crella (Crellidae) (14% of specimens). Subectosomal skeletal structure: very cavernous, well differentiated ascending plumose (extra-fibre) skeletal tracts (36%), paratangential, immediately subdermal tracts (21%), or intermediate of these conditions (43% of specimens). Choanosomal skeletal structure: irregularly reticulate (64%), regular fibre anastomoses (22%), or predominantly radial to plumose primary fibre architecture (14% of specimens). Fibre meshes: wide, rectangular, with uncored secondary elements (67%), with central pith component (7%), close-meshed, irregularly oval to eliptical fibre reticulation with uncored secondary fibres (7%), or with both cored primary and secondary fibre elements (19% of specimens). Spongin fibres: very light (7%), moderately heavy, yellow (48%), very heavy, yellow (43%), or extremely heavily, dark brown in colour (2% of specimens). Echinating acanthostyles: concentrated on peripheral fibres, more sparsely dipersed on axial fibres (69%), evenly distributed throughout fibres (17%), rare (14% of specimens). Extra-fibre spicules: typically abundant, dispersed without order within mesohyl (74%), very light or absent entirely (24%), or organised into ascending extra-fibre tracts (2% of specimens). Megasclere geometry: Choanosomal principal styles: predominantly fusiform (76%), distinctly hastate (abruptly pointed) points (7%), or mixture of both types of terminations (17% of specimens); bases of spicules predominantly smooth (90%), with both smooth and spined bases (8%), or only microspined bases on choanosomal styles (2% of specimens); bases of spicules tapering, non-tylote (46%), rounded, slightly subtylote (52%), or with both conditions (2% of specimens); megascleres slightly curved (83%), exclusively straight (7%), or both conditions (10% of specimens). Subectosomal auxiliary subtylostyles: tapering, rounded or very slightly subtylote bases (69%), or distinctly subtylote (31% of specimens); exclusively straight (69%), predominantly curved (7%), or both conditions (24% of samples). Ectosomal auxiliary subtylostyles: subtylote (76%), tapering nontylote (10%) or both conditions (14% of specimens); bases predominantly microspined (71%), exclusively smooth (3%), or with both conditions (26% of specimens); ectosomal spicules slightly curved near base (54%), completely straight (29%), or with examples of both (17% of specimens); most megascleres with fusiform points (95%), or also including few hastate spicules (5% of specimens). Echinating acanthostyles: typically short, fusiform (69%), significantly smaller, stouter (19%), or longer, thinner (12% of specimens); bases subtylote (81%), or distinctly non-tylote (19% of specimens); spines characteristically long, thick, recurved (74%), or reduced, granular spination (26% of specimens). Microsclere geometry: Larger isochelae: common (70%), uncommon (26%), rare (2%), or common but poorly silicified (2% of specimens); contort larger isochelae present but uncommon (10%), present, common (2%), or absent (86% of specimens). Smaller isochelae: abundant (7%), common (62%), uncommon (21%), rare (3%) or absent entirely (7% of specimens); 0-10% of smaller chelae were contort (2%), 10-30% contort (5%), 30-50% contort (12%), 50-70% contort (24%), 70-90% contort (43%), or 100% contort (7% of specimens). Toxas: typically very abundant (82%), common (3%), uncommon (5%), rare (3%), or absent entirely (7% of specimens); only symmetrical toxas present, with large central curvature and relatively straight points (52%), with both symmetrical and asymmetrical, sinuous toxas (41%), or with both these and wing-shaped toxas (7% of specimens). Variability in spicule dimensions: With few exceptions, and despite apparent morphological incongruence in many other features, spicule dimensions were surprisingly closely comparable between specimens despite that they were col- 300zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM lected from widely separated geographic localities. Results were not analysed statistically because bathymetric, habitat and season data were not available for the majority of material. However, scatterplot comparisons between samples found that for each spicule category there were only few anomolous specimens which consistently differed from typical forms of the species. These anomalies are discussed further below. REMARKS. Clathria (Thalysias) cactiformis is predominantly temperate. It is characterised by its typically flabellate or flabello-digitate growth form, a smooth surface with a detachable thick ectosomal peel which is usually paler than the choanosome, relatively thin spongin fibres heavily echinated but poorly cored, and especially its short, thick, squat acanthostyles with a bare neck, bare points and large recurved spines over the remainder of the spicule. There is considerable morphological variability between regional populations (see Variation, above), to some extent vindicating Hallmann's (1912) subdivision of the species into 'varieties' (subspecies), based mainly on differences in growth form. In fact some specimens examined were assigned to this species with hesitation given their diverse external morphologies, but no other consistent skeletal or other characters were found to correlate with these morphotypes. Nevertheless, this taxon is probably composed of several cryptic sibling species (morphospecies) which might only be reliably differentiated using biochemical, genetic and other non-skeletal data. NMVRN0436 and RN0551 from Port Phillip Bay, Vic. have acanthostyles which are long, evenly spined, spines are sparse and small; echinating spicules occur in heavy concentrations and are evenly dispersed throughout the choanosome, not predominant in the peripheral skeleton; ectosomal skeleton is very well developed and well differentiated from subectosomal plumose tracts; the subectosomal region is noticeably more cavernous than in most typical forms; the choanosome is distinctly plumoreticulate; mesohyl matrix is very heavy and fibres are heavily invested with spongin; the smaller category of isochelae and toxas are absent. In some respects these features correspond to Spongia appendiculata, perhaps justifying the retention of that species separately form cactiformis, but other general morphological features indicate that these specimens cannot be otherwise reliably separated. WAM133-82 from the Northwest Shelf of WA has consistently basally spined choanosomal styles, and isochelae are completely unmodified. The ectosome and subectosomal regions are poorly developed and incompletely differentiated, but the specimen is otherwise comparable with typical forms. Some further comment is necessary on the synonymy above. Spongia appendiculata is included here as a new synonym of C. (T) cacnformis (cf. Hooper & Wiedenmayer, 1994) with virtually identical growth form, spicule geometry and skeletal structure as typical forms of this species. However, there are some notable differences between these nominal species, especially in the markedly smaller dimensions of most spicules in appendiculata, acanthostyles have a vestigial granular spination, and they are also almost entirely incorporated into the fibres themselves (i.e., enveloped by collagen). These differences are not presently considered sufficient to recognise it as a distinct taxon. Incorporation of echinating acanthostyles into spongin fibres is also known for several other species of Clathria (termed here the `phorbastformis' species complex, including C. (T) phorbasifonnis, C. (D.) dura, C. (D.) myxilloides, and others). However, in Spongia appendiculata acanthostyles are inserted perpendicular to fibres, much like typical echinating megascleres, but they have 'sunk' into them, whereas in these other species acanthostyles run parallel to fibres and replace or supplement the usual coring spicules. Clathria acanthodes is considered a synonym of C. (T) cactifonnis, despite having slightly different acanthostyle morphology ('neck' often spined). The characters used by Hentschel (1911) to differentiate C. acanthodes from other species (i.e., similar geometry between principal and auxiliary spicules, presence or absence of microspines on bases of principal and auxiliary spicules, presence of two size classes of palmate isochelae, and possession of rhaphidiform toxas), are found in most other populations of C. (T) cactifonnis and are not considered to be of primary importance in differentiating species. Clathria typica var. porrecta, also described by Hentschel (1912) from Indonesia, is a synonym of Clathria (T) reinwardti not of C. (T.) cactiformis (this syonymy was overlooked by Hooper & Wiedenmayer, 1994). The type material of Clathria indurata and Echinonema anchoratwn var. dura from WA have a uni- or paucispicular core of choanosomal styles in secondary fibres, whereas typical populations of C. (T) cacnformis

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