MEMOIRS OF THE QUEENSLAND MUSEUM VOLUME 40 BRISBANE ] JULY 1996 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 CONTENTS INTRODUCTION smsa o bieti pa aa ta la e a A aas Rm pna 1 MATERIALS AND METHODS ............... asse a 2n 2 ABBREVIATIONS .......... sse a lr ee eee 3 ACKNOWLEDGEMENTS ........... 4s sh es 5 DEFINITION OF CHARACTERS ... 2... 2 0. ee 6 SYSTEMATICS: Aege mora dp ete ta a dit pose be ctae a de 19 GENERIC NAMES INCLUDED IN MICROCIONIDAE . ........ 005005. 21 DESCRIPTION OF AUSTRALIAN SPECIES... ..... 2l ees 88 Cisthtlá eJ. uuum ye cee ee Tu pee ee ee eS Sa a 88 Clathria (Clathria) t tao e i btaa hao epi te 88 OTHER SPECIES OF CLATHRIA (CLATHRIA) ...... 4... 171 TRANSFER otto A tte h fal al de teh bte Ot a dm ie a e ibm i] el de d d 178 Clathria (Wilsonell) |... ....... ee 179 OTHER SPECIES OF CLATHRIA (WILSONELLA) ........ 22s 202 Clathria (Microciona) ......... a 4 zs 202 OTHER SPECIES OF CLATHRIA (MICROCIONA) ...... 2... 22er 218 TRANSFERS eoeta ideoda aen aea pe eet yay hi ow eas tee ite qai 226 Clathria (Dendrocia) |... 1 ee ee d 227 Clathria (Axociella) .......... cars 247 OTHER SPECIES OF CLATHRIA (AXOCIELIA) ............--.2+-.- 267 Clathria (Isociella) 2... ee ee es 267 OTHER SPECIES OF CLATHRIA (ISOCIELLA) .....005 5052 50452-25280 Clathria (Thalysias) ...... ee ee ee ras 280 OTHER SPECIES OF CLATHRIA (THALYSIAS) .. 0.00... 00 0404 2 ee eee 408 TRANSFERS areh bats Se a pb 4 GE ba ed buoxaddees He bE opa 412 Antho (Antho) .......... reap rye 413 OTHER SPECIES OFANTHO(ANTHO) ......... e 420 Antho(Plocamia) ........... o ra 422 OTHER SPECIES OFANTHO (PLOCAMIA) ..,..... e mere 430 Antho (Isopenectya) ........ er emn 433 OTHER SPECIES OF ANTHO (ISOPENECTYA) ...... ene 439 Echinodathria .....:.........:2 24a hh mn 439 OTHER SPECIES OF ECHINOCLATHRIA ........ 2e 480 Holopsamma .............5:2»... rs a ras s se yim eos n 482 Echinochalina. ........... eaa ar ras 513 Echinochalina (Echinochalina) ............. lll ln 513 Echinochalina (Protophlitaspongia) ........... eere 543 Artemisina ......... csl RR mh RR rms 558 OTHER SPECIES OFARTEMISINA .. 06 0 esane 566 Pandaros ................ s sr sss. 968 DISCUSSION-, inrer uu abe rta ma e n la sb ee ru eh a Reino le lal feb ol 569 LITERATURECITED .......... or aa a ttn 591 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, noarlun- gae,biclathrata, borealis, burioni, sarai, saraspinifera). C. (Wilsonella) (6, 8, abrolhosensis, ensiae, lindgreni spp. nov.), C. (Microciona) (5, 91, illawarrae, lizardensis, simae, brondstedi, campecheae, claudei, hentscheli, leighensis, stephensae, tunisiae, urizae, vacelertia 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, phorbasifarmis, styloprothesis, tingens, wesselensis, amiranteiensis, hechteli spp. nov.); Antha (with 3 subgenera: A, (Antho) (12, 10), A. (Plecamia) (2, 17) and A, (Jsopenectya) (3,1, punicea, saintvincenti spp. nov.); Echinoclathria (14, 15, bergquistae, levii, notialis, parkeri, riddlei spp. nov.); Holopsamma (9 species endemic to Australia, | indeterminate species); Echinochalina (with 2 subgenera: E. (Echinochalina) (10, 2, felixi sp. nov.), E. (Protophlitasporigia) (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 exira-axial regions). Brief zoogeographical comparisons are made between con- tinental Australian and adjacent Indo-west Pacific faunas. Australian species comprise abou! 32% of the world’s microcionid diversity; about 75% of species are endemic for the Australian region, and temperate species (8156) have higher levels of endemism than tropical species (5956). [ ] Porifera, Demospongiae, Poecilosclerida, Microcionidae, family revision, new species, raxonory, biogeography, Australia. John N.A. Hooper, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101, Australia; received 1 December 1995, INTRODUCTION Microcionidae is one of the largest families of Demospongiae, comprising about &% of all described (extant) Porifera species (Hooper & Lévi, 19933). 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 arc particularly abundant, with some species being dominant components of the shallow water macrobenthos. Previous works describing this fauna (and other literature containing extra-limi- tal records of Australian species) include: Lamarck (1814, 1815, 1816), Gray (1858, 1867, 1869, 1870), Bowerbank (1864, 1875, 1877), Bar- nard (1879), Kent (1871), Ridley (18843), Ridley & Dendy (1886, 1887), Lendenfeld (1888, 18892), Kieschnick (1896, 1900), Thiele (1898, 1899, 1900, 1903), Kirkpatrick (19003, b), Whitelegge, (1901, 1902). Hentschel (1909, 1911, 1912), Li] Hallmann (1912, 191]4à-c, 1916a-c, 1920), Dendy & Frederick (1924), Topsent (1897b, 1930, 1932, 1933), Burton (19343), Bergquist & Tizard (1967), Kelly-Borges & Bergquist (1988), Bergquist & Fromont (1988), Wiedenmayer (1989), Hooper (1990b), Hooper et al. (1991, 1992), Hooper & Lévt (1993a, 1994). A brief synopsis of the fauna 3s given by Hooper & Wicdenmayer (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 com- posite (consisting of several sibling species), or synonyms of other species. This study 1) describes [48 species (31 new), many new locality records for Australia and new taxonomic combinations; 2) provides an annotated synonymy for 311 other species worldwide (in- cluding 21 new species); 3) revises the mor- phometric characters used for classification and population vartability for particular species; and 4) determines levels of endemism amongst provincial faunas. MATERIALS AND METHODS COLLECTION AND HISTOLOGICAL TECH- NIQUES. 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 choanosomul 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 per- pendicular 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 orcard to raise the coverslip level. Phenol-xylene precluded the MEMOIRS OF THE QUEENSLAND MUSEUM necessity for dehydration through an alcohol series. Some microtome sections cut at 30-35m were made for cach 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 lo the extenur so as to include both the outer surface and inner skeleton relatively intact}, placed in clearing agent for an adequate period La dissolve wax and/or dewaxing on a hot plate, then soaked in ethanol until perfectly clear, floated onto albumen-coated slides, orientated and flat- tened, stained with basic fuccsin and mounted, Fragments of dry specimens (e.g., type material) were reconstituted in 596 buffered formalin for 12 hours, which produced rehydration of the mesohyl and enabled cleaner histological sec- tions to be made. MORPHOMETRIC | ANALYSES. Spicules 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 lengihs refer to chord length; isochelae are measured from apex of alae; width measurements of other spicules refer 1o maximum width. Spicule dimensions were sorted and statistical- ly compared for various parameters (c. 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. Linc-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, morphurneltric and binary characters, uulising the DELTA sys- tem (Dallwitz & Paine, 1986), SCANNING ELECTRON MICROSCOPY, Scc- tions were prepared as follows: 1) Cutat 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 Fall- ing apart. Delicate structures (plumose, halichundrvid, hymedesmoid skeletons) only re- guired a few seconds treatment with bleach; robust skeletons (reticulate, fibrous, articulated REVISION OF MICROCIONIDAE 5 skeletons) required several minutes: but general- ly 30 seconds was adequate. 3) Bleach was pipetted off at the appropriate time and 70% ethanol immediately added. Sec- tions 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. con- centrated hydrogen peroxide in place of sodium hypochlonrite, finally rinsing in ethanol. The hydrogen peroxide step was omitted for very delicate sections. 5) Sections were placed an 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 cornplete- ly (usually several days), then prior 1o use ex- posed 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 à 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 he viewed successfully under low ac- celerator voltage, but better results were general- ly 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 ectoseme and choanosome were placed in a durham tube (micro-test Lube), to which drops of concentrated nitric acid are added, using drop-by-drop addition 80 as to control the oxidation reaction and produc- tion of by-product oxides. 2) Upon completion of acid digestion the dur- ham tube was half filled with acid and gently heated over an alcohol llame, ensuring that onl small bubbles form (low heat, no boiling), for I- 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 supematant between each change nf 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 ento 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 com- pound or dissecting microscope (magnification depending on spicule size}. More drops of spicule solution added and this step repeated if toa 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 Megascleres (Table 28). Principal spicules ab- sent. Auxiliary subtylostyles (coring fibres) long, slender, invariably straight. with prominent, smooth subtylote bases and fusiform points. 280 Ectosomal auxiliary subtylostyles significant- ly smaller than coring spicules but otherwise of identical geometry. Echinating spicules absent. Microscleres (Table 28). Palmate isochelae un- common, unmodified, with long lateral alae en- tirely 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, echinat- ing 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 af- finities 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 Isociella and Thalysias. Clathria (Isociella) skia was initially thought to be Am- philectus 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 sub- isodictyal skeletal structure, but re-examination of the holotype showed that it does have echinat- ing acanthostyles, and differentiated principal and auxiliary megascleres (thus more ap- propriately included in Clathria (Clathria); see above), and it also has substantially different fibre characteristics from C. (/.) skia indicating that they are not conspecific. OTHER SPECIES OF CLATHRIA (ISOCIELLA) Clathria (Isociella) incrustans (Bergquist, 1961) Isociella incrustans Bergquist, 196 1a: 42-43, text-figs 15a-b [Ahipara Bay, N. New Zealand; originally assigned to the Suberitidae, Hadromerida]. Bergquist & Fromont, 1988: 114-116, pl.53, figs e-f, pl.54, figs a-b; Dawson, 1993: 36. MATERIAL. HOLOTYPE: NMNZ unregistered. NZ. MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 28. Comparison between of Clathria (Isociel- la) skia sp.nov. Measurements in jum, denoted as range (and mean) of spicule length x spicule width (N=25). Holotype SPICULE (QMG300449) Choanosomal absent principal styles Auxiliary 162-(248.4)-368 x (coring) styles 2.5-(4.5)-8 Auxiliary 97-(111.9)-129 x (ectosomal) styles 1.5-(2.4)-4 Echinating Absent acanthostyles Chelae T 72-(104.4)-145 x 65-(81.1)-108 x Ox ^] 202404-08 | 02495)08 | Paratype (AMGS5043) absent 154-(242.1)-318 x 2-(4.1)-6 87-(110.2)-132 x 1.5-(2.4)-4 absent 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. Stylotellopsis Thiele, 1905: 456; de Laubenfels, 19362: 112. Colloclathria Dendy, 1922: 74. Damoseni de Laubenfels, 1936a: 110. DEFINITION. Specialised ectosomal skeleton composed of two size classes of auxiliary (sub- tylo)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 be- tween 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 (Thalysias) abietina (Lamarck, 1814) (Figs 138-141, Tables 29-30, Plate 4C-D) Spongia abietina Lamarck, 1814: 450, 377. Clathria abietina; de Laubenfels, 1954: 141-142, text- fig.90; Hooper & Wiedenmayer, 1994: 267. REVISION OF MICROCIONIDAE Clathria aculeata Ridley, 18842: 443-444, pl.40, fig.1, pl.42, fig.k; Ridley & Dendy, 1887: 147, 246, 254; Topsent, 1897b: 447; Burton, 1934a: 558; Burton, 1959a: 243; Lévi & Lévi, 1989: 80-81, pl.7, fig.3. Rhaphidophlus aculeatus; Topsent, 1932: 115, pl.4, fig.10; Vacelet & Vasseur, 1977: 114; Vacelet et al., 1976: 73-74. Rhaphidophlus cervicornis, in part; Vacelet & Vas- seur, 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?1T'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, NTMZ1943, NTMZ1958, NTMZ 1963, QMG303373, QMG303382, NTMZ2089, NTM- Z2161, 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, NTMZ- 1217, NTMZ 1244, NTMZ1272, NTMZ1314, NTMZ1411, NTMZ1423, WAM151-82, WAMIS5- 82 (fragments NTMZ1731, NTMZ1732), NTMZ1770, NTMZ1801, NTMZ1820, NTMZ1824, NTMZ 1852, NTMZ2272, NTMZ2329, NTMZ2349, NTMZ2486, NTMZ3017, NTMZ3030, NTMZ 3031, NTMZ3032, NTMZ3033, NTMZ3396, QMG300448 (NCIQ66C- 1517-P), QMG300117 (NCIQ66C1518-Q) (fragments NTMZ3488, NTMZ3489), PIBOC04-595 (fragment QMG30005 1) 281 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 as- sociated 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 (Lévi & Lévi, 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 ex- panded basal attachment, never rhizomous; branches rarely anastomosing, branching mostly irregular, bushy, occasionally regular growing in 1 plane, sometimes flagelliform, occasionally ex- panded, club-shaped; total length and branch diameter variable (50-460mm; 2-22mm, respec- tively). 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 al- cohol soluble; yellow pigmentation more stable in ethanol but easily scratched from surface of living sponge, leaving a maroon or mauve pig- mentation beneath; pigmentation extends into periphery of choanosomal mesohyl. Oscules. Exhalent pores usually very small, 0.6- 2.0mm diameter, barely visible optically, dis- persed on distal ends of branches surrounded by surface conules, occasionally scattered along en- tire 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 ex- tremities of branches, 2-6mm maximum basal diameter. Ectosome and subectosome. Very dense, con- tinuous palisade of small ectosomal auxiliary subtylostyles forming discrete brushes overlay- 282 50 um MEMOIRS OF THE QUEENSLAND MUSEUM = a FIG. 138. Clathria (Thalysias) abietina (Lamarck) (holotype MNHNDT634). A, Choanosomal principal sub- tylostyle. 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 auxiliary subtylostyles in peripheral skeleton (the latter often obscuring ectosomal brushes); subec- tosomal spicule tracts arise from ultimate choanosomal fibres, intermingled with tufts of principal choanosomal subtylostyles echinating peripheral fibres; principal spicules not extending into ectosomal skeleton; mesohyl matrix heavy and pigmented in both ectosomal and subec- tosomal regions. Choanosome. Skeleton irregularly reticulate; heavy spongin fibres, 50-125.m diameter, heavi- ly anastomosing, not divided into primary or secondary elements but forming irregular, close- meshed reticulation; fibre meshes oval to elipti- cal, 100-350um diameter; fibres cored by multispicular tracts of larger auxiliary subtylos- tyles occupying 60-80% of fibre diameter; choanosomal principal subtylostyle uncommon within fibres, mainly found at fibre nodes protruding through fibres individually or in bundles; fibres also echinated by acanthostyles especially on peripheral fibres; mesohyl matrix heavy but only lightly pigmented, slightly granular; choanocyte chambers oval or eliptical, REVISION OF MICROCIONIDAE TABLE 29, Comparison in range of spicule dimensions between present and published records of Clathria (Thalysias) abietina. All measurements are given in jum and denoted as length x width (N=25), [secue | 1 | 2 | | Subectoso mal} 157.343x5-10| 350x85 | 19-276x2-5 | 120.250x 2-3 127-387 x 1-20 | IRE 61-94 x 4-10 53-74x46 | 5070K78 | 4l-109 x 1-19 | Chelae I 11-15 10-14. Lum [ m» | 91-210 EMNENE DT634. ‘De Laubenfels’ (1954:14 ) material ial USNM 2 22808, 23090. on ia abietina - holot ria aculeata Ridley (1 4. Rhaphidophlus cenicas in part, (Vacelet & Vasseur, 1597 1:06]. | 5. Present material (N—25 40-170jum diameter; larger auxiliary styles also dispersed between fihres in disorganised tracts. Megascleres. Choanosomal principal styles-sub- tylostyles usually uncommon, short, stont, slight- ly curved at centre or near base, occasionally straight, with slightly subtylote bases or evenly rounded bases; spicules usually completely smooth, exceptionally with minutely microspined bases, tapering to abruptly (hastate) sharp points. Length 121-(202,7)-300.5pm, width 5414.4)-244m (lectotype 165-(215.5)- 258m x 9,5-(14.3)- 15.5 m). Subectosomal auxiliary subtylosytles straight or slightly curved near base, with prominent sub- tylote, usually microspined bases, tapenng to fusiform points. Length 127-(258.4)-386.91.m, width 1.1-(8.2)-20, jum (lectotype 157-(274.4)- 343um x 5-(6.7)-]0pm). Ectosomal auxiliary subtylostyles short, thin, invariably straight, with prominently subtylote. typically microspined bases, tapering to fusiform points. Length 44.8-(114.6)-230j m, width 1,1- (3.8)-10.1 um (lectotype 62-(85.5)-132 x 2-(3.4)- Spm). Echinating acanthostyles moderately long, thick, straight, with prominently subtylote, spined base, virtually aspinose ‘neck’ (proximal to base), and lightly spined shaft; spines relatively large. Length 40.6-(82, 4)-L09p.m, width 1.1- (7.5)-191.m (lectotype 61-(74.4)-94 x 4-(6.5)- Oum). Microscleres. Palmate isochelae with long lateral alae completely fused to shaft, shorter entire front alac; two size categories present, both relatively ahundant, both with examples of contort shafts {more abundant in smaller form). Length I: 7.9-(12.1)-19.9j. m (lec- [Ts — 'etype 11-(12,5)- L5 | 14 Sim), length Il- 0.5- (5.8)-1Oj.m (lectotype | 4-(5)-6j.m). Toxas relatively un- common with 2 | Fenner forms; (i) wing-shaped, short, | thin, generously curved at central with reflexed points; (ii) accolada toxas long, thin almost | straight with small an- | gular central curvature and straight points. Length I: 5-(24.9)- 10-15 75ym, width 1.8-(2.4)- 4.1m (lectotype 15-(30,8)-61 x 2,042.9}3.5pm). Length II: 58- (184.6}-345um, width 0.3-(0.8)-1.5um (lec- totype 65-(120,8)-180 x 0.5-(0.9)-1.2um), Larvae. Parenchymella larvae observed in peripheral choanosomal skeleton of about 20 specimens, predominantly during the tropical wel season (February-April) and less so during the pre-dry season (May-July). Larval incubation was distinctly seasonal and occurred more-or- less evenly throughout the population (i,e., ir- respective of depth of collection; Fig. 141). Larvae were 600-950um diameter, elongate-oval in shape and many contained larval megascleres; cilia were not observed (preserved material). Associations, 3% of specimens had filamentous algae coring fibres in addition to longer auxiliary megascleres. Variation. Highly variable in live colouration, non-fibre skeletal development, megasclere size. relatively consistent in growth form, surface fea- tures, fibre skeleton, spicule geometry, Gross morphology; stalked, bushy, branching in more than ] plane, branches bifurcate, occasionally anastomosing (46%), planar branching (33%). dendritic planar branching (fans) (14%), or single digits with no or few bifurcations (7% of specimens). Atypical growth forms (few surface conules, few branches, thin branching) found predominently in deeper offshore coastal waters (40m depth). Live colouration: highly variable, no particular pigment considered to be typical, ranging from (i) maroon, evenly Pigmentation (Munsell 5R4/10-2,5R4/8-10), (ii) red-marnon, even (SR5/8-10), (11) bright red-orange, even 284 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 139, Clathria (Thalysias) abietina (Lamarck) typical growth forms. A, Lectotype of C. aculeata BMNH1882,2.23.258, B, BMNH1887.5.2.110. C, Lectotype MNHNDT634. D, Paralectotype MNHNDT3343. E, Paralectotype MNHNDT3342, F-G, Deeper water specimens NTMZ1820, NTMZI314. H, Shallow-water specimen NTMZ68. (5R5/12), (iv) bright orange, even (IORS/I2- 10R6/10-12-10R7/10), (v) orange-brown, even (10R4/8-10), (vi) orange-yellow, even (7.5YR7/12), (vii) pale brown-pink, even (10R7/4), (viii) light brown, with pink conules (10R7/A, 5R8/4), (ix) light brown, with dark grey- brown conules (2.5 YR7/4-5 YR7/2-6, 2.5YR6/6- 5YR3-5/2), (x) pale brown, even (SYR6/6-5YR7/4-6-5YR8/4), (xi) muddy grey, even (7.5 YR7/2-8/2) (xii) yellowish grey, even (2.5Y8/6), (xii) yellow, even (2.5 Y7/8), (xiv) yellow, with pink conules (2.5Y8/6, 10R7/6), (xv) lime-yellow, with dark brown conules (2.5Y7/10, 2.5Y 5/2), (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 side- by-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 MICROCIONIDAE 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. SPICULE Mrs PE (1950) 4.44 P«0.005 (775) 0.26 10.61 (1950) 189 | Poos | 40 | Poo PEN stylesL (1950) 6.21 P «0.001 Ectosomal stylesL (1950) (1950) Acanthostyles L Chelae I L (1950) (1950) (1925) Chelae II L (1925) (1925) 0.39 P>0.05 Number of groups: 1. 3 locality groups s (NWS, DAR, CP localities). 2. 4 depth groups (0-4m, 4-10m, 10-40m, 40m (MA, 3. 4 seasonal groups (Darwin region only: wet branching) having peripheral choanosomal fibres lying immediately below ectosomal crust, thin paratangential subectosomal region, and acan- thostyles echinating peripheral fibres piercing ec- tosomal skeleton. Development of extra-fibre skeleton: 4% with very abundant juvenile auxiliary subtylostyles distributed throughout mesohyl, 9096 with at least some interstitial auxiliary spicules, and 6% of specimens without any interstitial auxiliary spicules dispersed be- tween 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 thin- ner, 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 2.72 P<0.05 a925) | 042 549 | P<0.001 P>0.05 (775) 1.67 P>0.05 ESEA P<0.0005 (775) 2.46 (775) 2.08 P>0.05 (750) (750) (12% of specimens). Microsclere geometry: Proportion of contort to normal morphs of pal- mate 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 entire- ly. 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 col- lected 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 dis- tributions 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- 286 MEMOIRS OF THE QUEENSLAND MUSEUM i88um FIG. 140. Clathria (Thalysias) abietina (Lamarck) (A-B, NTMZ2642; C-1, 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 MICROCIONIDAE SAMPLES WITH | LARVAE | | | SEASON TOTAL SAMPLES [wer [| s /—— | PREDRY FIG. 141. Clarhria (Thalvsias) abietina (Lamarck) In- cidence of incubated larvae within sampled popula- tions from NW Australia. possible ta separate the influences of either fac- tors on this observed variability. Variability of spicule size between samples collected during different seasons were analysed for Darwin samples only (Table 30), with significant dif- ferences observed for lengths of ectosomal auxiliary subtylostyles and larger isochelae, and widths of acanthostyles. Data also indicate a higher level of vanability 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 considerahle morphological variability C. (T:) abietina is easi- ly recognised in the field with distinctive stalked digitate growth form and prominent surface con- wes. The most similar species in growth form is C. (2) cervicornis but this has a much thinner, stoloniferous branching morphology, lacks prin- cipal megascleres completely, fibres have less spongin, toxas are different in morphology and size, and spicule dimensions are different. Skele- tal 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 ec- tosomal skeleton: 2) dispersed between fibres 1n 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, includ- ing a higher proportion of choanosomal principal spicules found echinating fibres, the absence of contort isochelae, and specific details in some 287 spicule dimensions (Table 29). But given the large range of variation in some characters, the relatively large sample sizes from widely dis- persed populations, and the antiquity of the dried holotype this variability is insignificant. Re-ex- amination of Ridley's (1884a) holotype of C. aculeata confirmed that it is a synonym of C. (7.) abietina, with shape, texture, colour in spirit, spiculation and skeletal architecture virtually identical. Conversely, C. (T.) coralliophila (sec below) has different spicule geo and skele- tal 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 3n 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 al- most completely covered by subectosomal spicule brushes (as opposed to ectosomal spicule brushes). In fact de Laubenfels (1954) completely overlooked the presence of ectosomal ascleres, not differentiating between spicules coring fihres from those forming the peripheral skeleton. Clathria (Thalysias) aphylla sp. nov. (Figs 142-143, Plate 5D) MATERIAL. HOLOTYPE - QMG300477 (NCIQ- 66C-A640-K): NW. of E. Passage, Easter Group, Hout- man Abrolhos, WA, 28740" S, L13*50' E, 20m depth. 17,1x.1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION, Staghorn and plate coral fringing reef; 20m depth; known only frorn the type locality, Houtman Abrolhos (WA) (Fig. 142H). DESCRIPTION. Shape. Thin, leaf-like, foliaceous. convoluted, basically frondose bundles of lamellae covering coral substrate; in- dividual fronds attached directly to substrate, sometimes completely enveloping staghorn corals, or attached via small basal stalk, up to 24mm long, Smm diameter, or attached to ad- jacent lamellae; individual lamellae usually flat, elongate, oval oreliptical, up to fmm long, 2mm thick. with rounded or sinuous, convoluted mar- gins, 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 2mm diameter, flush with surface. 288 Texture and surface characteristics. Soft, com- pressible 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 paratangen- tial brushes of ectosomal auxiliary styles forming thin surface layer, with choanosomal principal styles protruding through ectosome perpen- dicular to surface, singly or several together, ex- tending up to 150um through surface; choanosomal fibres immediately below ec- tosome, 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 pig- mented, Choanosome. Regularly reticulate, slightly renieroid skeleton; without compression or major differences between peripheral skeleton and core; spongin fibres heavy, 20-70j.m diameter, not ob- viously 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 con- fined 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), 90-170u.m diameter, with some triangular meshes, relatively even throughout skeleton; mesohyl matrix light, with few scattered megascleres; choanocyte chambers oval to eliptical, 25-40j.m diameter, lined by abundant, minute isochelae. Megascleres. Choanosomal principal styles long, thick, straight or slightly curved towards basal end, rounded or slightly tapering bases, invariab- ly smooth bases, fusiform points. Length 152- (205.1)-252jum, width 4.5-(8.4)-11 jum. Subectosomal auxiliary styles long, slender, straight, with rounded or faintly subtylote smooth bases, fusiform points. Length 208-(258.1)- 297 um, width 2-(2.7)-4.5 jum. Ectosomal auxiliary styles short, slender, straight, rounded or slightly subtylote bases, smooth or microspined (or mucronate) bases, fusiform points. Length 162-(178.2)-204j1.m, width 1-(1.6)-2.5j.m. MEMOIRS OF THE QUEENSLAND MUSEUM 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)-102j.m, width 3- (5.5)-8p.m. Microscleres. Palmate isochelae very abundant, minute, with lateral alae completely fused to shaft and partially fused to front ala. Length 4-(5.1)- 6jum. 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)-106,.m, width 0.8-(1.3)-2.5jum. 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 (main- ly 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 dif- ferences 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, ves- tigial fibres with light spongin, cored by much smaller principal spicules in uni- or paucispicular tracts); a specialised ectosomal skeleton com- posed of two size classes of auxiliary spicules forming brushes and principal spicules protrud- ing 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 dif- ferentiated 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 MICROCIONIDAE 50 um 289 V TE jui / J. f -f 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 ectosornal skeleton. (from Torres Strait and Shark Bay), which has only one category of auxiliary spicule and dif- ferent ectosomal structure, wing-shaped toxas, different acanthostyle geometry and specific dif- ferences in spicule sizes. All three species probably have sister species relationship based on skeletal architecture, here referred to the ‘angulifera’ group. A few other Clathria species also have V- shaped 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) Rhaphidaphlus arborescens Ridley, 1884a: 450-451, pl.40, fig.L, pl.42, figs n-n'; Thiele, 1903a: 958. MEMOIRS OF THE QUEENSLAND MUSEUM 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 MICROCIONIDAE Clathria arborescens; Hooper & Wicdenmayer, 1994: 268. cf, Microciona prolifera tropus spinosa; Vosmaer, 19352: 610, 634, MATERIAL. HOLOTYPE - BMNH1881.10.21.272: Friday J., Torres Strait, Qld, 10°36°S, 142°1D°E, Ly.1881, call. HMS ‘Alert’ (dredge). HABITAT DISTRIBUTION, Ecology unknown: Tor- res Strait, Qld (Fig. 144). DESCRIPTION. Shape. Branching, arborescent, 125mm long, 60mm maximum width of branch- ing, 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 subectasomal striations, ir- regular conules and low ridges. Ectosome and subectosame. Ectosomal skeleton with very thick crust of discrete spicule brushes composed of ectosomal auxiliary subtylostvles, occasionally paratangential or tangential to sur- face, supported below hy thick, vaguely plumnse, slightly disorganised bundles of larger subec- tosomal auxiliary subtylostyles arising from ler- minal choanosomal fibres; subectosomal skeleton lacking any spongin fibres and only light mesohyl matrix; together ectosomal and subec- tosomal bundles extend up to 800um from peripheral fibre skeleton, and comprise up to 25% of branch diameter. Choanosome. Choanosarmal skeleton irregularly close-meshed reticulate. Spongin fibres very large, heavy dark yellow-brown, 110-170um diameter, forming tight oval or elongate meshes, 130-350j.m diameter: fibre reticulation without any clear distinction between primary or secon- dary components. although ascending fibres mar- ginally more abundantly echinated than transverse connecting fibres, and some smaller fibres between major meshes with uni- or paucispicular core; fibres generally all cored hy 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 291 relatively large acanthostyles, more-or-less even- ly dispersed throughout skeleton (not con- gregated in periphery as reported by Ridley); choanocyte chambers not observed (dry material). Megascleres. Choanosomal principal subtylos- tyles with similar geometry to larger auxiliary spicules, long, straight. usually with subtylote, smooth or microspined bases, rarely rounded bases, with tapering fusifonn points. Length 205- (225.3)-262um, width 8-(9.7}+-12um. 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)-282um, width 4-($.4)-7pm_ Ectosomal auxiliary subtylostyles relatively long, straight or slightly curved towards base, prominently subtylote, with smooth or microspined bases, fusiform points, Length 82- (132.7)- 147j.m, width 2-(3.3)-45.m. Echinating acanthostyles long, robust, sub- tylote, large conical (not recurved) spines. scut- tered mainly over base and midsection of shafi, frequently with bare point and ‘neck’, Length 68-(76.3)-88um, width 6-48.2)-1] um. Microscleres, Palmate isochelae small, frequent- ly contort, single size category, with long lateral alae completely fused to shaft and entire front ala of equal length. Length 6-(10.8)-I4j2m. Toxas short, robust, wing-shaped, with generous angular central curvature, recurved and slightly reflexed points Length 27-(41.8)-5Rj.m. width 1,5-(1,8)-2.0j.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 (18842). These include: presence of dit- ferentiated principal and auxiliary spicules, presence of toxas, spinatiom pattern on acanthos- tyles, and his spicule dimensions were innac- curate, These characters now provide better clues as to rts 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.) arbares- cens has a substantially better developed ec- tosomal peel occupying a significant proportion of branch diameter, a close-meshed reticulate 292 MEMOIRS OF THE QUEENSLAND MUSEUM 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:) cactiformis has a relative- ly thin ectosomal skeleton, cavernous subec- tosomal region, wide-meshed choanosomal reticulation, substantially thinner fibres with only primary ones cored). Also, echinating acanthos- tyles are much larger than those of cactiformis, they have erect conical spines rather than hook- like 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 marked- ly between the two, with arborescens having only thicker, wing-shaped toxas, and cactiformis with mainly very thin, raphidiform, sinuous or ac- colada toxas but occasionally also with small wing-shaped forms. Spicule dimensions also dif- fer considerably between the two species. Both species belong to the ‘juniperina’ species com- plex (refer to discussion below under C. (T.) cactiformis and C. (T.) hirsuta). Dendy (1922), Burton & Rao (1932) and Bur- ton (19382) 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 MICROCIONIDAE t3 Mel [ov] 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. 294 at least eight other species (see below under C. cactiformis)). Thiele (1903a) also compared C. ( T.) arborescens and C. (T) cervicomis, for the same reason (1.e.. supposed absence of principal spicules), but C. procera does have principal spicules (exclusively echinating peripheral flores). and those in C. (T.) arborescens are inter- mingled with auxiliary subtylostyles within fibres. Skeletal structure and forms alsa differ considerably between these lwo taxa. Clathria (Thalysias) cactiformis (Lamarck, 1814) (Figs 146-148, Table 31, Plate 4E-F) Spongia cactiformis Lamarck, 1814: 440, 370 Clathria cactiformis: Rudman & Avern, 1989- 335; Hooper & Wicdenmayer, 1994: 268. Spongia appendiculata, in part, Lamarck, 1814: 383, 2 Wilsonella appendiculata, Topsent, 1930: 46, pl2, fig.3; Topsent, 1933: 22. Echinonema typicum Carter, 1875; 195 [nomen nudum]; Carter, 1881: 362, 377, 388, 384. Thalassedendron typica; Lendenfeld, 1898. 223; Hallmann, 1912; 242; Hallmann, 19143; 267. Not Thalassodendron typica; Whitelegge; 1901: 86. Clathria typicu; Vosmaer, 1885b: 357; Ridley & Dendy, 1887: 153; Dendy, 1889c: LI;]in part] Dendy, 1896: 32; Whitelegge, 1901: 80, 117; Hentschel, 1911: 280, 364-367, text-fig. 43. Not Ciarhrio typica var. porrecta Hentschel, 1912: 298, 359-150. Not Clathria rypica; Kirkpatrick, 1903: 248-249; Kirkpuiek: 1904: 148; Vacelet & Vasseur, 197; 34-95, pl.3, fig.5, text-Fip. 48. Wilsonella typica; Hallmann, 1912: 203, 242. Not Tenavia typica var. porrecta Hallinann, 1920: 771. Rhaphidophlus typicus: Hallmann, 1912: 176-177, 184-203, 234, 299, 300, pl.27, pL 28, figs 1-4, pl.29, lext-figs 38-42 [et varr]; Shaw, 1927: 424-425; Top- sent, 1932: 88, pL4, fig.4; Burton, 19383: 12: Guiler, 1950: 8; Vacelet & Vasseur, 1977; 115; Vacelet et al., 1976: 73; Liaaen-Jensen et al., 1982- 170-172. Echinonema anchoratum Carter, 188 1a- 362, 379, 380: Lendenfeld, 1888: 219-220, Echinonema anchoratum var. dura Whitelegge, 1901: 81 Echinonema gnchoratwm var. lomellosa; Lendenfeld, [888: 219-220. Rhaphidophius oanchoratum; Bergquist & Fromon, 1988: 112; Dawson, 1993: 39. Wileonella ancheratum var. lameflosr; Hallmann, 1912: 211 [imperfectly known]. Not Echinonema — anchora Whitelegge, 1901: 82. Echinonema flabelliformis Carter, 1885f: 352. Evhinonema pectiniformis Carter, 1885f; 353, Fa ventilabrum var -australiensis Carter, 1886g: var. Jamellosa; MEMOIRS OF THE QUEENSLAND MUSEUM Thalassodendron brevispina Lendenteld, 1888: 225; Whitelegge, 1901: 87, Clathria favosa Whitelegge, 1907: 498-499, Clathria indurata Hallmann, 1912: 299. Clathria acanthodes Hentschel, 19) 1: 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. ct. Microciona pralifera; Vosmaer, 1935a: 609, 628- 629. 647, 667. MATERIAL. LECTOTYPE: MNHNDTSS0O: "Southern Seas’, Peron & Lesucur collection, no other details known. PARALECTOTY PE: MNHNDT3360: same details (= R. npicus var. cactiformis, Topsent, 1932; 88). LECTOTYPE of S. appendiculata; MNHNDT526 (fragment BMNH1953.4.9.83) (dry). PARALECTOTYPE of S, appendiculata: MNHNDT- 3394: King George Sound (Albany), WA. LEC- TOTYPE of T, brevispina: AMZ931; Port Jackson, NSW, 33*48'8. 131720'E, coll. R. von Lendenfeld (dredge). SYNTYPE of E. typicum: BMNH- 1877.5.21.149; Fremantle, WA, 32703 S, L15°38"E, coll. E. Clifton (dredge). HOLOTYPE of C. favesu: AMZD944: Off Port Jackson, NSW, 33*50'S, 151°30 B, 98-100m depth, coll. FIV ‘Thetis’ (dredge), LEC- TOTYPE of R. typicus var. proximus: AMZ930: Hen- ley Beach, St. Vincent Gulf, SA, 34°45°S, 137°S7°E, coll. T. Worsnop (dredge). PARATYPE of R. rypicus var. obesus: AMZ937: Tuggerah Beach, NSW, 33°18°S, 151°30°E, coll. NSW Fish Commission (trawl), HOLOTYPE of R, typicus var. geminus: AMZ9S28: specific locality unknown, WA, no olher details known, HOLOTYPE of &, typicus var. stellifer. AME?648: E, coast of Flinders 1, Bass Strait, Tus, 40*01'5, 148702'E, no other details known. SYN: TYPE of E. ancharanum: BMNH L1886.12.15.423: Port Phillip, Vic, 38°09'S, 144°52°E, no olher 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: BMNH- 1886.12.15.141 (fragment AMG2777): Port Phillip. Vic, 38*09' 8, 144752" E, 40m depth, no other details known. Fragment of HOLOTYPE C. ucanthodez: ZMB4A443; 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 - AMER24, AMZI415, AMZ2203, QMG311412 (NCIQG6GC- 3713-A) (fragment NTMZ3811). VICTORIA - AMZIA30, AMZIISS, NMV RN359, NMVRN43], NMVRN436, NMVRN 438, NMVRN551, NMVRN- 677, NMVRN797, NMVRN8&40, NMVRND9((, NMVRN959, AMZ3921 (RRIMPFN3527/000/02), AMZA271 (RRIMPFN 1906), NTMZ2832. S AUST - AMZAI, SAM unreg. (fragment NTMZ1657), SAM unreg. (fragment NTMZ 1693), SAM unreg. (fragment NTMZI628), SW COAST, WA -MNHN unregistered REVISION OF MICROCIONIDAE 25um 100 um FIG. 146. Clathria (Thalysias) cactiformis (Lamarck) (lectotype MNHNDT580). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthos- tyles. E, Smuous 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), QMG304660, AMZ4331 (RRIMP-ZW62), WAM639-81(1), WAM661-81(), WAM660-81, QMG300736 (NCIQ66C-4189-T), QMG300733 (NCIQ66C-4197-C), QMG300730 (NCIQ66C-4214-V). NORTHWEST SHELF, WA - WAM133-82, NTMZ1128, NTMZ3018, NCIQ66C- 1460-C (fragment NTMZ3347). ERITREA, REDSEA - PIBOCB20-125 (fragment QMG300055). SOMALIA, W. INDIAN OCEAN - PIBOCB12-388 (fragment QMG300060). SEYCHELLES - PIBOCOI- 140 (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 m 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 (sec 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. Shape. Typically flabeliate, 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, vary- ing from globular, cylindrical planar digitate, globular digitate, planar flabellate, lamellate branching in more than one plane, or thinly lamellate. Colour, Liye colouration pale orange-brown (Munsell SYR8/4) to bright red (5R5/10); beige, 296 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 147. Clathria (Thalysias) cactiformis (Lamarck) typical growth forms. A, Lectotype MNHNDT580. B, Paralectotype MNHNDT3360, C, Lectotype of S. appendiculata MNHNDTS526. D, Paralectotype of S. appendiculata MNHNDT3394, E-F, Variety cactiformis WAM656-81(1), WAM655-81(1). G, Variety stel- lifera NTMZ1708. H, Variety geminus WAM639-81(1). 1, Somalia specimen PIBOCB 12-388. J, Syntype of E. typicum BMNH1877.5.21.149. K, QMG300453. L,Variety obesus with Rostanga feeding (photo W. Rudman) NTMZ2832, REVISION OF MICROCIONIDAE 297 J 166um 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. 298 TABLE 31. Comparison between present and publish- ed records of Clathria (Thalysias) cactiformis (Lamarck), All measurements are given in jum, denoted as range, or as range (and mean) of spicule length x spicule width (N=25). = = i Choanosomal| i94419L2i- | joo ig), | 103.6(199.2)- | principal 232x 5-(7.6)- 1926 314x3.2- Mytes u i 9.7)-19.4 Subectosomal| 2044229.) | 1912977 | 1570357 auxiliary 268 x 4-(5.4)-6 455 356.6x L6- | styles H m (6.2)-16.8 Ectosomal 101-(124.3)- 73,8-(114,7)- auxiliary 148% L5- | 85-104 x 2-35 | 162.1 x 1.6- styles (25)3 4.4)-13.6 38.2-(61.3)- Echinating | 68-(74.4)-79% | y 4 ucanthostyles| 5.5-(8.8)-11 | 03/3345 | 93.1817 (6.7)-13.1 244078 | — T8 /— [654139)222 Cheer | esmo | m | 250-0 | 11342017295 | 82-1462 05- | 8-127.8)-355 | xü3412k2 | LO — [2035-13-25 Tortas brown, to dark brown in ethanol; ectosomal crust usually lighter colour than choanosomal regions. Oscules. Oscules typically large (up to 3mm dia- meter}, dispersed over entire sponge surface, or sometimes clumped 1n special pore areas; oscules usually flush with surface, without raised lip. Texiure and surface characteristics. Firm, com- pressible, rubbery; surface characteristically fleshy, smooth but uneven, with bumps, ridges, depressions and subdermal stnations commonly dispersed over surface; opaque skin-like covering present stretched between adjacent conules in live material; microscopic ridges and microconules abundant. Ectosome and subectosome. Ectosomal skeleton predominantly heavy, with a dense crust of smaller auxiliary subtylostyles forming mainly erect or less often paratangential brushes: oc- casionally ectosomal skeleton reduced to rela- lively thin tangential or paratangential crust; larger subectosomal auxiliary styles perched on ultimate choanosomal fibre endings, forming plumose tracts underlaying brushes of smaller ectosomal spicules, usually protruding through and obscuring ectosomal bundles; subectosomal region cavernous, with well differentiated, wide- ly spaced plumose tracts of larger auxiliary sub- tylostyles, Choanosome. Skeleton irregularly reticulate, vaguely radial, with clearly differentiated primary ascending and secondary transverse MEMOIRS OF THE QUEENSLAND MUSEUM skeletal components; fibre anastomoses produce widely spaced rectangular meshes formed by as- cending primary fibres, containing pauci- or mul- tispicular bundles of choanosomal principal styles which occupy only the central portion of fibres, and uncored secondary fibres; occasionál- ly secondary fibres uni- or paucispicular, usually with an optically diffuse pith reminiscent of dic- tyoceratid sponges; spongin fibres charac- leristically thin, without marked differences between primary and secondary fibre diameter, with pigmentation ranging from extremely light, yellow (specimens from Port Phillip, Vic) to ex- tremely heavy and dark brown (material from Port Hedland, WA); echinating acanthostyles very heavy, especially on peripheral fibres imine- diately below subectosomal skeleton; larger auxiliary subtylostyles usually abundant belween fibres, strewn without order within mesohyl; mesohyl matrix relatively light, choanoc yte chambers large, oval or elongate, up to 120m diameter. Megascleres (Table 31). Choanosomal principal styles morphologically similar to larger auxiliary subtylostyles but relatively shorter, thicker, slightly curved near base, usually with tapering, evenly rounded smooth bases but sometimes slightly subtylote, and fusiform points. Subectosomal auxiliary subtylostyles nearly completely straight, occasionally slightly curved or sinuous, usually with slightly subtylote smooth bases, less frequently with microspined bases, tapering to fusiform points. Ectosomal auxiliary subtylostyles more prominently subtylote, slightly curved ar centre, with microspined or smooth bases and fusiform points. Echinaung acanthostyles relatively short, sicut, prominently subtylote, with large spines mainly dispersed on base and midsection, leaving a bare ‘neck’ (below basal swelling) and bare point. Micrescleres (Table 31). Larger palmate isochelae variable in abundance, with lateral alae completely fused to shaft, wide gap between lateral and front alae, and fused front ala; usually non-contort. Smaller isochelae common, fre- quently contort. Toxas differentiated into two main morphs, larger often found in toxodragmala, accolada toxas most common, long, very thin, slight an- gular central curvature, straight arms, sometimes sinuous asymmetrical, Small toxas less abundant, relatively thick, widely curved at centre, slightly reflexed points. REVISION OF MICROCIONIDAE Larvae. Reproductive products observed in only four specimens. Incubated parenchymella larvac large, oval to eliptical in shape, 220-420j.m diameter, containing well developed centrally ap- gregated larval styles. Associations. Heavy infestations of zoophytic or- ganisms observed on surfaces of several specimens (both compound ascidians and zooan- thid polyps); numerous microalgal cells seen in sponge mesohyl of few samples; sponge ap- parently serves as a food source lo Restanga 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 pee) reminiscent of Crella (Crellidae) (14% of specimens), Subec- tosomal skeletal structure: very cavernous, well differentiated ascending plumese (extra-fibre) skeletal tracts (36%), paratangential, immedi- ately subdermal tracts (21%), or intermediate of these conditions (43% of specimens). Choanosomal skeletal structure: irregularly reticulate (64%), regular fibre anastomoses (2296), or predominantly radial to plumose primary fibre architecture (14% of specimens), Fibre meshes: wide, rectangular, with uncored secondary elements (67%), with central pith com- ponent (7%), close-meshed, irregulariy oval to eliptical fibre reticulation with uncored secon- dary fibres (796), or with both cared 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: con- centraled on peripheral fibres, more sparsely dipersed on axial fibres (69%), evenly distributed throughout fibres (17%), rare (14% of specimens), Extra-fibre spicules: typically abun- dant, dispersed without order within mesohyl (74%), very light or absent entirely (24%). nr organised into ascending extra-fibre tracts (2% of specimens). Megasclere geometry: Choanosomal principal styles: predominantly fusiform (7696). distinctly hastate (abruptly pointed) points (7%), or mixture of both types of terminations (1746 of 299 specimens); bases of spicules predominantly smooth (90%), with both smooth and spined bases (895), or only microspined bases on choanosormal styles (2% of specirnens); bases of Spicules tapering, non-tylote (46%), rounded. slightly subtylote (525€), or with both conditions (2% of specimens): megascleres slightly curved (83%), exclusively straight (722), or both condi- tions (10% of specimens). Subectosomal auxiliary subtylostyles: tapering, rounded or very slightly subtylote bases (6945), or distinctly sub- tylote (31% of specimens); exclusively straight (69%), predominantly curved (7%), or both con- ditions (2496 of samples). Ectosomal auxiliary subtylastyles> suhtylote (76%), tapering non- ivlote (10%) er both conditions (14% of specimens); bases predominantly microspined (719), exclusively smooth (3%), or with both condiltons (26% of specimens); ectosomal spicules slightly curved near base (54%), com- pletely straight (299%), or with exaniples of both (17% of specimens); most megascleres with fusiform points (95%), or also including few has- iate spicules (5% of specimens). Echinating acan- thostyles: 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 churactenstically long, thick, recurved (749%), or reduced, granular spanation (26% of specimens). Microsclere geometry; Larger isochelae; common (70%), uncommon (26%), rare (296), orcommon but poorly silicified (2% of specimens); contort larger isachelae present but uncommon (10%), present, common (2%), or absent (86% of specimens). Smaller isachelac: abundant (756), common (62%), un- common (21%), rare (396) or absent entirely (7% of specimens); 0-10% of smaller chelae were canarr (2%), 10-3096 contort (596), 30-50% con- tort (129), 50-70% contort (249%), 70-90% con- tort (43%), or (00% contort (7% of specimens). Toxas: typically very abundant (82%), common (3%), uncommon (5%), rare (396), or absent eh- tirely (7% of specimens); only symmetrical toxas present, with large central curvature and relative- ly straight points (52%). with both symmetrical and asymmetrical, sinuous toxas (41%), or with both these and wing-shaped toxas (796 of specimens). Variability in spicule dimensions: With few cx- ceptions, and despite apparent morphological in- congruence in many other features, spicule dimensions were surprisingly closely comparable between specimens despite that they were col- 300 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 characiensed hy 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 heavi- ly echinated but poorly cored, and especially its short. thick, squat acanthostyles with à 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 ex- lent 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 exter- nal 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 (morphespecies) which might only be reliably differentiated using biochemical, genetic and other non-skeletal data. NMVRN(Ü436 and RNOSS1 from Port Phillip Bay, Vic. have acanthostyles which are long, evenly spined, spines are sparse and small; echinating spicules occur tn 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 subes- tosomal plumose tracts; the subectosomal region is noticeably more cavernous than im most typical forms; the choanosome is distinctly plumo- reticulate; mesohyl matrix is very heavy and fibres are heavily invested with spongin; the smaller category of isochelac and toxas are ab- sent. In some respects these features correspond io Spongia appendiculata, perhaps justifying the retention of that species separately form cactifor- mis, but other general morphological features indicate that these specimens cannot be otherwise reliably separated. MEMOIRS OF THE QUEENSLAND MUSEUM WAM133-82 from the Northwest Shelf of WA has consistently basally spined choanosomal styles, and isochelae are completely unmodified. The ectosome and subectosomal regions arc poorly developed and incompletely differen- tiated, but the specimen is otherwise comparable with typical forms. Some further comment is necessary on the synonymy above. Spongia appendiculata is in- cluded here as a new synonym of C. (7:) cactifor- nus (cl. Hooper & Wiedenmayer, 1994) with virtually identical growth form, spicule geometry and skeletal structure as typical forms of this species. However, there are some notable dif- ferences between these nominal species, espe- cially 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 them- selves (1.e,, enveloped hy collagen). These dif- ferences dre 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 "phorbasiformis' species con- plex, including C. (T.) phorbasiformis, C. (D.) dura, C. (D.) myxilloides, and others), However, in Spongia appendiculata acanthoslyles are in» serted perpendicular to fibres, much like typical echinating megascleres, but they have 'sunk' into them, whereas in these other species acanthos- tyles run parallel to fibres and replace or supple- ment the usual coring spicules. Clathria acanthodes is considered à synonym of C, (T) cactiformis, despite having slightly different acanthostyle morphology ('neck' often spined). The characters used by Hentschel (1911) to differentiate C. acanthodes from other species (1e. 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 rhaphidif'orm toxas), arc found in most other populations of C. (7) cactiformis and are mot considered to be of primary importance in differentiating species. Clathria typica var. porrecta, also described by Hentschel (1912) from Indonesia, is à synonym of Clathria (T. ) reinwardtii rot of C. (1) cactifer- inis (this syonymy was overlooked by Hooper & Wiedenmayer, 1994), The type material of Clathria indurata and Echinonema anchoratum var. dura from WA have a uni- or paucispicular core of choanosomal styles in secondary fibres, whereas typical populations of C. (T. ) cactiformis REVISION OF MICROCIONIDAE generally have an aspicular secondary fibre sys- fem, but otherwise spiculation and architecture is similar between these morphs and they are ob- vious synonyms. The species 1s included in the juniperina com- plex (see discussion of C. (T) hirsuta) all of which have reduced skeletal and fibre charac- teristics. This group includes 12 species (with about 20 synonyms): C. (T. arborescens, C_(T.) caciiformis, C. (T.) cervicornis, C. (T.) clathrata, C. (1) corneolia, C. (T.) craspedia sp. nov., C. (C.) decumbens, C. (T.) fusterna sp. noy., C. (7:) hirsuta, C. (T.) juniperina, C. (T) rubra and C. (T.) placenta. Many of these species barely differ in their gross morphologically, and Hallmann (1912) suggested they should be combined into one or few species, but this contention cannot be resolved without access to live populations of all species, to determine accurately their field char- acteristics, nor on the basis of preserved material. Clathria (Thalysias) cancellaria (Lamarck, 1814) (Figs 149-150, Plate 5A) Spongia cancellaria Lamarck, 1814: 382, 361. Rtiapiidophlus cancellarius, Topsent, 1930: 43, pl.2, 1g.6. Clathria cancellaria; Hooper & Wiedenmayer, 1994: 270, MATERIAL. HOLOTYPE: MNHNDT528: locality unknown, Peron & Lesueur collection, OTHER MATERIAL: WA - NTMZI249 (fragment QMG300430), QMG300594 (NCIQ66C-4272-K), QMG300536 (NCIOG6C-4667-0). HABITAT DISTRIBUTION. Rock substrate, on or under ledges, or exposed on isolated rock in sand substrate; 13-40m depth; known only from Australia: Northwest Shelf, Wallaby Is and Houtman Abrolhos (WA) (Fig, 14914). DESCRIPTION. Shape. Lamellate, massive, mostly planar growth form, with or without basal attachment, consisting of more-or-less fused erect thin digits forming a contmuous lamella, 370mm maximum span, with some free or barely anastomosing sections; lamellae are 116-135mm high, 4.5-7mm thick, forming irregular meshes between branch anastomoses, 3-14mm diameter. Colour, Salmon-pink (Munsell SRP 8/2) to orange-red alive (5 YR 6/10), ight grey-brown in ethanol. Oscules. Small, 2-3mm diameter, mainly be- tween ridges; pores minute, 0.5- Imm diameter, scattered over entire surface, Texture and surface characteristics, Firm, com- pressible, difficult to tear. Lamellae prominently 3I striated with longitudinal ridges, grooves and dis- continuous conules; margins of lamellae digitate, microconulose. Ectosome and subectosome, Relatively thin, dense ectasomal crust, 60-95 ym thick, composed of erect ectosomal auxiliary subtylostyles form- ing continuous erect dermal palisade; immedi- ately below ectosome, protruding through palisade, are mostly paratangential subectosoriyal auxiliary sublylostyles; in thicker, more elon- gated sections larger auxiliary subtylostyles form plumose ascending tracis supporting ectosonial skeleton, but in most sections they lie paratangen- tial to surface, arising from ends of peripheral choanesomal fibres; subectosomal region rela- tively cavernous; in growing points of sections in peripheral skeleton elongate conulose protrusions extend from surface for up to 0.5mm, fully cored by dense tracts of subectosomal sub- tylostyles, Choanosome, Skeletal architecture irregularly reticulate, without axial or extra-axial differentia- tion; spongin fibres relatively heavy, imperfectly separated imlo primary and secondary elements, forming ovoid or elongate, relatively cavernous. meshes, 140-680j.m diameter; primary fibres, 92-165um diameter, predominantly ascending, cored by multispicular tracts of choanosomal principal styles occupying about 75% of fibre diameter; secondary fibres uni- to paucispicular, 26-71m diameter, predominantly transverse; all libres heavily echinated by acanthostyles protrude at various obtuse angles from fibres; mesohyl matix heavy but only lightly pigmented, slightly granular, with abundant larger auxiliary megascleres throughout, sometimes completely obscuring skeletal architecture; choanocyte chambers, small, oval, 42-68pum diameter. Megascleres. Choanosomal principal styles short, thick, slightly curved towards basal end, with rounded or very slightly subtylote smooth bases, fusiform points. Length 166-{185.5)- 218pm, width 7.5-(12.6)-l6pm (holotype 161- (198.1)-224 x 6-(12.1)- 16pm). Subectosomal auxiliary subtylostyles long, thick, straight or very slightly curved, with slight- ly suhtylote or rounded bases, usually microspined, fusiform points. Length 134.6- (218.8)-2864.m, width 3.5-(6.5)-8.6.m (holotype 148-(229.7)-265 x 3-(5.9)-Bpum). Ectosomal auxiliary subtylostyles short, thin, straight, with prominent. subtylote bases, in- variably microspined, fusiform points. Length 86-(97.6)-114,5j. m, width 2.5-(3.3)-4.5jum (holotype 85-(93,45- 102 x 2«(3.T)-5j&m). MEMOIRS OF THE QUEENSLAND MUSEUM 302 E SN 100 um ai cS Dm ER Ñ N J FIG. 149. Clathria (Thalysias) cancellaria (Lamarck) (holotype MNHNDT528). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Accolada and sinuous toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. J, NTMZ1249. REVISION OF MICROCIONIDAE FIG. 150. Clathria (Thalysias) cancellaria (Lamarck) (NTMZ1249), A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of subectosomal and ectosomal auxiliary subtylostyles. G-H, Palmate isochelae. I, Accolada toxas. 304 Acanthostyles. long, thick, with slightly sub- tylote rounded bases, large spines dispersed over apical and basal extremities, usually with uspinose neck, occasionally evenly spinose; spines conical, barely recurved. Length 67- (79.8)-92 5um, width 3.8-(8.6)-11.64m (holotype 62-(80.2)-88 x 3-(8.1)-LOj.m), Microscleres. Palmate isochelae divided into two size classes, larger usually unmodified, smaller often contort; long lateral alae completely fused to shaft; front alae complete. Length T: 9-(10.9)- I6y.m (holotype 9-(12.3)-l6pm), length II; 3- (4.4)-8.m (holotype 3-(4.5}-6.m). Toxas thin, rhaphidiform, accolada to sinuous. varying from small forms with slightly curved centres and slightly reflexed points, to long forms with only very slightly rounded central curvature and straight or slightly reflexed points. Length 11-(121.1)-l664m, width 0.4-(0.8)-T.2,m (holotype 17-(102.3)-186 x 0.5-(1.1}-1 5pm). REMARKS. Clathria (Thalysias) cancellaria is most closely related to the New Caledonian C. (T) flabellifera, both with similar skeletal ar- chitecture, spicule geometry and spicule size, and also (vaguely) growth farm, surface features and live colouration (Hooper & Lévi. 19933). The two populations differ in several cryptic characters. which Hooper & Lévi (1993a) suggested were indicative of sibling species relationships rather than population variability. Clathria (T.) cancel- laria has only a single category of toxa mucrosclere, resembling neither of the two found in C. (T.) flabellifera; t has two size categories of isochelae microsclere (whereas the latter species has only one); acanthostyle spines are much larger, the spicules more robust, and spines are more evenly dispersed over the spicule (whereas those on the latter species are small and found mainly towards the extremities of the spicule): basal spines are present on most of the smaller and larger auxiliary spicules (whereas most auxiliary spicules are completely smooth in the latter species); and spicules of most categories arc substantially thicker in the WA population. Clathria (Thalysias) cancellaria also superti- cially resembles C. (T) raphana and iregular rowth morphs of C. (T) coppingeri, with lamel- ale. planar growth form, although fibre charac- teristics, spicule geometry and spicule sizes are otherwise different between all these species. The species has a dense crust of erect ectosomal spicules, very unlike the pauctspicular dermal skeleton of C. (7) coppingeri, although both species conform to the Thalysias condition in MEMOIRS OF THE QUEENSLAND MUSEUM having differentiated ectosomal and subec- tosomal megascleres. Clathria (Thalysias) cervicornis (Thiele, 1903) (Figs 151-152, Table 32, Plate 5B) Rhaphidophlus cervicornis Thiele,1903a: 959, 968, p1.28, fig.24a-e; Hallmann, 1912: 177; Brondsted, 1934: 22-23, fig.23; Simpson, 19682: 70, pl.15. Not Rhaphidophlus cervicornis, Vacelet & Vasseur, 1971: 96-97, text-fig.50, pl.4, fig.3. Thalysias cervicernis, de Laubenfels, 1954: 135-137, text-fig. 86. Clathria cervicornis; Bergquist, 1965: 165-167, fig. 14. MATERIAL. HOLOTYPE: SMF679 (fragments ZMB3141, MNHNDCL2312) Ternate, Moluccas, In- donesia, 0°48'N, 127?23'E, 1894, coll. W. Kükenthal (dredge). OTHER MATERIAL: QLD - QMG300707 (fragment NTMZ4045). CAROLINE IS., CENTRAL W PACIFIC - USNM22892, USNM22905, QMG304828 (NCI OCDN-0488-O). HABITAT DISTRIBUTION. Usually at base of coral reef slope. on coral rubble, rock or seagrass beds, forming tangled thickets; intertidal - 45m depth; Howick ls (FNQ), (Fig. 151G), Aru Is, Arafura Sea, Moluccas Sea, Marshall Js, Palau Is, Truk Atoll (Chuuk), Ponape. DESCRIPTION, Shape, Stoloniferous, thin, cylindrical branches, 10-25mm diameter forming single digits. erect oc creeping over substrate. to dense tangled thickets with numerous, thin, sioloniferous, anastomosing, cylindrical branches, and cither small, central, single basal stalk, or multiple points of attachment to substrate via branches, Colour. Pale orange or orange-red exterior (Mun- sell 7.5YR 7/10-2.5 YR 7/8), bright vermillion or brown interior in life (2.5R 4/2-4/8); grey-brown in ethanol. Oscules. Large, up to 8mm diameter in life, with large membraneous lip, collapsing when preserved, dispersed mainly on lateral surface of branches in regular rows. Texture and surface characteristics. Firm. flexible, compressible surface, wiry axis difficult to tear, surface usually optically smooth, even, with subectosomal ridges and drainage canals radiating away from raised oscules, occasionally sparsely conulose, microscopically hispid in life in thicker branches; surface even. glabrous, unor- namented when preserved, Ectosome and subectosome. Ectosomal skeleton a dense palisade nf discrete, erect spicule brushes composed of smaller ectosomal auxiliary sub- tylostyles, supported by underlying, ascending, REVISION OF MICROCIONIDAE 305 AR 1 FIG. 151. Clathria (Thalysias) cervicornis (Thiele) (holotype SMF679). A, Subectosomal auxiliary subtylostyles and bases. B, Ectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Ectosomal and subectosomal skeletons (QMG300707) I, QMG300707. 306 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 152. Clathria (Thalysias) cervicornis (Thiele) (QMG300707), specimen with reduced spiculation. A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Polytylote bases of subectosomal auxiliary subtylostyles. F, Wing-shaped toxa. G, Palmate isochelae. REVISION OF MICROCIONIDAE 307 TABLE 32. Comparison between spicule dimensions of Clathria (Thalysias) cervicornis, from present material and published records. All measurements given in jum, as range, or range (and mean), of lengths x widths. Principal megascleres Absint Absent Ectosomal 112-0028.3)-1355 B6-(117.3)-151 105-(118.3)-135 subtylostyles £143.95 x2(3.2)4 x 3441-5 mane as anie 95-32 3465 38-(62 0)-74 52-(55.6)-61 47-(56.2)-63 og ( -82 x 23-5: -73x 4-6, | Acsumostyles x1454)7 x4.63)9 x344.25 60-82 x 23.5.9 50-73 x 4-6,5 | Chelae 1 10-(12.4)-15 | 10411.6414 8-13.6 Chelae lf 3(5.3)-8 3-(4.4)-6 abseni Nov recorded Nor recorded 12-(35.3)-55 16-(54.5)-112 | | - ^ | ) $ x05409-13 | x10412-20 Al “x Mm plumose tracts of larger subectosomal auxiliary subtylostyles, identical to those coring fibres which also protrude through surface; no fibre component in peripheral skeleton but plumose tracts in subectosomal region arise directly from terminations of ascending primary fibres; together ectosomal and subectosomal skeletons occupy up to 30% nf branch diameter. Choanosome. Skeleton regularly reticulate, with even, rectangular or ovoid, relatively cavernous meshes, 180-270j.m diameter; spongin fibres very heavy, 90-125 jum diameter, thicker at nodes, up to 2001.m diameter, cored by multispicular tracts of subectosomal auxiliary subtylostyles which occupy only 60% of fibre diameter; echinating acanthostyles moderate to lightly dis- persed over fibres, predominantly clustered around fibre nodes. sometimes rare or absent completely in some specimens; mesohyl matrix moderately light, with numerous subectosamal auxiliary megascleres scattered between fibres; choanocyte chambers small, oval, 35-55pm diameter. Megascleres (Table 32), Choanosomal principal megascleres absent or completely undifferen- tiated from subectosomal spicules. Subectosomal auxiliary subtylostyles long, slender or robust, straight or slightly curved near point, with prominently subtylote, palytylote slightly subtylote, or rarely rounded bases, smooth or minutely spined, fusiform points. Ectosomal auxiliary subtylostyles relatively large, robust, straight, with prominently subtylote bases, usually microspined occasionally smooth, fusiform points. Echinating acanthostyles moderately short, robust or slender, subtylote, with smooth point mE NEM - ae En - - i Holotype (N23) (N=1) (N=) (de (N-1) (Bergquist, | SPICULE (SMF679) (N=1) (GBR) (Micronesia) (Brondsted, 1934) | Laubenfels, 1954) | 1965; Simpson, (Indonesia) ARS (ArL,ndonesis)| — (Marshall Is.) Absent 180-4258.9)-304 | 232-(274.9).225 | 205-(236.7)-262 subtylostyles x 4-(5.8)-9 x4-(4.7-7 x 341.3)-6 ]9682) (Palau Is.) | abone | abet Absent Absent 165-315 » 6-10 135-325 x 2.4-7.6 and 'neck' below basal swelling: spines moderately large, recurved. Micrascieres (Table 32). Palmate isochelae occur in Iwo size classes, wih some smaller contort forms; lateral alae long, completely attached to shaft, front ala entirely fused. Toxas wing-shaped, long or short, thin, with large central curvature, slightly reflexed points, REMARKS. This widely distributed Indo-west Pacific species is easily recognisable in the field by its growth form, resembling prolific tangles oi thinly hranching thickets. In the western Pacific and Indonesian archipelago this species is a prominent member of the fringing coral recf com- munity, commonly found at the base af reef slopes in the rubble and sediment, whereas on the Great Barier Reef it has been found only it the far northern sector where itis rare and occurs as isolated thin single branches. The species has consistent skeletal architecture, probably dictated by its persistent cylindrical growth form, and similar spicule dimensions (Table 32), notwithstanding its relatively widespread Indo-west Pacific distribution, al- though there ts some variability in spicule geometries. Specimens from Chuuk luck microscleres and often have very thin megascleres, whereas those in the Great Barrier Reef specimen there is only sparsely echinating acanthostyles. Brondsted's (1934) material rs reported to have differentiated principal and auxiliary megascleres —the former with rounded smooth bases coring fibres and the latter with slightly subtylote smooth bases and found ex- clusively in the ectosomal skeleton or scattered between fibres. Similarly, acanthostyles in 308 Brondsted's material are evenly spinose, whereas in the holotype these have aspinose ‘necks’ and points. It is possible that Brondsted's material represents a different species, or it has simply been misdescribed, but no formal diagnosis can be made until his specimens are found and these differences confirmed or refuted. The presence of a second, smaller category of isochela, and frequent contortion of isochelae have not been recorded previously for this species, although these features were commonly seen in most material examined, including the holotype, and appear to have been overlooked by Thiele (19032) and subsequent authors. Conse- quently, there is no justification in separating Moluccan populations from others described by Bergquist, Brondsted, de Laubenfels and Simpson, whereas material described by Vacelet & Vasseur (1971) as Rhaphidophlus cervicornis belong to C. (T.) abietina. Clathria (Thalysias) cervicornis is closely re- lated to C. (T.) corneolia Hooper & Lévi (from New Caledonia), C. (T.) craspedia sp. nov. (from the Tweed River region) and C. (T:) fusterna sp. nov. (from the Gulf of Carpentaria), all having similarities in their skeletal structure and diver- sity of spicule types, but with major differences in their growth forms, live colouration and live surface features, some also in their spicule geometries (the latter three species with differen- tiated principal and auxiliary megascleres), and spicule dimensions. These are discussed further below for the respective species (and see also Hooper & Lévi, 1993a). Clathria (Thalysias) coppingeri Ridley, 1884 (Figs 153-154, Tables 33-34, Plate 6A) Spongia juniperina, in part (variety beta only); Lamarck, 1814: 444. Not Spongia juniperina (variety alpha); Lamarck, 1814: 444, Clathria coppingeri Ridley, 1884a: 445-446, pl.40, figs f-P, pl.42, figs i-i’; Fristedt, 1887: 459; Hallmann, 1912: 215; Hentschel, 1912: 298, 358, 361-362; Topsent, 1932: 99, pl.5, fig.1; Hooper, 1984a: 55; Hooper & Wiedenmayer, 1994: 270. Thalysias coppingeri; de Laubenfels, 1936a: 105. bs lendenfeldi, Brondsted, 1934: 19-20, text- ig.9. Not Clathria coppingeri var. aculeata; Hentschel, 1912: 363. cf. Microciona prolifera, Vosmaer, 1935a: 610, 633. MATERIAL. LECTOTYPE: BMNH1881.10. 21.246 (dry): Albany I., N. Qld, 10?44'S, 142°37°E, 6-8m depth, coll. HMS ‘Alert’ (dredge). PARALEC- TOTYPE: BMNH1881.10, 21.330 (spirit): same MEMOIRS OF THE QUEENSLAND MUSEUM locality. SYNTYPES of var. thuyaeformis: MNHNDTS71, DT3353: precise locality unknown, Indian Ocean, Turgot collection, no other details known (dry). OTHER MATERIAL: QLD - QMG4731 (fragment NTMZ1557). WA - NMV unregistered (fragment NTMZ1493). WA - NTMZ670, NTMZ1 152, NTMZ1861, NTMZ1173, NTMZ1155, NTMZ 1221, NTMZ2269, NTMZ2283, NTMZ2301, NTMZ2311, NTMZ2316, NTMZ2317, NTMZ 2363, NTMZ2463, NTMZ3040. INDONESIA - SMF1702 (fragment MNHNDCL2325), SMF1265 (fragment MNHNDCL2251). HABITAT DISTRIBUTION. Deeper offshore rock reefs, dead coral, coral heads, probably restricted to harder substrates (as indicated by presence of smooth encrusting basal attachment (peduncle); growth form consistently planar and likely that orientation of fan is towards direction of predominent currents; 15-94m depth; Port Headland region, Bedout I., Lacepede Is, Amphinome Shoals, Barracouta Shoals, Northwest Shelf; Ashmore Reef, Sahul Shelf (WA); off Moreton Bay (SEQ) (Ridley, 1884a; Hooper, 1984a; present study) (Fig. 153H); also Aru and Kai Is, Indonesia (Hentschel, 1912; Brondsted, 1934), DESCRIPTION. Shape. Growth form invariably thin, planar reticulate flabellate, 150-460mm long, 185-290mm maximum span, with short thick, cylindrical stalk (22-54mm long, 12-30mm diameter) and small peduncle; branching planar- dendritic reticulate, with ascending radial primary branches and interconnecting secondary branches; branches form tight meshes (4-9mm diameter); primary branch diameter 12-16mm, 6.5-9.5mm midway, 1.5-4mm near apex of fan; connecting branches 1.5-3mm diameter. Colour. Consistent, unpigmented, light beige- brown in both life and ethanol (Munsell 2.5Y 8/2-5Y 8/4). Oscules. Not observed in live or preserved material. Texture and surface characteristics. Firm, barely compressible, particularly on woody basal stalk; surface optically smooth, without conules or other visible processes, little flesh; microscopi- cally hispid, irregular, consisting of tight reticula- tion of compacted fibres and protruding megascleres. Ectosome and subectosome. Poorly developed ectosomal skeleton, without obvious spicule brushes but with sparse paratangential smaller ectosomal auxiliary subtylostyles scattered over surface; ascending plumose choanosomal prin- cipal styles usually protrude through ectosomal skeleton with bases embedded in peripheral spon- gin fibres, especially at fibre junctions and on fibre endings; detritus also often present on sur- REVISION OF MICROCIONIDAE 309 FIG. 153. Clathria (Thalysias) coppingeri Ridley (NTMZ670). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Accolada toxa. F, Palmate isochelae. G, Section through peripheral skeleton. H, Known Australian distribution, I, NTMZ3040. J, Unregistered live specimen. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 154. Clathria (Thalysias) coppingeri Ridley (QMG300106). A, Choanosomal skeleton. B, Fibre charac- teristics, C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of principal and auxiliary subtylostyles. G, Palmate isochelae. H, Accolada toxas. REVISION OF MICROCIONIDAE 311 TABLE 33. Comparison between present and published records of Clathria (Thalysias) coppingeri. All measurements given in jum, as range, or range (and mean), of lengths x widths (N=25). Holotype S. juniperina var. beta (MNHNDTS71) 179-(254.1)-332 x 200-340 x 15 9.62)21 134-(196.5)-279 x 1.6-(4.6)-8 Holotype (Ridley, 18842) SPICULE Choanosomal principal styles Subectosomal auxiliary styles 150-250x 5-63 Specimens (N22) (Indonesia) (Hentschel,1912; Brondsted,1934) Specimens (N=16) (Indian Ocean, Arafura Sea) Specimen (N=1) (Pacific Ocean) 150-(228.4)-318 x 3.5-(13.6)-26 136-(202.2)-323 x 1-(4.9)-12 164-340 x 10-20 201-520 x 4-24 164-362 x 1-9 Ectosomal 75-(108.8)-133 x auxiliary styles 1.1-(3.1)-5 Echinating 66-(99.5)-118 x acanthostyles 10x63 1.1-(6.3)-10 TAE L0504)25 face, sometimes replacing dermal skeleton en- tirely; subectosomal peripheral skeleton not sharply delineated from deeper choanosomal regions with choanosomal fibres usually imme- diately subdermal; meshes formed by fibre anas- tomoses often wider in peripheral skeleton. Choanosome. Skeletal architecture irregularly reticulate, without differentiation between axial or extra-axial regions; spongin fibres heavy, 40- 110p.m diameter, without any clear division be- tween primary or secondary elements; fibres form oval or rectangular meshes, 100-220m diameter in axis, 280-405p.m diameter near periphery; fibres uncored, usually heavily echinated by both acanthostyles and principal spicules especially at fibre junctions; very few fibres have unispicular core of subectosomal auxiliary megascleres; extra-fibre spicule development minimal, where present consisting of plumose ascending tracts of subectosomal auxiliary subtylostyles; mesohyl matrix light and collagen found mostly around fibre nodes, with few microscleres and sometimes detritus scat- tered throughout. Megascleres (Table 33). Principal choanosomal subtylostyles long, thick, slightly curved near the basal end, with slightly subtylote microspined bases, fusiform points; occasionally completely smooth styles, sometimes both lightly spined shafts and bases intermediate between choanosomal spicules and acanthostyles (both echinating fibres). Subectosomal auxiliary subtylostyles straight, thin, slightly subtylote microspined bases, oc- casionally smooth, fusiform points. 69-(110.8)-142 x 1-(3.7)-7 144-256 x 4-6 61-(97.3)-150 x SEHU 1.2-(6.5)-14 6-(13.1)-25 28-(140.6)-245 x ST 72-149 x 2-10 Ectosomal auxiliary subtylostyles smaller, thinner than subectosomals, invariably with microspined bases. Acanthostyles variable in size, charac- teristically curved at centre, lightly spined, spines small, recurved more-or-less evenly dispersed, bases slightly subtylote, points hastate or rounded. Microscleres (Table 33). Palmate isochelae moderately common, relatively small, never con- tort, with lateral and front alae of equal size, lateral alae completely fused to shaft, front ala entire, sometimes alae vestigial reduced to a ridge on shaft. Toxas accolada, very common in some specimens, rare in others, mostly thin, rhaphidiform, long, with slight central curvature, tapering arms, little apical reflexion, less often short and curved. Associations. Scyllidae polychaete worms (Typosyllis spongicola) abundant in 60% of specimens examined, identical to commensals seen in C. (T.) reinwardti and C. (T.) lendenfeldi. Variation. Growth form and colour relatively consistent. Growth form: consistently planar, stalked, with more-or-less tightly anastomosing branches, although two morphs recognised; typi- cal morph (6496 of specimens, including type material) with thicker ascending almost dendritic (primary) branches radiating outwards to produce an arborescent appearance; second morph (3646 of specimens, including type material of S. juniperina) have even branching, even branch sizes and mesh sizes closely resemble Echinodic- tyum cancellatum (Raspailiidae). Foreign detritus in skeleton: abundant in choanosomal mesohyl of deeper water specimens (70 m depth), 312 Tare in samples from shallower habitats (38-46 m depth). Ectosoma! skeleton: occasionally well developed, thick, typical of Thalysias condition (but also including principal spicules protruding through ectosome) (13% of specimens), more sparsely developed (20%), variable ectosomal development (well formed in some regions, such as surface irregularities. completely absent from other regions) (75€), or consistently poorly developed (60%). Subectosomal skeleton: poorly developed with peripheral choanosomal fibres lying immediately below the surface and thin paratangential spicule tracts (87% of specimens), or well formed (wide) plumose extra-fibre skeleton (13%). Choanosomal skeleton: fibres heavy (67% of specimens), or much lighter in construction (33%), Coring spicules: irregularly reticulate thick spongin fibres largely uncored (81% of specimens). more regularly renieroid- reticulate with thicker fibres cored by paucispicular tracts of choanosomal styles (12%), or fibres mostly cored by choanosomal styles (7%). Fibre diameter: even, consistent throughout skeleton (8156 of specimens), with distinct primary (ascending) and secondary (transverse) fibre elements (1256), or noticeably heavier fibres in the axis than an the peripheral skeleton (7%), Echinating spicules: moderately heavy acanthostyles and choanosamal styles forming dense plumose tufts at fibre nodes. producing ascending extra-fibre tracts extending into peripheral skeleton (80% of specimens). or with more poorly developed echinating spicules seemingly dispersed al random throughout skeleton (2096). Mesohyl matrix: light with cal- lagen found mainly around fibre nodes (93% of specimens), nr heavy, granular (7%). Megasclere gcometry: Principal spicules predominantly basally spined, but with variable proportion of entirely smooth spicules in specimens, ranging from 0-10% of spicules sampled (47% of specimens), 16-30% (27%), up to 56% of spicules (6% of specimens). Larger auxiliary subtylos- tyles usually with microspined bases, and only 0-10% of spicules sampled (73% of specimens) were smooth, 12-20% with higher proportion basally spined (20%), up to 32% of spicules (7% of specimens). Smaller auxiliary spicules mostly common in histological preparations, although exceptionally producing well-formed dermal structures (93% of specimens), or scarse in both Sections and spicule preparations. Microscleres: Isochelae abundant (26% of specimens), com- mon (33%), rare (21%), or absent entirely (2085). MEMOIRS OF THE QUEENSLAND MUSEUM Toxas: abundant (26% of specimens), common (2096), rare (47%) or absent (6%). Variability in spicule dimensions: Discounting the anomolous southern Queensland specimen (QMG4731) discussed further below, spicule dimensions were relatively consistent throughout the species’ geographical distribution (Table 33). For all categories of megascleres, spicule length was on average higher for the Queensland specimen than samples from the northwest and west coast, whereas the mean width of most megascleres, and the length of isochelae were relatively more homogeneous between all samples. Toxas were also significantly shorter in the southem Queensland specimen. Analysed by locality, variation in spicule lengths and widths did not conform to any obvious latitudinal gradient in sample distribution (the three major sampling localities for this species were 19°, 16°and 12°S Jatitudes) for the west coast matenal (P>0.05), for all spicule types, although the in- clusion of Queensland material (27°S) in analyses did reveal significant differences (com- monly at P«0.005) in all spicule categories except isochelae (Table 34), Excluding Queensland material (collected at 94 m depth). there were no significant differences in mean spicule lengths ur widths between specimens collected from 38- 46m or 70-90m depth ranges (P2005) for all spicule categories except acanthostyles. REMARKS. This species has à characteristic planar, flabellate growth form similar to C. (C.) loveni Fristedt (1887: 459) (from North America) and C, (C.) ulmus Vosmaer (1880: 151; 1935a: 633) (from an unknown locality). Previous descriptions omitted to mention thin, raphidiform toxas or presence of two size categories of auxiliary megascleres (Table 33), These were seen in all type and recent material, and posses- sion of two size categories of auxiliary spicules places the species in the Clashria (Thalysias) groupalthough itis atypical of most other species in having only rudimentary ectosomal structure. with sparse spicule brushes. This species belongs lo the spicata complex (Hallmann, 1912; Hooper et al., 1990) based on skeletal architecture (virtual absence of coring megascleres in fibres, inclusion of choanosomal principal spicules echinating fibres, imperfect differentiation of principal and auxiliary megascleres, the semi-plumose (or spicate) arran- gement of choanosamal megascleres protruding through fibres, and dense echination of peripheral fibres). Included in this group are: C, (T.) lenden- REVISION OF MICROCIONIDAE 313 TABLE 34. Summary of results of one-way ANOV's (Model I), testing for variability in spicule lengths and widths between locality and bathymetric distribu- tion of Clathria ( Thalysias) coppingeri. Lem as Lr 3s m ESL (350) Ectosom — les L ra ie [| va tate points (whereas in typical material they are fusiform); ectosomal spicules occur in light | Prob, | Paratangential bundles near the surface (versus not forming brushes); | oos f fibres are lightly cored ETC E, paucispicular tracts of choanosomal styles, | Poos | which produce an ir- regular renteroid reticulation (whereas most specimens have [fum | men mb fibres]; acun- thostyles are only spar- sely spined, and | Poos | (350) 037 P20.05 082 P20.05 — L (275) 0.69 P20.05 0.02 P20.05 Toxas L 1148 P«().0005 (335) together wrth | mos | Choanosomal styles, al- ways form ascending extra-fibre tracts wilhin 0.03 | Poo | 45 P24,05 Number of groups: L 4 locality groups (27, 19, 16, 12°S latitude) . 2 depth groups (38-46m, 70-90m depth feldi, C. (T.) major, C. (C.) caelata, C. (C.) in- anchorata, C. (T) clathrata, C. (T.) costifera (in- cluding the nominate species Clathria spicata, C. bispinosa, C. whiteleggei and Ophlitaspongia membranacea, which have since been shown to be conspecific with other species of the spicata group: Hooper et al., 1990)). This assemblage of species does not appear to constitute a natural taxon because it cuts across a classification based on other (possibly more important) characters (viz. Clathria and Thalysias). Nevertheless, all species are very close in fibre construction and spicule geometry, and in fact C. (T.) coppingeri and C. (T.) lendenfeldi can only be easily differen- tiated on the basis of their respective growth forms, which is very characteristic for the former, but relatively variable in the latter species (see Hooper et al., 1990). The single known specimen from southern Queensland differs in many respects from other populations, accounting for most of the variability documented above. In shape it is close to Lamarck's (1814) variety thuyaeformis (Top- sent, 1932: pL5, fig.1), whereas most samples examined resemble type material (Fig. 1531-I). This specimen also has a well developed ec- tosomal skeleton (consisting mainly of larger subectosomal megascleres which surround the bases of protruding choanosomal styles): true (smaller) ectosomal auxiliary spicules have has- the skeleton. This specimen was initially assigned to C. (T) coj pingeri witb some hesitation, but there is no doubt that it belongs to this species and probably represents the southem- most extent of its geographical range. There is a nomenclatural complexity thut re- quires brief comment. Wiedenmayer (1989) sug- gesied that the senior ‘varicty’ name nf S. Juniperina (viz. thuyaeformis) had priority over Ridley's (18842) subsequent species name, but this is not accepted here. Lamarck (1814 described several distinct "varieties! of 5. juniperina, tor which he used Greek symbols bur not a trinomen. It was Topsent (1932) who sub- sequently elevated Lamarck's "variety. B' to a subspecific rank by using it in a trinomen (ICZN Article 45fii), using the name thuyaeformis us a noun whereas it was previously used as an adjec- tive by Lamarck. This subsequent designation is therefore considered to be infrasubspecilic, not suhspecific (ICZN, Article 45f), and does not take precedence over Ridley's (1884a) species desig- nation. Further support of this opinion is that S. juniperina thayaeformis is a composite taxen as some of Lamarck's syntypes belong to different species. Thus the choice of the name coppiageri over thuyaeformis is also supported by ICZN Article 57g, and the pragmatic argument that Ridley's (1884a) name has now become well known for this species. 3i4 Clathria (Thalysias) coralliophila (Thiele, 1903) (Figs 155-156, Table 35, Plate 5C) Rhaphidophlas corailiophilus Thiele, 19033: 959, 968, pl.28, fig.25a-d. Rhaphidophlus coralliophita, Hallmann, 1912: 177. Tenacia coralliophila, Burton, 19348- 360. Clathria coralliophila; Hooper & Wiedenmayer, 1994- 270 cf. Microciona prolifera, Vosmacr, 19353: 611, 645. MATERIAL. LECTOTYPE: SMF1784 (fragment ZMB3145): Temate. Moluccas, Indonesia, 0°48"N, 127?23'E, 1894, coll. W. Kükenthal (dredge). PARALECTOTYPE: SMF787 (fragment MNHNDCL2309): same locality. OTHER MATERIAL: QLD - BMNH1930.8.13.107. PNG - QMG300377 (NCIQ66C-4518-A]). HABITAT DISTRIBUTION. Growing on on Seriotepora coral, bivalves and dead coral substrates; shallow intertidal 10 8m depth: Low Is (NEQ) (Fig. 155Hx; also Moluccas, Indonesia; Matupore T., S. PNG. DESCRIPTION. Shape. Thin, thick, or massive- Ly encrusting, covering up to 80mm* surface. Colour, Red, dark orange or pale pinkish alive (Munsell SR 8/2 - IOR 6/10), grey in ethanol. Oscules. Large. up to 4mm diameter, scattered over surface, slightly raised above surface with membraneous lip. Texture and surface characteristics. Firm, com- pressible; surface even, smooth, fleshy, without obvious sculpturing, microscopically hispid. Ectosome and subectosome, Well developed, dense, continuous palisade of ectosomal auxiliary spicule brushes in 1 or more plumose layers (several consecutive brushes of spicules overlaying one another), ectosomal skeleton sup- ported by well developed series of discrete sub- ectosomal plumose brushes, the latter not proteuding through ectosome; subectosomal region cavernous with numerous plumose brushes of larger auxiliary subtylostyles. Choanosome. Skeletal structure ranges from thin basal layer of spongin on substrate (hymedes- mid), to large, erect, non-anastomosing, single fibre nodes arising from encrusting basal layer of spongin (microcionid); basal (hymedesmoid) fibres very heavily echinated by erect choanosomal principal styles and acanthostyles; digitate (microcionid) fibre nodes cored by erect mulüspicular tracts of choanosomal principal spicules, congregated especially on ends of fibre nodes, forming plumose brushes and producing a series of ascending plumose oroccasionally anas- tomosing tracts; subectosornal tracts mostly per- MEMOIRS OF THE QUEENSLAND MUSEUM pendicular, less commonly longitudinal in thin- ner sections, always plumose; echinaling acan- thostyles concentrated primarily in basal part of skeleton, rarely near surface, forming very dense erect layers on basal (hymedesmoid) fibres; mesohyl matrix moderately heavy, granular, abundant subectosomal auxiliary styles dispersed between choanosomal spicules; choanocyte chambers small, oval or elongate. 120-2504 m diameter. Megascleres (Table 35). Choanosomal principal styles long ur short, slightly curved, with rounded or only slightly subtylote, smooth bases, tapering tn shárp fusiform points, barely differentiated fram subectosomal auxiliary spicules, being only marginally thicker and more curved and with predominantly smooth bases. Subectosomal auxiliary suhtylostyles relauve- ly long, straight, thin, sharply pointed, with most- ly microspined subtylote bases, Ectosomal auxiliary subtylostyles short, rela- tively thick, slightly curved, usually with sub- tylote microspined bases, Acanthostyles relatively long, thin, subtylote with aspinose necks, spines slender, long, slightly recurved, Micrescleres (Table 35). Palmate isochelae very abundant, in two size classes, the smaller often contort; larger with lateral alae marginally smaller than front ala, with lateral alae not com- pletely fused to shaft, and front ala widely separated from lateral alae. Toxas very abundant, short. thin, mostly wing- shaped with slightly curved centre and slightly unreflexed points, sometimes u-shaped with nearly straight arms and slight central curvature, Larvae. Incubated larvae large, spherical parenchymella, ap to 42521 diameter, with light matrix and no larval spicules. REMARKS, The presence of two size classes of isochelae microscleres, some with geometric modifications, was not previously described for this species, but in other spicule measurements Thiele's (19033) and Burton's (19342) specimens agree closely. The two type specimens differ slightly in their gross morphology, and this provides some evidence to illustrate the effect of growth form directly determining skeletal development. Whereas the lectotype is thinly encrusting and hymedesmoid in architecture, the paralectotype is thick and has a very well developed microcionid choanosomal structure, with ascending fibre nodes closely resembling C, (Microciona) seriata (sensu Simpson, 1968a). REVISION OF MICROCIONIDAE 315 PA jl ! Wal NAIK i 500 um 100 um Ni } ‘ FIG. 155. Clathria (Thalysias) coralliophila (Thiele) (lectotype SMF1784). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Wing-shaped - U-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Burton (1934) BMNH1930.8.13.107. MEMOIRS OF THE QUEENSLAND MUSEUM Dye URS ii SI Mi d N as ENS FIG. 156. Clathria (Thalysias) coralliophila (Thiele) (BMNH1930.8. 13.107). A, Choanosomal skeleton. B, Fibre characteristics (x219). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of subectosomal and ectosomal auxiliary subtylostyles. G-H, Palmate isochelae. I, Wing-shaped and u-shaped toxas. REVISION DF MICROCIONIDAE Were it not for the presence of a specialised ectosomal skeleton this specimen , would be included in | the Micreciona group, | illustrating the dif- [principal sti 145-(252,8)-406 x | 164-(279.8)-349% | 285-(347.5)-408x | 224-(345.2)-452 n ficulty in recognising | principal styles 7-410.5)-13 10-(11.5)-14 12-(13.8)-15 10-(12.4)-15 generic concepts sole- | Subectosomal 321-(355.4)-390 x | 233-(302.7)-350x. | 312-(3904)-488 x | 223-(347.3)-430 lyonthebasisofskele- HA Earysles | 53k 601.2): 547.9)-10 54139 nl aTtizt bobsrE: ERAI 104-(150.1)-208 x. | 102-(115.8)-134x | 118-(1409)-174x | 109-(121.5)-138 \ lathriid, h '" | auxiliary styles 2-(3.1)-4 | 2-(3.2)-4 2-(3.1)4.5 2-(3.5)-5 dts es Tod Wu Echinatin: 48-(56.3)-71 x j 52-(66.8)-78 x 84-(95.0)-105.& 76-(85.8)-95 x edesmoid or micro- E | aie d ( "th acanthostyles 445.26 345.2)-8 4-(6.3)-8 5. BETON See a t Bo- Ehe [cse BETTTEE toclathtia, Microciona, (Chemen ae Tee WB eu Specimens from the ^ 2 | 0540710 | — 05414-2 | — 05412)2 Great Barrier Reef and Papua New Guinea also differ from the In- donesian population in having some larger spicules (notably toxas, principal styles and acan- thostyles; Table 35), but generally spicule size and spicule geometry are comparable between all specimens, and there is no doubt that these populations are conspecific. This species is related to the New Caledonian C. (T) araiosa, differing in the distribution of spines on acanthostyles and auxiliary spicules. and specific dimensions of megascleres (which are generally smaller in C. (T.) araiosa), and Hooper & Lévi (19933) suggested that these similanties in skeletal architecture and spicule geometry may be indicative of a sibling species relationship. Clathria (Thalysias) costifera Hallmann, 1912 (Figs 157-158) Clathria costifera Hallmann, 1912; 215-218, p.31. fig.2, text-fig. 44; Vosmaer, 1935a: 648; Guiler. 1950: 6; Hooper & Wiedenmayer, 1994; 270, Pseudanchinoe costifera; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: AMES650: E. coast of Flinders I.. Bass Strait, Tas, 40°01'S, 148°02'E, coll FIV ‘Endeavour’ (dredge), OTHER MATERIAL S, AUST - AME!03S (dry), VIC - OMG300666 (NCIQ66C-3633-P) (fragment NTMZ3798). HABITAT DISTRIBUTION. Rock reef: 15-60m depth; Bass Strait (Tas), Furncaux Is (Vic), Kingston (SA) (Fig. 157G). DESCRIPTION. Shape. Large, thinly flabeliate, plannar, 170-220mm high, 120-270mm wide, with small cylindrical basal stalk. 24-55mm long, 18-38mm diameter, corrugated apical margins. 317 TABLE 35. Comparison between present and published records of Clathria (Thalysias) coralliophila (Thiele). All measurements are given in pm, denoted as range (and mean) of spicule length x spicule width (N25). Lectotype aralectotype |. GBR (BMNH 1 SRICULE SMF1784) 1930.8.]3.107) | (NCIQS6 suu) Colour Dark red alive (Munsell 2. brown in ethanol. 5R4/10); grey- Oscules. Small, up to 2.5mm diameter, scattered over entire surface, not apparently confined to any particular region. Texture and surface characteristics. Harsh, firm, compressible, flexible; surface rugose with prominent striations, raised ridges and subec- tosomal grooves running longitudinally and radially. Ectosome and subectosome. Dense ectosomal skeleton consisting of erect plumose brushes of smaller ectosomal auxiliary subtylostyles form- ing a continuous palisade on surface; subec- tosomal skeleton plumose with erect brushes of larger auxiliary subtylostylex ansing and diverg- ing from ends of choanosomal spicules in peripheral fibre skeleton; subectosomal megascleres prorude into and partially inter- mixed with smaller ectosomal spicules although both spicule categories with distinctly localised distribunum. Choanoseme. Choanosomal skeletal architecture irregularly reticulate. with heavy fibres and as- cending primary and transverse secondary fibre components; primary fibres have paucispiculur core of both choanosomal principal and subec- tosamal auxiliary megascleres. and choanosomal styles also protrude through fibres to [orm as- cending, plumose brushes near periphery; secon- dary fibres entirely aspicular; echinuting acanthostyles relalively sparse at core, slightly more numerous towards peripheral skeleton; mesohyl matrix moderately light, with few megascleres scattered between fibres. 318 Megascleres. Choanosamal principal styles long or short, moderately slender, straight or only slightly curved at centre, with rounded or slightly subtylote smooth bases, fusiform points, Length 110-(189.6)-305pm, 4-(5.8)-7um width (holotype 129-315 x 6-91.m]. Subectosomal auxiliary subtylostyles slender, straight ar very slightly curved near basal end, with smooth well developed subtylote bases, fusiform points. Length 296-(321,3)-3424um, width 3-(3.8)-5j1.m (holotype 198-336 x 3-7um). Ectosomal auxiliary subtylostyles slender, straight or slightly curved at centre, wilh prominently subtylote smooth bases, fusiform points. Length 117-(152.2)-175pm, width 2- (2.6)4,0m (holotype 112-158 x 2-3.5j.m). Echinating acanthostyles slender, usually slightly curved at centre, with subtylole bases, fusiform points, rudimentary spination, small spincs, aspinose *neck' region proximal 1o base, Length 76-(£5.4)-95pm, width 4-(4.3)-5um (holotype 56-92 x 4-7 um). Microscleres. Isochelae absent. Toxas accolada, abundant, long, thin or rhaphidiform, with slightly angular central curva- ture, straight arms, unreflexed points, sometimes completely straight. Length 176-4(215,2)-264 um, width 0.5-(0.8)- 1.Ujum (holotype 110-315 x 0.5- 1.20 m). REMARKS. Vosmaer's (19352) remark that C. (T) costifera was a synonym of C. (C.) caelata is not supported here. although both species do belong to Hallmann's (1912) ‘spicata’ group (with spicules protruding from fibres in plumose brushes). They differ in toxa geometry (rhaphidiform accolada versus thicker accolada plus oxhom, respectively), and C. (C.) caelata has only asingle size categary of auxiliary spicule (whereas C, (T.) costifera obviously belongs to C. (Thalysias) in having two categories). Although apparently initially identified only by superficial comparison with the holotype (ac- cording to its specimen label], another specimen found in general collections of the AM(E1035) also belongs to this species. The more recent material described above from the Furneaux Is- lands is surprisingly only the third known record for this large, conspicuous, brightly coloured species. It differs only slightly from the holotype in having abundant, very small sand grains scat- tered throughout the mesohyl. and accolada toxas that are nearly straight (oxeote) or have only very slight, angular, central curvature (whereas in the holotype they are more generously curved). MEMOIRS OF THE QUEENSLAND MUSEUM Clathria ias) craspedia sp. nov. (Figs 159-160, Plate 3E-F) MATERIAL. HOLOTYPE: QMG301436: Wommin Reef. S. of Cook Island, Tweed Heads, NSW, 28°12.1°S, 153934,8'E, 22m depth, 04.ii.1993, coll. J.N.A. & S.D. Cook (SCUBA). PARATYPE: QMG301452: Guy Rock, NW. side of Cook I.. Tweed Heads, NSW, 28°11,7°S, 153°34.6'E, 15m depth, 04.11.1993, coll. J.N.A. Hooper & S.D. Cook (SCUBA). HABITAT DISTRIBUTION. In sand, coral rabble coral substrata at base of granite boulders; 15-22m depth: Tweed River region (NSW) (Fig. 1596). DESCRIPTION. Shape. Growth form erect, lamellate, digitate or bulbous-lobate, 120- 230mm long, 8-50mm diameter, partially bur- rowed into soft sediments with rhizomous root-like attachments found below the surface; digits slightly flattened, irregularly shaped, usually branching, typically anastomosing with or entirely fused to adjacent digits forming con- tiguous lamellae, occasionally tsolated, single, completely attached or only partially attached to substrate, with tapering and frequently bifurcate apex. Colour. Yellow-orange (Munsell 7.5YR 7/10) to red-brown alive (1OR 6/10), grey-brown in ethanol. Oscules. Large, 2-5mm diameter, shghtly raised above surface, with membrancous lip, scattered over lateral margins of digits or on apex of digits, Texture and surface characteristics. Firm, com- pressible, flexible; surface fleshy, mostly smooth, relatively even in cylindrical specimens, or with crenellated margins in erect bulbous specimens. Eciosome and subectosome. Ectosomal skeleton composed of relatively dense but discrete bundles of smaller ectosomal auxiliary styles, supported beneath by paratangential, occasionally plumose brushes of larger subectosomal auxiliary suh- tylostyles arising from terminal choanosomal spongin fibres: mesohyl matrix light in choanosome but more darkly pigmented in peripheral skeleton. Choanosome. Regularly reticulate, wide- meshed, with heavy spongin ftbres differentiated into primary and secondary elements, but no axial compression or differentiation berween axial and extra-axial regions of skeleton; fibre diameter relatively homogeneous throughout skeleton. with fibres distinguished mainly by numbers of coring spicules, whereas fibre nodes prominently bulbous, up to 160j.m diameter: primary ascend- ing fibres, 45-90, diameter, cored by 4-8 REVISION OF MICROCIONIDAE 319 100 um FIG. 157. Clathria (Thalysias) costifera Hallmann (holotype AME650). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Accolada toxa. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype. I, Live NTMZ3798, 320 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 158. Clathria (Thalysias) costifera Hallmann (holotype AME650). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinated fibres. D, Echinating acanthostyle. E, Acanthostyle spines. F, Oxeote accolada toxas. REVISION OF MICROCIONIDAE 321 FIG. 159. Clathria (Thalysias) craspedia sp.nov. (paratype QMG301452). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle/ style. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Paratype. spicules abreast; secondary mainly transverse, ous, triangular or oval meshes, 180-360pm connecting fibres, 40-65j.m diameter, with 1-3 diameter; echinating acanthostyles absent; spicules abreast; fibre reticulation forms cavern- mesohyl matrix heavy but only lightly pigmented 322 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 160. Clathria (Thalysias) craspedia sp.nov. (holotype QMG301436). A, Choanosomal skeleton. B, Fibre characteristics. C, Palmate isochelae. D, Wing-shaped toxas. REVISION OF MICROCIONIDAE thoughout choanosome, with numerous auxiliary spicules scattered between fibres; choanocyte chambers oval, 30-45jum diameter. Megascleres. Choanosomal principal styles long or short, slender, straight, with evenly rounded smooth bases, telescoped points. Length 103- (221.4)-305 jum, width 3-(4.1)-6m., Subectosomal auxiliary subtylostyles long, slender, straight, with smooth subtylote bases, fusiform points. Length 255-(316.6)-361 um, width 2.5-(3.8)-4.5u m. Ectosomal auxiliary styles short, slender, straight, with smooth subi ylote or evenly rounded bases, fusiform points. Length 107-(135,4)- 174m, width 1.542.3)-3 um. Echinating megascleres absent. Microscleres. Palmate isochelae in single size class but variable length, with front ala slightly longer than lateral alae, lateral alae fused com- pletely to shaft, front ala entire, and apex of spicule characteristically constricted, pointed. Length 5-(10.7)-144um. Toxas moderately short, thick, wing-shaped, with rounded or slightly angular central curva- ture, non-reflexed arms. Length 15-(42.7)-764.m, width 0.8-(1.9)-2.5].m. ETYMOLOGY. Greek kraspedon, edge or border; occunng in the transition zone between the Salanderian and Peroman biogeographic provinces. REMARKS. This is a sibling species of C. (7) cervicornis, initially referred to that species based on skeletal structure and spicule types; it is separate by its different erowth forms, surfacc features, live colouration and several subtle but important skeletal characters (i.e., C (T) craspedia lacks echinating megascleres, the bases of all megascleres have different termina- tions such as telescoped points, smooth subtylote swellings, principal and auxiliary megascleres are differentiated within fibres, and isochelae have a terminal tooth-like canstriction). These differences in skeletal characters might be inira- specific variability, and the 4 populations (i.e., including C. (T.) fusterna sp. nov. and C. (T.) corneolia) may represent à single, widely dis- persed species. I consider major differences in growth forms, surface features and live coloura- tion are consistently correlated to skeletal dif- ferences, supporting distinct taxa for the Tweed Riverand Gulf of Carpentaria populations (C. (7:) fusterna below and Hooper & Lévi, 19933). Whereas C. (T.) cervicornis is habitually long, thin, cylindrical, digitate, attached to the substrate 323 at one or few points, and characteristically forms extensive tangles or thickets. Clathria (Thalysias) darwinensis sp, nov. (Figs 161-162, Plate 6B) MATERIAL. HOLOTYPE - QMG303375: Stephen's Rock, West Arm, Darwin Harbour, NT, 12°29,2'°5, 130°47.0'E, 19m depth. 24.ix.1993, coll. J.N.A. Hooper, L.J. Hobbs & B. Alvarez (SCUBA), HABITAT DISTRIBUTION, Coral pinnacle pear mouth of estuary, high sediment, turbid water; [$m depth; Darwin Harbour (NT) (Fig. 161H). DESCRIPTION. Shape. Arborescent, very thinly branching, reminiscent of an Axinella ( Axincl- lidae), 220mm high, 340mm maximum breadth of branches, with short basal stalk and point of attachment, 85mm long, 35mm diameter, main branches long, subcylindrical, up to 22mm diameter, slightly flattened, producing numerous smaller branches, up to 14mm diameter, con- voluted, bulbous branch nodes, and branches repeatedly bifurcate, decreasing in size, towards tapering, pointed branch tips. Colour. Pale cream alive (Munsell 5YR 8/2), darker yellow-brown in air, pale brown in ethanol. Oscules, Small, on lateral sides of branches, up to 3mm diameter, surrounded by sliphtly raised membrancous lip. Texture and surface characteristics. Saft, com- pressible, flexible branches, more harsh in ethanol; surface optically hispid, fleshy alive, even, bulbous, turgid, non-porous, but contract- ing greatly in ethanol producing porous, microconulose, uneven, irregular surface with scattered sharp conules. Ectosome and subectosome. Ectosome dominated by long, single. erect principal styles at regular intervals on surface, 400-500).m apart, extending 300-450p.m from surface, surrounded at base by paratangential tracts of both larger and smaller auxiliary subtylostyles, sometimes in plumose brushes surrounding base of principal spicele, more often in tangential or paratangential tracts: echinating acanthostyles also erect peripheral fibres, protruding through surface; subectosomal skeleton usually reduced with penpheral choanosomal fibres immediately below ectosome, whereas on surface conules auxiliary spicules produce more-or-less erect bundles associated with protruding principal spicules; no obvious localisation of smaller (ec- tosomal) or larger (subectosomal) auxiliary spicules, both appearing to be intermingled in 324 surface brushes; mesohyl matrix moderately heavy in ectosomal skeleton. Choanosome. Choanosomal skeleton irregularly reticulate, more regular (subreneiroid) in peripheral region. slightly compressed at axis; spongin fibres heavier in axis (110-160j1.m diameter) than at periphery (60-90p.m diameter), producing wide-meshed reticulation and slight axial compression: fibres imperfectly divided into primary and secondary elements; primary fibres ascend to surface with little or no bifurca- tion and relatively few transverse connecüng fibres, producing a nearly subreneiroid peripheral skeleton; primary fibres cored by 2-5 principal spicules, confined entirely to centre of each fibre, not protruding through fibres except at surface; secondary fibres short, more-or-less transverse, coned by 1-3 principal spicules abreast, intercon- necting primary fibres mainly in axial region of skeleton, producing oval or elongate fibre meshes, generally smaller at core (120-190um diameter) than periphery (170-240;.m diameter); fibres moderately heavily echinated by acanthos- tyles, evenly distributed over fibres although pas- sihly more abundant on exterior surface of fibres, especially in peripheral skeleton; mesahyl matrix moderately light, including some auxiliary spicules scattered between fibres; choanocyle chambers small, oval, 12-24jum diameter. Megascleres. Choanosomal principal styles long, robust, slightly curved near base, entirely smooth, evenly rounded bases without any tvlote swell- ing, long, tapering, fusiform points. Length 188- (M)1.8}-492 um, width 4-(12.8)-21 um. Subectosomal auxiliary subtylostyles long. slender, straight, slightly subtylote microspinecd bases, fusiform points, Length 210-(282.2)- 365um, width 3-{4.3)5um. Ectosomal auxiliary subtylostyles short, slender, straight, slightly subtylote microspined bases, fusiform points, Length 115-(135.3)- 153m, width 1.542. 1)-2,5 pum. Echinating acanthostyles long, slender, straight or slightly curved at centre, slightly sub- tvlote bases, evenly spined except for aspinose ‘neck’ proximal to base; spines large, recurved. sharp; points sharp or slightly rounded, spinose. Length 96-(104.8)-1160m, width 3-(5.6)-1] um. Microscleres, Palmate isochelae abundant, single size class. unmodified, with Lateral and front alae approximately same length. long, lateral alae en- tirely fused to shaft, front ala detached along lateral margin. Length 15-(16,.8)-1 8pm. Toxas abundant, wing-shaped, thick, variable in length, with wide central curvature, slightly MEMOIRS OF THE QUEENSLAND MUSEUM reflexed arms. Length I: 73-(111.2)-124j.m, width 2-(3.8)-8p.m; length II: 17-(26.2)-361. m. width 0.5-(1.7)-2.04um. ETYMOLOGY. For the type locality. REMARKS. Clathria (T) darwirensis is similar to C. (T) lendenfeldi, C. (C.) inanchorata and C. (I) coppingeri of the spicata group having choanosomal principal spicules protruding through peripheral spongin fibres forming a hispid surface. It differs from these species, and lo some extent the concept of the spicata group, having smooth principal styles enclosed within spongin fibres, only protruding through fibres ar the surface, and with all fibres more-or-less fully cored by principal spicules. This species is also similar to C (C) transiens in ectosomal structure (with prominent, individually protruding, smooth principal spicules), and also in having a vaguely sub-remeroid skeletal architecture, and toxa mor- phology, but the two differ in their acanthostyle geometry (in C. (T) darwinensis these are long, slender, unevenly spined, with large, recurved spines, whereas in C. (C. ) transiens they are short, unspined, or evenly lightly spined with vesugal spines}, possession of 2 size classes of auxiliary styles (versus one size class), thinly branching gross morphology (versus bulbous branches), and spicule dimensioas. Clathria (Thalysias) dubia (Kirkpatrick. 1900) (Figs 163-164) Mierociona dubia Kirkpatrick, 19003; 128, 136, 141, pl. 12, fig.3,3a, pl.13, fig.2a-f. Cionanchora dubia; de Laubenfels, 1936a; 108. Clathria dubia; Hooper & Wiedenmayer, 1994: 270. cf. Microciona prolifera, Vosmaer, 1935a: 608, 643. MATERIAL. HOLOTYPE: BMNHI898.12. 20.37: Flying Fish Cove, Christmas I.. Indian Ocean, 10°25.5°S, 105*4Q' E, coll. Mr Andrews (dredge). HABITAT DISTRIBUTION. Coral mbble; probably intertidal; Chrisimas I. (Indian Ocean) (Fig. 163H). DESCRIPTION. Shape. Thickly encrusung lamella, 12mm diameter, on eroded bi valve shell. Colour. Yellow preserved. Osciles. Not seen. Texture and surface characteristics. Compres- sible: optically smooth surface. Ectoseme and subectosome. Ectosome micros- copically hispid, with bundles of ectosomal auxiliary megascleres protruding through sur- face, forming a relatively thick dermal palisade, arising from subdermal brushes of subectosomal REVISION OF MICROCIONIDAE 325 FIG. 161. Clathria (Thalysias) darwinensis sp.nov. (holotype QMG303375). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyle/ styles. D, Echinating acanthos- tyle. E, Wing-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. J, Ectosoml specialization. spicules; subectosomal skeletal architecture plumose. with subectosomal auxiliary subtylos- tyles arising from ends of choanosomal megascleres. Choanosome. Choanosomal skeletal hymedes- moid, with a thin layer of spongin lying on sub- strate, in which bases of erect choanosomal principal subtylostyles and acanthostyles are em- bedded; small amounts of detritus scattered within skeleton; mesohyl matrix relatively heavy. Megascleres. Choanosomal principal subtylos- tyles long or short, slightly curved, usually with prominently microspined bases, occasionally smooth, bases subtylote, points fusiform. Length 132-(195.6)-2924um, width 7-(10.6)-16j1.m. Subectosomal auxiliary subtylostyles long, straight, with prominent subtylote, microspined 326 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 162. Clathria (Thalysias) darwinensis sp.nov. (holotype QMG303375). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E, Base of ectosomal auxiliary subtylos- tyle. F, Palmate isochela. G, Wing-shaped toxas. REVISION OF MICROCIONIDAE bases, fusiform points. Length 218-(280.8)- 314m, width 4.5-(5.9)-7 jum. Ectosomal auxiliary subtylostyles straight, with well formed tylote, microspined bases, fusiform points. Length 86-(110.2)-153pm, width 3-(3.9)-5j.m. Acanthostyles short, thick, tapering club- shaped, with large recurved spines on basal por- tion of shaft, aspinose point; spines on basal swelling often bifurcate, greatly recurved. Length 35-(46.4)-54jum, width 5-(8.2)-11 jum. Microscleres. Isochelae modified palmate, very small, often anisochelate, with lateral alae com- pletely fused to shaft, front ala often bifurcate or trifurcate, producing multiple, partially fused teeth. Length 2-(4.8)-8jum. Toxas divided into two morphs - I: very short, ? oxhorn, relatively thick, slightly curved at centre, with slightly reflexed points. Length 4- (6.6)-91.m, width 0.5-(1.1)-1.5j.m. IT: Accolada, long, thin, with gently rounded or angular curva- ture, with straight points. Length 112-(195.2)- 295m, width 0.5-(1.4)-2j.m. REMARKS. De Laubenfels (1936a) assigned this species to Cionanchora because it supposed- ly had anchorate (rather than palmate) isochelae, although differing in no other respect from typical species of Clathria (Thalysias). Scanning electron micrographs show that these chelae have modified lateral alae completely fused to the shaft and front alae often split into several 'teeth', producing an anchorate-like appearance, but they are obviously palmate in origin. The species is well characterised by it megasclere and microsclere geometry, although the species is so far known only from a single specimen from Christmas Island. Clathria (Thalysias) erecta (Thiele, 1899) (Figs 165-166, Table 36) Rhaphidophlus erectus Thiele, 1899: 14-15, pl.2, fig2; Thiele, 1903a: 957; Hallmann, 1912: 177; Lévi, 1961a: 136-137, text-fig.10, pl.1. cf. Microciona prolifera; Vosmaer, 1935a: 611. MATERIAL. LECTOTYPE: NMB19 (fragment BMNH1908.9.24.163): Kema, Minahassa, Celebes (Sulawesi), Indonesia, 2°S, 120°30’E, coll. P. & F. Sarasin (dredge). PARALECTOTYPE: NMB18 (frag- ment BMNH1930.7.1.7): same data. OTHER MATERIAL: NT - NTMZ3113 (fragment QMG- 300579), NTMZ3146 (fragment QMG300219). INDONESIA - SMF1788. HABITAT DISTRIBUTION. Coral reef and coral rub- ble; 16-20m depth; known Australian distribution: 327 Parry Shoals, Timor Sea (Fig. 165H); also Moluccas and Sulawesi, Indonesia (Thiele, 1899, 19032), Viet- nam (Lévi, 19612). DESCRIPTION. Shape. Elongate, arborescent, 90-240mm high, with a short cylindrical stem, 25-75mm long, 15-25mm diameter, bifurcate and relatively thick cylindrical branches, up to 35mm diameter, or lamellate, fused, erect digitate branching pattern. Colour. Orange to dull brown alive (Munsell 5YR 7/10 - 7.5YR 7/6), beige in ethanol. Oscules. Numerous, small, up to 2mm diameter, scattered over all sides of branches, below surface conules. Texture and surface characteristics. Firm, flexible, compressible; highly conulose, rugose surface, pocked with holes and drainage canals. Ectosome and subectosome. Thin but prominent discrete brushes of small auxiliary subtylostyles standing more-or-less perpendicular to surface; subectosomal region cavernous, with plumose tracts of larger subectosomal auxiliary, and choanosomal principal megascleres supporting ectosomal skeleton and protruding through sur- face. Choanosome. Very irregularly reticulate, cavern- ous, with very large primary fibres running lon- gitudinally through branches, up to 140m diameter, interconnected by smaller tangled secondary fibres, up to 70jum diameter, produc- ing vaguely triangular skeletal meshes, up to 450p.m diameter; both primary and secondary fibres heavy, fully cored by multispicular tracts of choanosomal principal styles, and lightly echinated by acanthostyles, the latter slightly more abundant at fibre nodes; mesohyl matrix moderately heavy, with auxiliary megascleres dispersed between fibres. Megascleres (Table 36). Choanosomal principal styles characteristically curved near basal end, hastate pointed, with rounded or faintly subtylote, smooth bases. Subectosomal auxiliary subtylostyles long, thick, straight, fusiform pointed, faintly subtylote smooth bases, or minutely microspined bases. Ectosomal auxiliary subtylostyles small, slender, prominently subtylote, with microspined bases. Acanthostyles long, thick, with subtylote bases, fusiform points, heavily spined on base and point but unspined neck; spines characteristically large, recurved, heavily concentrated at point of spicule. Microscleres (Table 36). Palmate isochelae in two size classes, the smaller sometimes contort. TABLE 36. Comparison between present and publish- ed records of Clathria (Thalysias) erecta (Thicle). Measurements in pum, denoted as range (and mean) of spicule length x spicule width (N=25). A i Holotype | Specimens (2) | SOME ‘Timor Sea 142-4197,7)-259 x 6-(10,9)-14 (Lévi, 1961 y Vietnam} 190-300 110-22 165-(226.9)-262 x 4-(6.6)-9 255-310 x 6-7 924 138.5)-198 V 3-(4.2)5 | 85-(102,7)-134 100-200 x 3-5 RHAG | Ekhinatin 6472.678 61-468.5)-75 x 6-168) | 10955910 | 5567-8 | Chele L 446.309 | 12-(134)-15 10-17 10-12.:)-14 12-(197.0265 > |25-13&3)-304 Toras xoxd.pi15s| 999? [osi s] Toxas accolada, relatively long, thin, with small. angular central curvature, or rounded centrally, straight arms and reflexed points; juvenile forms resemble oxhorn toxas. Larvae. Incubated parenchymella larvae small. spherical, 140-190,m diameter, with light miesohyl matrix and larval toxas dispersed within axis. REMARKS. This species is a sibling of C. (T) reinwardtii based on similarities in skeletal struc- ture (even-meshed, cavernous primary and secondary tracts), geometry of some spicules (smooth, curved principal styles; robust subec- tosomal auxiliary subtylostyles; slender accolada toxas with slightly angular central curvature; 2 sizes of jsochelae), and live colouration (orange- brown). They differ significantly in growth form (C. erecta is arborescent, thickly branching; C, reinwardti has only thin or slightly thick cylindn- cal branches), surface features (prominent sur- face conules versus completely smooth or irregularly rugose surface), and acanthostyle geometry (tapering, sharply pointed and sub- tylote bases versus rounded ‘points’ and only slightly subtylote or rounded bases). Spicule dimensions are also comparable (Tables 36, 39). These differences are consistent for the six known specimens of C. (T.] erecta and for the present these species are maintained as distinct. Another species, C. (T) fasciculata Wilson. from Indonesia and the central west Pacific ( Wil- son, 1925; de Laubenfels, 1954), is alse very simular to both C. (T. ) erecta and C. (T) reinwardti MEMOIRS OF THE QUEENSLAND MUSEUM in the diversity and geometry of its spicules, hut it differs again in growth form (being bushy, flattened branching), skeletal structure (more ir- regularly reticulate) and toxa morphology (in- cludes asymmetrical sinuous forms). lt is possible that all three species are extreme morphological variants of a single species, in which case rein- wardti would have priority, but all these mar- phological differences are consistent within each growth form type (morphospecies) and probably represent fixed genetic differences. Clathria (T.) erecta is also vaguely similar to C. (T) vulpine in ihe overall structure of spongin fibre skeleton and spicule skeleton, both species having a characteristic, more-or-less triangular skeletal network of fibre meshes fully cored by principal styles, although this fibre reticulation 1s much more regular in the latter species. This structural feature is prominent and their inferred similarities are immediately obvious upon casual observation, but the two species differ from each other in most other respects. The presence of two size classes of isochelae, including contort forms, has not been recorded previously for C. (T.) erecta but are consistently present in all specimens including the type material, Thiele's (1899, 1903a) Indonesian specimens are identical to the Timor Sea specimens in most respects (see Table 36), whereas Lévi's (19612) material from Viemam differs slightly in growth form (compare Lévi's (1961a) Plate 1 with Fig. 165I-I of the present study), and spicule dimensions are relatively Jarger. Clathria (Thalysias) fusterna sp. nov. (Figs 167-168) MATERIAL. HOLOTYPE: QMG303240: NE. Cape Grenville, Shelburne Bay, Qld., 11^03'S, 143*14* E, 27m depth, 04.iv.1993 (dredge). PARATYPES: QMG300862; NW. of Vrilya Point, Gulf of Carpen- taria, Qld, 11°12.7°S, 142?05.9'E, 21m depth, 30.xi.199] (dredge). QMG301L008: NW. of Pon Musgrave. Gulf of Carpentaria. Qld, 11^18.9'S. 140?55.8 E, 41m depth, 27.xi.199| (dredge). OTHER MATERIAL: GULF OF CARPENTARIA, - QMG- 301013, 0MG303462. RED SEA - PIBOCO4-17 (frag- ment OMG300064). HABITAT DISTRIBUTION Soft sediments, mud, shell gril; 21-58m depth; Shelburne Bay, Torres Strail and Gulf of Carpentaria (FNQ), Gove Peninsula (NT) (Fig. 167H); also Eritrea, Red Sea (present study). DESCRIPTION. Shape. Erect, club-shaped growth form 190-280mm long, with long thin, cylindrical stalk, 80-150mm long, up to 12mm REVISION OF MICROCIONIDAE 329 FIG. 163. Clathria (Thalysias) dubia (Kirkpatrick) (holotype BMNH1898.12.20.37). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthos- tyles. E, Accolada and oxhorn toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. diameter, bifurcating several times towards base becoming filamentous, rhizomous (for embed- ding in soft sediments); rhizomous roots encrusted with sand and shell fragments; apex of club usually large, 50-100mm diameter, 60- 110mm long, composed of fused, tightly anas- tomosing digits; adjacent digits fused with dense fleshy surface; apex of digits tapering slightly pointed. Colour. Grey-brown on-deck (Munsell 2.5 Y 8/2), grey in ethanol. Oscules. Few small pores, 0.5mm diameter, pos- sibly oscules, scattered near apex of digits (seen in preserved material only). Texture and surface characteristics. Stalk tough, wiry, flexible, apex of club softer, more compres- sible but with firm axis; slightly convoluted sur- face with sparse conules, up to 3mm high, low ridges or occasional folds on anastomosing digits. Ectosome and subectosome. Dense ectosomal skeleton composed of discrete bundles of smaller ectosomal auxiliary subtylostyles supported below by paratangential or plumose brushes of subectosomal auxiliary subtylostyles; mesohyl matrix moderately lightly pigmented in ec- tosomal skeleton; ectosomal and subectosomal spicule skeletons very dense but together com- prise only 10-2046 of branch diameter. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 164. Clathria (Thalysias) dubia (Kirkpatrick) (holotype BMNH1898.12.20.37). A, Choanosomal skeleton. B, Basal fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Bases of principal and auxiliary subtylostyles. F, Modified palmate isochelae. G, Accolada toxas. H, Juvenile oxhorn toxa. REVISION OF MICROCIONIDAE 100 um FIG. 165. Clathria (Thalysias) erecta (Thiele) (holotype NMB19). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary style. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Accolada toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. J, NTMZ3113. Choanosome. Heavily reticulate architecture; spongin fibres short, thick, 90-200j.m diameter, heavily collagenous, forming tight oval or rectan- gular meshes, 150-400j.m diameter; fibres not obviously differentiated into primary or secon- dary elements, but meshes slightly more cavern- ous in peripheral skeleton than in axis; fibres virtually fully cored by multispicular tracts of both subectosomal auxiliary subtylostyles and choanosomal principal styles, together occupy- ing 80-90% fibre diameter, interconnected by very large, bulbous fibre nodes, 160-400j.m diameter; fibre nodes contain larger bundles of spicules than in connecting fibres, indicating that fibres ascending through branches are heavier than fibres running from axis to peripheral skeleton; echinating acanthostyles abundant, concentrated mainly on fibre nodes; mesohyl 332 MEMOIRS OF THE QUEENSLAND MUSEUM na T MEM 4 A à st Wes (9 hy col i P. p- i "E y Pih A i.) t b T is à SM A if tn 1 ) $7 za ES A M i m 7 y F + L2 be a EY 1 D © L) © © FIG. 166. Clathria (Thalysias) erecta (Thiele) (QG300219). A, Choanosomal skeleton. B, Fibre characteristics (x389). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of subectosomal and ectosomal auxiliary subtylostyles. G, Accolada and juvenile toxas. H, Palmate isochelae. REVISION OF MICROCIONIDAE 333 FIG. 167. Clathria (Thalysias) fusterna sp.nov. (paratype QMG301008). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyles. E, Wing-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. 1, Holotype QMG303240. 334 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 168. Clathria (Thalysias) fusterna sp.nov. (paratype QMG300862). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E, Base of choanosomal principal sub- tylostyle. F, Wing-shaped toxas. G, Palmate isochelae. REVISION OF MICROCIONIDAE martrix heavy. with few auxiliary spicules scat- tered between fibres but abundant microscleres lining small oval choanocyte chambers, 20- 45pm diameter. Megascleres. Choanosomal principal styles robust short or long, thickest towards middle of spicule, straight or slightly curved towards pointed end, with slightly constricted bases, smooth or faintly microspined, fusiform points. Length 185-(264.3)-355u& m, width 5-(13.9)- 25j.m. Subectosomal auxiliary subtylostyles long, thick or thin, mostly straight, with subtylote microspined bases, onlv occasionally smooth bases, and fusiform points. Length 211-(369,2)- 385m, width 4-(9.5)-12].m. Ectosomal auxiliary subtylostyles short, straight or slightly curved near basal end, sub- tylote microspined bases, fusiform points. Length 99-[127.7)-163,um, width 2-(3.2)-41um. Echinating acanthostyles thick, robust, rela- tively short, heavily spined but with bare neck and point; spines moderately large, conical erect (not recurved), Length 73-(82,8)-96p. m, width 3-(7.4)-12u m. Microscleres. Palmate isochelae abundant, small, single size class, some contort, with lateral and front alae approximately equal length, lateral alae completely fused to shaft, front ala entire, slight constriction at apex of chela. Length 5-(11.8)- I4um. Toxas wing-shaped, relatively thick, with slightly angular central curve, arms at wide angles from centre, straight or slightly reflexed points. Length 18-44 1.4)-63ju.m, width 1.5-(1.9)- 25pm. ETYMOLOGY. Latin fusrerna , club or knotty part of a tree. REMARKS. This species is possibly a very atypi- cal, highly specialised population of C. (7°) cer- vicornis with a specialised. peculiar growth form adapted to living in soft sediments (long stalk, rhizomous roots, club-shaped apex). Its live colouration, gross skeletal structure and spicule diversity are closely comparable with typical populations of C. (T.) cervicornis. However, there are subtle differences in skeletal characteristics that consistently differentiate the two popula- tions: possession of differentiated principal and auxiliary spicules (whereas cervicornis has un- differentiated structural megascleres), acanthos- tyle spines are erect, conical (not recurved), a single size class of palmate isochelae (not two), and slightly subtylote bases on principal and 335 auxiliary sprcules (not prominently subtylote as in most C. cervicornis). These subtle differences correlate with the major differences in growth forms and are consequently considered here to justify the recognition of the Gulf of Carpentaria population as a distinct species in a species com- plex of four: the cylindrical C. (T.) cervicornis from the Indo-Malay - western Pacific region; the lamellate C. (T.) craspedia sp. nov. from the southern Solanderian province of Australia, and the New Caledonian species C. (T. ) comeolia (see Hooper & Lévi, 19932), This species is dis- cussed further in the remarks under C. (T.) craspedia. Clathria (Thalysias) hallmanni sp. nov. (Figs 169-170, Plate 6C) MATERIAL. HOLOTYPE: NTMZ2218: Vesteys Beach, Fannie Bay. Darwin, NT, 12°26.2'S, 130°49.9 E, intertidal, 21.1.1985, coll. J.N.A. Hooper, HABITAT DISTRIBUTION. Encrusting under beach rock and coral rubble; intertidal pools; NT (Fig. 169H). DESCRIPTION. Shape, Thinly encrusting, upto I.5mm thick, extending approximately 7cm across rock and dead coral substrata. Colour. Dark grey-brown orange-brown in life (Munsell 2.5R 5/4); pale grey in ethanol. Oscules. Minute, less than 1mm diameter, scat- tered evenly over surface. Texture and surface characteristics. Spongy, easily torn from substrate; surface has a dull slimy appearance due to production of small amounts of clear mucous upon exposure to air; surface optically smocth, even, without conules, ridges or canals, and encrusiation conforms exactly with contours of substrate. Eciosome and subectosome. Opaque in hie, slightly pellucid, subdermal canals or cavities not visible; ectosomal skeleton with extensive plumose brushes of small auxiliary subtylostyles. through which protrude ascending, plumose tracts of larger subectosomal auxiliary subtylos- les; moderate quantities of detritus in ectosomal skeleton; subectosomal region extensive, oc- cupying 70% of sponge thickness. composed of mostly paratangential tracts of larger auxiliary subtylostyles gradually ascending and diverge al surface, Choanosome. Skeletal architecture hymedes- moid in choanosomal (basal) region, but distinct- ly plumose towards peripheral skeleton; spongin fibres consist of a basal layer of spongin lying ugainst substrate, 18-35p.m thick, with bases of thounosomal principal subtylostyles and acan- 336 thostyles embedded in spongin and standing per- pendicular to substrate; choanosomal principal subtylostyles morphologically close to subec- tosomal auxiliary subtylostyles, and so difficult to determine exactly where basal mineral skeleton ends and where subectosomal skeleton begins, but extra-fibre multispicular tracts appear to begin close to basal layer; moderately common acanthostyles echinate basal spongin, whereas principal megascleres less common; choanocyte chambers 35-48jum diameter; mesohyl matrix heavy, granular, with small amounts of detritus Megascleres. Choanosomal principal subtylos- tyles long, fusiform, with slightly constricted bases or subterminal bases, entirely smooth or with low apical conules (? vestigial spines), and typically slightly curved towards basal end. Length 312-(385.5)-419.5um, width 8-(9.3)- llum. Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, almost indistinguishable from choanosomal megascleres but with prominent spined subtylote bases. Length 284.5- (362.2)-450um, width 2-(3.2)-4.5jum. Ectosomal auxiliary subtylostyles short, thin, fusiform, with microspined subtylote bases. Length 94-(121.2)-151j.m, width 0.8-(1.4)- 2.5m. Acanthostyles subtylote, fusiform, relatively evenly spined although spines less heavily con- centrated in ‘neck’ region, proximal to base, heavier on apical and distal extremities; spines relatively small, weakly formed. Length 52- (59.5)-72yum, width 3-(4.5)-6.5 jum. Microscleres. Palmate isochelae relatively com- mon, variable in size but not easily differentiated into two size classes, unmodified, with lateral alae entirely fused to shaft, approximately equal in length to front ala, and entirely free from front ala except in juvenile forms. Length 5-(10.8)- 17m. Toxas accolada, moderately common, long, thin, almost straight, with only slight angular central curvature, straight arms, straight (un- reflexed) points. Chord length 174-(208.0)- 481m, width 0.4-(0.8)-2.0.m. Associations. Single known specimen growing next to encrusting sponges (Renieria, Haliclona, Mycale), polychaete worm tubes (Pomatoleios kraussii) and simple ascidians. ETYMOLOGY. For E.R. Hallmann in recognition of his contributions to Australasian microcionids. REMARKS, It is difficult to define C. (T) hallmanni in any single unique character apart MEMOIRS OF THE QUEENSLAND MUSEUM from the close resemblance between choanosomal principal and subectosomal auxiliary subtylostyles. As far as can be ascer- tained from personal knowledge of the Australasian sponge fauna and Indo-west Pacific literature its field characteristics are unique. It is acknowledged that many older published descriptions of encrusting microcionids, espe- cially those from the Indo-Malay archipelago, rarely include details on live colouration or sur- face details. But none of these species match the present one in spicule geometry either. Conse- quently, C. (T.) hallmanni can be differentiated from other encrusting (hymedesmoid) Clathria (Thalysias) species in: grey-brown live colour; even (unornamented) surface, i.e., lacking subec- tosomal drainage canals commonly found in thin- ly encrusting species; plumose ectosomal and subectosomal skeletal structure as well as exten- sive paratangential tracts composed of both sorts of auxiliary spicules in the periphery; entirely smooth, relatively short and thin choanosomal subtylostyles, barely different from the subec- tosomal auxiliary subtylostyles except for pattern of spination; evenly spinous acanthostyles, un- modified palmate isochelae, and thin, nearly straight toxas with unreflexed arms. None of these features are unique or particularly distinc- tive by themselves but their combination is uni- que for this new taxon. Clathria (Thalysias) hesperia sp. nov. (Figs 171-172, Plate 6D-E) MATERIAL. HOLOTYPE: QMG300213 (fragment NTMZ3041): N. of Amphinome Shoals, Northwest Shelf, WA, 19?19,7-23.3'S, 119?08.8-12.2'E, 50m depth, 19.vii.1987, coll. J.N.A. Hooper (beam trawl), PARATYPE: NTMZ3327 (NCIQ66C-1407-U, frag- ment QMG30499]1): 1.8km N. of Bessieres I, Anchor Is, Exmouth Gulf, WA, 21?30.6' S, 114?45.4'E, 17m depth, 23.viii.1988, coll. D. Low Choy & NCI (SCUBA). HABITAT DISTRIBUTION. Coral rubble and deeper rock reefs exposed amongst gravel and shell grit sub- strates; 17-50m depth; NW. coast (WA) (Fig. 171G). DESCRIPTION. Shape. Thickly flabellate, simple planar fans resembling a Phakellia, or slightly cup-shaped with convoluted, concentric, smaller lamellae inside larger lamellae, resem- bling species of Cymbastela (Axinellidae); mar- gins pointed digitate (paratype) or convoluted folded (holotype); lamellae up to 380mm wide, 235mm high, 15mm thick; holotype probably lying on, or parallel to, substrate, with con- REVISION OF MICROCIONIDAE 337 100 um FIG. 169. Clarhria (Thalysias) hallmanni sp.nov. (holotype NTMZ2218). A, Choanosomal principal subtylos- tyle. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Palmate isochelae. F, Accolada toxa, G, Section through peripheral skeleton (hatched area coralline substrate), H, Australian distribution. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 170. Clathria (Thalysias) hallmanni sp.nov. (holotype NTMZ2218). A, Choanosomal skeleton through thick region. B, Hymedesmoid basal skeleton. C, Echinating acanthostyle. D, Acanthostyle spines. E-G, Bases of principal and auxiliary subtylostyles. H, Palmate isochelae. I, Accolada toxas. REVISION OF MICROCIONIDAE voluted, concentric ndges or small digitate processes arising from upper surface; paratype erect, perpendicular to substrate, with folded ridged running longitudinally; both specimens with differentiated osculiferous (upper) and porous surfaces, latter relatively even, smooth. Colour Pale orange-red alive (Munsell 5R 8/4), khaki-brown in ethanol Oscules. Small. up to 3mm diameter alive, smaller in preserved specimen, slightly raised above surface, with membraneous lip. only found on 1 surface of lamellae. Texture and surface characteristics. Firm, harsh, flexible, slightly compressible, difficult to tear; lower surface s: , even, upper surface more conulose (with terminal oscules), longitudinal folds, ridges ar convaluted folds and small digits. Ectosome and subectosame. Discrete surface brushes produce specialised ectosomal skeleton: brushes. composed of ectosomal auxiliary sub- tylostyles on outer surface forming thick, erect bundles bit not continuous palisade; with 1 or several choanosomal principal styles also protruding through surface associated with ec- tosomal brushes; subectosomal auxiliary sub- tylostyles intermingled with ectosomal spicules but originating slightly lower in peripheral skeleton; subectosomal region greatly reduced with peripheral choanosomal fibres lying imme- diately below ectosome: mesohyl matrix moderately heavy in peripheral region. Choanosome. Choanosomal skeleton almost regularly renieroid reticulate although renicroid pattern severely disrupted by heavy concentra- tions of echinating spicules (both principal styles and acanthostyles); spongin fibres very well developed, dark brown, imperfectly divided into primary (90-140pm diameter) and secondary (2545 diameter), and very large fibre nodes (up to 220m diameter); primary ascending fibres cored by multispicular tracts of principal styles. 2-5 spicules abreast, with spicules protruding slightly through fibres, particularly at fibre meshes. producing nearly plumose tracts; secon- dary more-or-less transverse fibres relatively short, interconnecting primary elements, cored by 1-3 spicules abreast; spicules occupy only 40-70% of fibre diameter for secondary and primary fibres, respectively; near peripheral skeleton principal styles distinctly plumose, with those on ultimate fibres contributing to ectosomal structure, whereas at core skeleton more renieroid reticulate; echinating acanthostyles very abun- dant, particularly at fibre nodes, also contributing to ectosomal spicule brushes, with only small 339 portion of base of acanthostyle embedded in spongin fibre and consequently protruding a long way into choanosomal mesohyl: fibre meshes oval or squarish, more cavernous in periphery (45-11 5pm diameter) than at core (170-250um diameter): choanocyte chambers oval, 35-55. m diameter, often lined by isochelae; mesohyl matrix moderately heavy but only lightly pigmented. Megascleres. Choanosomal principal styles suaight or slightly curved near centre, with rounded or very slightly subtylote bases, bases usually smooth, occasionally microspined, long tapering fusiform points. Length 162-(187.3)- 213jum, width 8-(11.1)-]4ym-_ Subectosomal auxiliary subtylostyles straight or very slightly curved near basal end, slightly subtylote bases lightly microspined, fusiform points. Length 121-(138.1)-168um, width 4- (4.6)-5.S5pum. Ectosomal auxiliary subtylostyles only slightly shorter than subectosomal spicules but consis- tently thinner. with smooth subtylote bases, fusiform points, Length 97-(121.1)-147jum. width 2-(2.9)-4 5pm. Echinating acanthostyles long, slender, mostly straight, sometimes slightly curved near point, with subtylote bases, fusiform points, heavily spined on bases, shaft and points, aspinose on ‘neck’ proximal to base; spines short, sharp, recurved. Length 97-(103.6)-J]2pm, width 4- (5.3)-6j. m. Microscleres, Palmate isochelae very abundant. poorly silicified, sigmoid, with short, sharp, ves- uigial unguiferous alae. Length 11413.7}1 Sum. Toxas absent. ETYMOLOGY, Latin hesperius, western; from WA, REMARKS. This species is borderline between Clarhria and Thalysias given that the ectosomal skeleton consists of spicule brushes composed of auxiliary spicules of relatively homogenous lengths (i.e. not clearly differentiated into smaller auxiliary spicules supported by larger auxiliary spicules, characteristic of other Thalysias). Nevertheless, ectosomal and subec- tesomal spicules can be consistently differen- halted by their thickness as well as the absence or presence of microspines on their base, respective- ly, even though there is no marked difference in length between the two categories. Clathria (T) hesperia has a distinctive lamel- late growth form with differentiated osculiferous and porous faces. [ts skeleton is a mixture of plumose tracts (reminiscent of C. (M) coccinea, particularly its plumose fibre nodes, or the MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 171. Clathria (Thalysias) hesperia sp.nov. (paratype NTMZ3327). A, Choanosomal principal style and base. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Ectosomal auxiliary subtylos- tyle. E, Modified palmate isochelae, F, Section through peripheral skeleton. G, Australian distribution. H, Holotype QMG300213. ‘scabida’ species group with principal spicules protruding through the peripheral skeleton), with an underlying renieroid reticulate skeleton. Pal- mate isochelae are vestigial, unguiferous, sig- moid reminiscent of C. (7:) michaelseni (which is an encrusting, hymedesmoid species, has toxas, and differs from this species in virtually every other respect). Clathria (Thalysias) hirsuta Hooper & Lévi, 1993 (Figs 173-175, Table 37, Plate 6F, 7A) Clathria (Thalysias) hirsuta Hooper & Lévi, 1993a: 1259-1264, figs 19-20, table 10; Hooper & Wieden- mayer, 1994: 270. MATERIAL. HOLOTYPE: QMGL2746 (fragment NTMZ1551): Cairns region, Qld, 16°56'S, 146°00'E, 1982, coll. A. Kay (trawl). PARATY PES: QMGL2750 REVISION OF MICROCIONIDAE 341 FIG. 172. Clathria (Thalysias) hesperia sp.nov. (holotype QMG300213). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of subectosomal auxiliary subtylos- tyle. F, Reduced sigmoid palmate isochelae. 342 (fragment NTMZIS555). QMGL2754 (Fragment NTMZ1560): Cairns region, Qld, 16°56°S, 146?DXY E, coll. A. Kay (trawl). OTHER MATERIAL: QLD - QMG300328 (NCIQ66C-1893-X; fragment NTMZ3513), NTMZ3494, QMG303040, QMG- 304767, QMG300081, QMG303971. NSW - QMG300771 (NCIQ66C-1185-F). NEW CALEDONIA - QMG301274, QMG301325, QMG301340. HABITAT DISTRIBUTION. Rock, dead coral and caral rabble substrates, usually on broken substrates, sides of bommies, or in gullies; 7-30m depth; Shel- burne Bay, Howick Is (FNQ), Whitsunday Is (NEQ), Noosa Heads, Stradbroke Í. (SEQ), Solitary Is (N. NSW) (Fig, 1730). Also New Caledonia lagoon (Hooper & Lévi, 19932). DESCRIPTION. (See Hooper & Lévi, 19933). DIAGNOSIS (Tahle 37). Tubular, lobo-digitate, reliculate-honeycombed, excavated growth forms superficially resembling Phakellia caver- nosa; bright red (or orange-red) conules, paler pink or white between conules, prominent sub- dermal drainage canals; large oscules scattered between surface projections; texture firm, com- pressible, slightly arenaceous; surface prominently conulose, conules pointed (or rounded, fleshy); ectosome with irregular. tan- gential or paratangential layer of intermixed ec- iosamal and subectosarnal subtylostyles (or with light palisade of smaller auxiliary styles focming erect brushes arising from ends of larger auxiliary spicules); thick choanosomal fibres immediately below ectosome (or subectosome cavernous); choanosomal skeleton irregularly reticulate (or regularly renieroid reticulate), with fibre skeleton dominant over spicule skeleton; primary fibres multispicular, meaning longitudinally through branches, ascending to surface, interconnected hy shorter uni- or paucispicular secondary fibres, cored by both shorter choanosomal principal styles and longer subectosomal auxiliary styles; acanthostyles dispersed evenly over fibres; choanosomal principal styles straight, with smooth, rounded or slightly subtylote bases und fusiform points: subectosomal auxiliary sub- tylostyles long, slender, straight, fusiform, with rounded or subtylote, smooth or microspined bases (or with simply rounded, smooth bases); ectosomal auxiliary subtylostyles short, straight, very slender, subtylote smooth or microspincd bases (or with simply rounded, smooth hases); acanthostyles small, subtylote, light or vestigial spines, aspinose ‘neck’ proximal to base; palmate isochelae small, unmodified. imperfectly divided into two size categories; lateral alae completely fused to shaft, completely detached from front ala MEMOIRS OF THE QUEENSLAND MUSEUM for whole of length. longer than front ala; ioxies accolada and wing-shaped morphs, very thin. sometimes slightly sinuous, rarely raphidiform, only slightly curved at centre, with straight non- Teflexed arms or only slightly reflexed points (or exclusively raphidiform with small angular central curve and straight arms). REMARKS. Clathria (T) hirsuta was originally described and illustrated from both New Caledonian and Queensland populations (Hooper & Lévi, 19932), in which it was reported that the New Caledonian population possessed ectosamal auxiliary subtylostyles (i.e., belonging to Thalysias) whereas Queensland specimens did not (i.e., belonged to Clathria). Since this publi- cation several more samples have been collected along the Queensland coast and Great Barrier Reef (Fig. 1730) in which specialised ectosomal spicules were discovered. Conversely, isochelae were originally reported only from the Queensland populations but absent in New Caledonian samples, but these have now also been observed in a recent sample collected from Noumea (albiet rare). Other dilferences between these two populations are discussed in Hooper & Lévi (19933. Two ‘atypical’ specimens from SE. Australia (QMG300328, G300771) (which are included in the diagnosis above, in brackets) fur- ther illustrate the considerable variability of this species (toxas are only raphidiform, auxiliary spicules lack tylote bases, and a structured cc- tosomal skeleton is present (Figs 174-175)). These specimens are reminiscent of the ‘reduced New Caledonian population. At first glance this species lacks any remark- able or unique feature that stands it apart frum other Clathria (Thalysias), but it possesses an unusual combination of characters not seen in any other species. It ts superficially similar to C. (7.) vulpina m growth form, but spiculation and fibre characteristics are quite different between the two specics. Its skeletal architecture and sptculation is also very similar to C. (T.) schoena Ge, USNM22404: which may be different again from Rhaphidephlus schoenus of authors; e.g., Simpson, [968a: Alcolado, 1980: Van Soest, 1984h), bur these species differ significantly im their growth form, spongin fibre architecture and cctosomal characteristics. Clathria (T.) hir- suta has very lightly spined acanthostyles, com- parable with those of C. (T.) rransiens, and it is also closely related to that species in its fibre characteristics and spiculation, although they REVISION OF MICROCIONIDAE 343 Dj Du VN Aa en VATI Y DY 100 um FIG. 173. Clathria (Thalysias) hirsuta Hooper & Levi comparison between typical and reduced populations (A-G, paratype QMGL2750; H-M, QMG300771). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyle. E, Accolada and wing-shaped toxas. F, Palmate isochelae. G, Section through typical skeleton. H, Choanosomal principal styles. I, Subec- tosomal auxiliary style. J, Ectosomal auxiliary style. K, Echinating acanthostyles. L, Raphidiform toxa. M, Palmate isochelae. N, Section through reduced specimen. O, Known Australian distribution. P, Holotype QMGL2746. Q, Atypical QMMG300328. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 174. Clathria (Thalysias) hirsuta Hooper & Levi typical population (A-C, F-G, QMG300081; D-E, H-J, paratype QMGL2750). A, Choanosomal skeleton. B, Fibre characteristics. C, Ectosomal skeleton. D-E, Echinating acanthostyle and spines (Cairns population). F, Echinating acanthostyle and spines (Moreton Bay population). H-I, Palmate isochelae. J, Raphidiform - accolada toxa. REVISION OF MICROCIONIDAE FIG. 175. Clathria (Thalysias) hirsuta Hooper & Levi atypical QMG300771. A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Palmate isochelae. F, Raphidiform toxa. TABLE 37. Comparison between Australian and New Caledonian populations of Clarhria (Thalysias) hir- suta Hooper & Lévi. measurements in jum, denoted as range (and mean) of spicule length x spicule width ( Nus» Specimens (N=3) (New Caledonia) | Specimens (N=6) (Queensland) Holotype ce (Caims x 447.812 R3 x2-(48)R Stes 179-(241.8)-284 | 141-216.9)-293 | 163-(204.3)-248 pron pa x15434)5 | xL$-G.D-5 | x 1.8-G.1)-40 74-(100.4)-115| 72-(92.0)- 111 x 1,542.13 | x D8-(1.92,5 34-(51.9)-72 x 2«(3.55 [ChelacI | I | 344.8)-6 | B)-6 3-(4.3).6 6-9, rare Chele l — | 9410:8)-12 | 9411514 | 10-12 rane | agoma auxiliary styles Echinating 37457,8)-73 | 41-(60,4)-78 remm les gr m 4)3 x2,5-(4 4)-8 differ considerably in growth form and geometry of choanosomal styles. Clathria (T.) hirsuta belongs to the juniperina group. These species have choanosomal (coring) megascleres which are only slightly differen- tiated from the subectosomal auxiliary spicules, 4n irregular heavy fibre skeleton, and rhaphidiform toxas, but each species differs in one or more other significant features. More detailed comparisons between C. (T-) hirsuta and other members of the juniperina group, are given hy Hooper & Lévi (19932). Most specimens of Clathria (T) hirsuta have a nearly vestigial ectosomal skeleton, unlike most of the other juniperina species, with ectosomal and subectosomal spicules intermingled in paratangential tracts on the surface, and conse- quently their placement in either Thalysias or Clathria is equivocal. However, the two atypical specimens from SE Australia mentioned above have much better structured ectosomal skeletons than most other known samples (Fig. 175), more reminiscent of the usual Thalysias condition. Moreover, the possession of two categories of auxiliary spicules in most specimens indicates that it belongs with C. (Thalysias), whereas those without specialised ectosomal spicules could be included in Clathria (Clathria). This is further evidence to question the distinction between these taxa at the generic level. MEMOIRS OF THE QUEENSLAND MUSEUM Clathria (Thalysias) juniperina (Lamarck, 1814) (Figs 176-177) Spongia juniperina Lamarck, 1814: 444; Lamarck, 1816: 373. ja juniperina; Hooper & Wiedenmayer, 1994; pot Pasdarr à juniperina; Duchassaing & Michelotti, 1864: 90, pl. 19, fig.3, Not Thalysias juniperina, de Laubenfels, 1936a: 105- 107 (see synonymy for T. virgulipsa below). Rhaphidophlus clathratus; Hallmann, 1912: 209; Top- sent, 1920b: 17-18; Topsent, 1932: 97, pl.5, fig.6, text-fig.3. Not Tenacia clathrata Schmidt, 1870: 56, 80. MATERIAL. LECTOTYPE: MNHNDTS70: SW, cosst of Australia, Peron & Leseur collection. PARALECTOTYPE - MNHNDT3354: same details. HABITAT DISTRIBUTION. Dead coral and rock substrates, shallow subtidal to 10m depth: SW coast WA (Fig. 1763). DESCRIPTION. Shape. Growth form ranging from thickly encrusting to frondose, lamellate, clathrous, with or without free or anastomosing branches. Colour. Bright red to deep red alive, brown dry. Oxcules. Not observed Texture and surface characteristics. Harsh, firm in dry state; surface characteristics range from relatively smooth, even, with white subdermal canals in encrusting forms, to irregularly microconulose or clathrous in more massive forms. Ectosome and subectosome. Ectosomal skeleton crust-like, easily detachable, relatively thin but dense palisade of erect or paratangeritial brushes supported by paratangential tracts of larger sub- ectosomal auxiliary megascleres immediately below surface; peripheral fibres immediately subectosomal with vaguely ascending multi- spicular subectosomal tracts arising to surface. Choanosome. Choanosomal skeleton irregularly reticulate, with very heavy spongin fibres form- ing oval meshes; fibres usually with paucispicular core of subectosomal auxiliary styles occupying only a small proportion of fibre diameter, and fewer choanosomal principal styles which are entirely enclosed in, or project from fibres; in some cases fibres completely uncored, whereas others contain abundant, disorganised auxiliary megascleres; fibres Lypically heavily echinated, some enveloping echinating megascleres entirely, some fibres without echinaling megascleres; mesohyl matrix light, REVISJON OF MICROCIONIDAE FIG. 176. Clathria (Thalysias) juniperina (Lamarck) (lectotype MNHNDT570), A. Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Sinuous, accolada and U-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Lectotype. I, Paralectotype MNHNDT3354. J, Australian distribution. with numerous choanosomal styles dispersed be- tween fibres. Megascleres. Choanosomal principal styles straight or slightly curved near basal end, with smooth, rounded or very slightly subtylote bases. Length 170-(244.4)-280jum, width 9-(10.1)- 12Known only from Australia: m. Subectosomal auxiliary subtylostyles straight or curved, sometimes with multiple curves (sinuous), with smooth subtylote bases. Length 169-(253.5)-310p.m, width 4-(5.4)-6.5,um. Ectosomal auxiliary subtylostyles with prominent subtylote, smooth bases. Length 93- (102.3)- 11Oqum, width 2-(3.7)-4.5jum. Acanthostyles small, stubby, with rounded or only slightly subtylote bases, with few spines and extensive aspinose regions on necks and points; 348 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 177. Clathria (Thalysias) juniperina (Lamarck) (lectotype MNHNDT570). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Palmate isochelae. F, Accolada, sinuous and U-shaped toxas. REVISION OF MICROCIONIDAE spines Jarge, bulbous, erect. Length 45-(57.0)- 65pm, width 5-(6.1)-S1.m. Microscleres. Palmate isochelae unmodified. in- completely differentiated 1nto two size classes; lateral alae entirely fused to shaft, approximately same length as front ala and completely detached from front ala, Length I: 6-(7.1)-9um, length Il: 12-(13.8)-16.5j.m. Toxas very variable in length, mostly thin, ranging from accolada forms with large central curvature and slightly reflexed points, large cur- vature and simply u-shaped, to asymmetrical sinuous forms. Length 55-(122.43)- 180j.m, width 0,8-(1.1)-1.5pm. REMARKS. C. (T.) juniperina differs from that of Hartman (1955), Simpson (19683), Wieden- mayer (1977) and Van Soest (1984h), who in- cluded Spongia juniperina, Spongia virgultosa, Clathria copiosa, Microcioria plana and Clathria clathrata in synonymy, That ‘species’ concept was based on Caribbean populations, with only a single record from the E. Indian Ocean (i.c., nominotypical population of Spongia juniperina) inferring a widely disjunct geographical distribu- tion. Three explanations are possible: 1) W. In- dian Ocean and Caribbean populations are not conspecific. but represent two cryptic sibling species populations with similar morphology; 2) the published province of the original material (‘Indian Ocean, possibly Australia’; Topsent, 1932) is erroneous; 3) the species is widely dis- tributed ‘cosmopolitan’ and these disjunct populations are conspecific. The latter two hypotheses are rejected (specimen labels record one of the types from SW Australia), so the first is considered to be the most probable explanation. The present interpretation is that C. (T) juniperina is restricted to the Indo-west Pacific (and the synonymy given above), whereas the the most senior name for the Caribbean population is C. (T) virgultosa (including several other nominal species in synonymy; see C. (T.) virgul- tosa below). This conclusion conflicts with Topsent's (1932) revision of the relevant (preserved) type material, but this present action is more preferrable than the unlikely alternative that such widely disjunct populations are con- specific. Clathria (T.) juniperina is similar to C. {T} cactiformis and several other species included here in a "juniperina' species complex (spanning the groups Clathria and Thalysias; see comments for C. (T.) cactiformis), which has a depuuperute skeleron (fibres shed some or all of their coring 349 spicules) and principal and auxiliary spicules are similar in geometry, Clathria (Thalysias) kieschnicki Hooper, in Hooper & Wiedenmayer, 1994 Rhaphidephlus cylindricus Kieschnick, 1900 569, pl, fig, 10, Not Esperiopsis cylindrica Ridley & Dendy, 1886: 340 Clurhría (Thalysias) kieschiicki Hooper, in Hooper & Wiedenmayer, 1994; 271. MATERIAL. None. Halotype PMJ missing; (F. Wiedenmayer, pers. comm.). HABITAT DISTRIBUTION. Ecology unknown; Thursday L, Torres Strait (FNQ). DESCRIPTION. Shape. Bifurcate digitate, with short cvlindrical stem, up to 40mm long, branches tapering towards apex, up to 80mm long. Colour, Live colouration unknown, ash-grey in ethanol, Oscules, Small, scattered between surface con- ules, Texture and surface characteristics. Fragile; sur- face with prominent, uregularly distributed con- ules, and detachable skin-like crust. Ectosome and subectosome. Ectosome relatively thick, with discrete plumose bundles of ec- tosoma] auxiliary subtylostyles, forming a con- linuous palisade, below which plumose tracts of subectosomal auxiliary subtylostyles protrude through ectosome. Choanosome. Choanasomal skeletal architecture irregularly reticulate, with heavy, lamellated spongin fibres, 60-160).m diameter, not obvrous- ly divisible into primary or secondary elements, forming ovoid meshes, 90-150jm diameter; fibres cored by irregular multispicular tracts of cheanosómal principal styles and also fewer sub- ectosomal auxihary subtylostyles; echinating acanthostyles abundant, evenly distributed. Megascleres. Choanosomal principal styles straight or slightly curved, with smooth bases. Length 90-180um, width 9-15pm, Subectosomal auxiliary subtylostyles straight, with microspined bases. Length up to 270jum, width upto [8um- Ectosomal auxiliary subtylostyles are fusiform, straight or slightly curved, with microspined bases. Length 135-230j.m, width 4-8um. Acanthastyles subtylote, with bare necks. Length up to 135m, width up to 18yum, Microscleres. Palmate isochelae. Length 15pm. 350 Toxas thin (but of unknown geometry), occur- ring in trichodragmata or singly. Length 70- 100jum. REMARKS. This species is poorly characterised because Kieschnick's (1900) description is brief and does not differentiate it from other arbores- cent, branching Clathria (Thalysias). From his description spicule geometries (which were never figured). and growth form are similar to C. (T.) abietina, but its true affinities remain a mystery given that ihe holotype is missing from PMJ collections. Maurice Burton (note on BMNH1887.5.2.104 specimen label) suggested that it was similar to C. (T) filifera, but any relationship is unsubstantiated. The specific name cylindrica is preoccupied by C. (Axociella) cylindrica (Ridley & Dendy, 1886). Clathria (Thalysias) koltuni Hooper, in Hooper & Wiedenmayer, 1994 Stylotellopsis antarcticus Koltun, 1964a; 66, text- fig.16 Not Anchinoe toxifera antarctica Topsent, 1917: 43. pl.4, fig.5, pl.6, figs. Clathria (Thalysias) kolmi Hooper, in Hooper & Wiedenmayer, 1994: 271. MATERIAL. None: ‘Syatypes’ (ZIL 10637, 11437) (no seen]. HABITAT DISTRIBUTION. Subsirale unknown; 610-860m depth; Budd Coast, Wilkes Land, An- tarctica. DESCRIPTION, Shape. Thinly encrusting, up to only Imm thick. Colour, Red alive, Oscules. Not seen. Texiure and surface characteristics. Even sur- face. Ectosame and subectosome. Erect choanosomal principal subtylostyles protruding through sur- face with bundles of smaller auxiliary subtylos- tyles dispersed around principal spicules. Choanosome. Hymedesmoid, with choanosomal principal and subectosomal auxiliary subtylos- 1yles erect on basal spongin: bundles of echinat- ing acanthostyles clumped around erect structural megascleres. Megascleres. Choanosomal principal subtylo- styles long, straight, fusiform, with subtylote bases and evenly microspined in basal third of spicule. Length 400-750pm, width 26-364m. Ectosomal and subectosomal auxiliary styles very long, slender, straight, fusiform, with MEMOIRS OF THE QUEENSLAND MUSEUM microspined subtylote bases. Length 430- 630m, width 8-121um. Echinating acanthostyles short, club-shaped, fusiform, with prominent subtylote base and evenly spined over entire length of spicule, Length 100-260pm, width 10-14m. Microscleres, Absent. REMARKS. This species is a lipochelous Clathria, but it is uncertain from Koltun’s (1964a) brief description what subgenus it belongs to. It is retained in Thalysias (following Hooper & Wiedenmayer, 1994). It differs from other hymedesmoid species (especially those pre- viously referred to Pseudanchinoe), in spicule dimensions and spicule omamentation, but has few other noteworthy features. antarcticus 1s preoccupied by C. (M.) antarctica (Topsent, 1917). Clathria (Thalysias) lendenfeldi Ridley & Dendy, 1886 (Figs 178-179, Plate 7B-E) Clathria lendenfeldi Ridley & Dendy, 1886: 474; Rid- ley & Dendy, 1887: 148, pl.28, fig.5. pl.29, fig.6, p.47, fig.5; Whitelegge, 1889: 186; Whitelegge, 1901: 86; Whitelegge, 1907: 492-494; Burton & Rao, 1932: 334; Rudman & Avern, 1989: 335; Hooper et al., 1990: 126-133, figs 1, 2, 4, 6; Hooper & Wiedenmayer, 1994: 271. Not Clathria lendenfeldi; Brondsted, 1934: 19-20, text- fig.19. Thalysias lendenfeldi; de Laubenfels, 19363; 105. Micraciona lendenfeldt, Dawson, 1993; 37, Spongia abietina, in part, Lamarck, 1814: 450. Echinonema anchoratum var. lamellosa; Whitelegge, 1901: 82. Not Echinanema anchoratum var. lamellosa Lenden- feld, 1888: 219, Clathria spirans Hallmann, 1912; 210; Dendy, 1922; un pl.5, fig.2, pl.13, fig.4a-f; Burton, 19594: Pia diechinata Hallmann, 1912: 211; 1914a: 268 [nomen nudum]. Thalysias spicata; de Laubenfels, 1936a: 105. Clathria whiteleggii Dendy, 1922; 67, pl.7, fig.1, pl. 13, figs 5a-f; Burton, 1931a: 344-345; Burton, 19592: 245; Lévi, 1963: 66. Thalysias whiteleggei; de Laubenfels, 1936a: 105, Clathria coppingeri var. aculeata Hentschel, 1912: 3, Rhaphidaphilus bispinosus Whitelegge, 1907: 503- 504. Clathria bispinosa; Hallmann, 1912: 177,211. cf. Microciona prolifera; Vosmaer, 1935a: 610, 636, 669. MATERIAL. HOLOTYPE: BMNH1887.5.2. 107: off Port Jackson, NSW, 33?40'S, 151"40'E, HMS REVISION OF MICROCIONIDAE 100 um FIG. 178. Clathria (Thalysias) lendenfeldi Ridley & Dendy (NTMZ2095). A, Choanosomal principal subtylos- tyles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthos- tyles. E, Wing-shaped and accolada toxas. F, Palmate isochelae. G, Australian distribution. H, Section through peripheral skeleton. I, Trawled specimens from NW Australia. "Challenger' (dredge). OTHER MATERIAL (refer to Hooper et al., 1990 for list of additional specimens used in this study): QLD - QMG303025, QMG303039, QMG304777, QMG305138, QMG300784, QMG- 300830, QMG303507, QMG303523, QMG304946. NT - NTMZ2821. WA - NTMZ3060, NTMZ3384, QMG310535 (NCIQ66C-1518-Q) (fragment NTMZ3489), QMG310423 (NCIQ66C- 1291-T) (frag- ment NTMZ3463), QMG310423 (NCIQ66C-1318-X) (fragment NTMZ3468). TAS - QMG311436 (NCIQ- 66C-3745-M) (fragment NTMZ3822). ANDAMAN SEA, THAILAND - NTMZ3657, NTMZ3659. RED SEA - PIBOC04-11 (fragment QMG300057). SOMALIA, EAST AFRICA - PIBOCB 12-367 (frag- ment QMG2300062). HABITAT DISTRIBUTION. Rock reefs and dead coral heads; intertidal to 108m depth; widespread Indo- Pacific; Port Jackson, Botany Bay (NSW); Shelburne Bay, Howick Is, Direction Is, Gulf of Carpentaria (FNQ); Cairns, East Frankland Is, Pandora Bay (NEQ); Darwin Harbour, Bynoe Harbour, Melville I., Beagle Gulf, Port Essington, Cobourg Peninsula, Cootamundra Shoals, Wessel Is (NT); Broome, Port Hedland, Bedout I., Dampier Archipelago, Monte Bello Is, Exmouth Gulf, Northwest Cape, Amphinome Shoals, Northwest Shelf (WA); Bicheno (Tas)(Fig. 178G); also Gulf of Manaar (Burton & Rao, 1932), Aru Is, Indonesia (Hentschel, 1912), Cargados Carajos (Dendy, 1922), Gulf of Aden (Burton, 1959a), Arabian coast (Burton, 1959a), Andaman Sea (present study), 352 MEMOIRS OF THE QUEENSLAND MUSEUM (5 mcd FIG. 179. Clathria (Thalysias) lendenfeldi Ridley & Dendy (A-B, NTMZ2701; C-G, QMG303039). A, Choanosomal skeleton. B, Fibre characteristics (x424). C, Echinating acanthostyles. D, Acanthostyle spines. E, Bases of principal and auxiliary subtylostyles. F, Wing-shaped and accolada toxas. G, Palmate isochelae. REVISION OF MICROCIONIDAE 35 Red Sea and Somalia (present study), Saya de Malha (Dendy, 1922) to the Natal coast (Burton, 19313). DESCRIPTION. (See Hooper et al., 1990). DIAGNOSIS. Variable growth form ranging from bushy clathrous reticulate-branching to bushy lamellate planar digitate fans, usually with woody basal stalk and cylindrical branches; vivid red to pale red-brown alive in shallow waters to slightly turquiose or unpigmented in deeper waters; oscules small, congregated into special pore areas on points of digits or scattered between surface processes; surface usually micro- conulose; ectosome ranging from sparsely scat- tered smaller auxiliary spicules to dense, erect, continuous crust; choanosomal principal sub- tylostyles also protrude through surface; subec- tosomal skeleton poorly developed, paratangential, composed of larger auxiliary sub- tylostyles: choanosomal skeleton irregularly reticulate, heavy spongin fibres divided into primary (ascending) and secondary (transverse) components, producing regular or irregular meshes: fibre-meshes heavier in axis; fibres generally uncored, some with uni- or pauci- spicular tracts of principal spicules, and abun- dantly echinated by both acanthostyles and principal subtylostyles especially at fibre junc- tions (*spicate’); choanosomal principal sub- tylostyles long. curved or straight, sharply pointed, usually with heavily spined bases (119- (229.6j-49211m x 1.8-(12.9)-35u m); subec- tosomal auxiliary subtylostyles long, straight, fusiform, slightly subtylote, microspined bases (136-(241.5)-404um x 2-(4.6)-I5um); ec- tosomal auxiliary subtylostyles short, thin, straight, fusiform, with microspined subtylote bases (62-(123.4)-198ym x 2-(3.0)-10pm); acanthostyles slender, long or short, fusiform, slightly stbtylote. evenly and lightly spined, spines small, recurved (49-(88.1)-151j.m x 2- {6.4)-14,4m); palmate isochelae small, rarely modified, narrow lateral alae completely fused to shaft, approximately same length as front ala; lateral alae completely and widely separated from front ala (6-(12_5}-25j.m long); toxas accolada to wing-shaped, yery thin (hair like); larger toxas usually accolada, straight, with slight but sharp angular central curvature and unreflexed arms; smaller toxas usually wing-shaped, with large central curvature and slightly reflexed arms; toxas found singly and in bundles (dragmata) (7-(136.4)-361 um x 0,4-(1.3)-3.6 um). A REMARKS. Variation has been comprehensive- ly investigated from many living specimens and type material (Hooper et al., 1990), The synonymy above also includes several new synonyms added to the species since that earlier paper. This species is a cryptic sibling species of CAT) major. differing only substantially by its hair-like toxas, sharply pointed auxiliary megascleres, statistical differences in spicule dimenstons and various biochemical features (Hooper ct al., 1990), and both are members of Hallmann's (1912) ‘spicata’ group. Clathria (Thalysias) major Hentschel, 1912 (Figs 180-181) Clathria frondifera var, major Hemschel, 1912: 36]. Clatheia (Thalysias) major; Hooper et al., 1990: 133- 135, figs 1, 3, 5, 6; Hooper & Wiedenmayer, 1994; 272. MATERIAL. HOLOTYPE: SMF977 (fragment MNHNDCL2303): Straits of Dobo, Aru L, Indonesia, 6°S, 134^ 50" E, 40m depth, 20.ii1.1908, coll. H, Merton (dredge), OTHER MATERIAL: (Hooper et al., 1990) WA - NTMZ3338, NTMZ3360, CSIROEMGOOL. HABITAT DISTRIBUTION. Rock reefs and dead coral heads; intertidal to 82m depih; Bedout L, Port Hedland, Mary Anne I., Direction Is, Exmouth Gulf (WA): Bynoe Harbour, Darwin Harbour, Port Es- sington (NT) (Fig. 180H); also Aru Is, Indonesia (Hentschel, 1912], DESCRIPTION. (See Hooper et al., 1990). DIAGNOSIS. Vanable growth forms ranging from low, foliose, bushy, subspherical, clathrous digitate, to flabellate or digitate Fans, usually with Jong basal stalk and flattened or irregularly cylindrical branches; bright red to orange-red alive; oscules small, congregated into special pore areas on lateral sides of branches or between surface conules; flabellate specimens may have Phakellia-like pores grouped into stellate pore- areas; surface irregularly microconulose with close-set subdermal ridges and stnations; ec- tosomal skeleton ranges from very few tangen- tially placed ectosomal auxiliary subtylostyles to dense, erect or paratangential brushes of ec- tosomal spicules; subectosomal skeleton umose, paratangential tracts of larger auxiliary subtylostyles; both larger auxiliary and principal spicules protrude through surface singly or in plumose bundles; choanosomal skeleton ir- regularly reticulate; fibre characteristics, skeletal siructure and distribution of megascleres and microscleres identical to C. (T) lendenfeldi; 354 MEMOIRS OF THE QUEENSLAND MUSEUM 100 um FIG. 180. Clathria (Thalysias) major Hentschel (specimen NTMZ858). A, Choanosomal principal subtylostyle. B, Intermediate principal-echinating subtylostyle. C, Echinating acanthostyles. D, Subectosomal auxiliary styles/ quasi-tornote. E, Ectosomal auxiliary style/ quasi-tornote. F, Wing-shaped and accolada toxas. G, Palmate isochelae. H, Australian distribution. I, Section through peripheral skeleton. J, Trawled specimens from NW. Australia. choanosomal principal styles thick, slightly curved, fusiform, rounded or subtylote, usually with microspined bases, sometimes smooth (187- (250.5)-38p. m x 5-(15.1)-361.m); subectosomal auxiliary subtylostyles long, straight, fusiform pointed, usually subtylote, microspined bases, or commonly with rounded apex (quasi-tornotes) also bearing microspines (156-(287.8)-4391.m x 2-(5.7)-14jum); ectosomal auxiliary subtylos- tyles short, straight, subtylote microspined bases, usually with rounded apex (quasi-tornotes) and terminal spines (84-(136.8)-193,1.m x 2-(3.8)- 9pm); acanthostyles relatively slender, long or short, fusiform, subtylote, with large spines on base and apex but nearly aspinose 'neck' (77- (112.7)-144m x 3-(7.8)-15jm); palmate isochelae small, unmodified, wide lateral alae completely fused to shaft, approximately same REVISION OF MICROCIONIDAE m tA m FIG. 181. Clathria (Thalysias) major Hentschel (QMG300153). A, Choanosomal skeleton. B, Fibre charac- teristics. C, Echinating acanthostyles. D, Acanthostyle spines. E-F, Bases and apex of principal and auxiliary subtylostyles. G, Palmate isochelae. H-1, Accolada toxas. 356 MEMOIRS OF THE QUEENSLAND MUSEUM 100 um i FIG. 182. Clathria (Thalysias) michaelseni (Hentschel) (fragment of holotype SMF969T). A, Choanosomal principal subtylostyles. B, Echinating acanthostyles. C-D, Subectosomal auxiliary subtylostyles and polytylote forms. E, Oxhorn - U-shaped toxas. F, Sigmoid anchorate-like isochelae. G, Section through peripheral skeleton. H, Australian distribution. length as front ala; lateral alae completely separated but close to front ala (6-(10.4)-161.m long); toxas accolada or wing-shaped, the former long, very thick, with large rounded central cur- vature, straight or reflexed arms, the latter short, thin, widely curved at centre with reflexed arms (27-(108.9)-390j.m x 0.6-(2.3)-5 2m). REMARKS. The species is a cryptic sibling of C. (T.) lendenfeldi with a sympatric but more restricted distribution. In gross morphology, sur- REVISION OF MICROCIONIDAE face features, live colouration and in many of its skeletal characters C. (T:) major is indistinguish- able from its sibling. However, they can be reliab- ly differentiated by spines on the points of many of the auxiliary spicules (especially most of the smaller ones), marginally thicker and longer toxas, and statistically (but not absolute) larger size of most other megascleres in C. (T.) major. The importance of these apparently 'relatively minor' morphological differences is indicated by clear differences between the two species in their biochemical fingerprints (Hooper et al., 1990). In a recent survey of several Western Australian species, C. (T:) major was found to contain sig- nificant quantities of the chemical 2,6- dibromophenol of potential commercial impor- tance as an ‘iodoform’ or ‘fresh sea’ flavour used in the production and marketing of commercial prawns (F. Whitfield, CSIRO, pers.comm.). Clathria (Thalysias) michaelseni (Hentschel, 1911) (Fig. 182) Hymeraphia michaelseni Hentschel, 1911: 351-352, text-fig.34; Hentschel, 1912: 385. Damoseni michaelseni; de Laubenfels, 1936a: 110. Clathria michaelseni; Hooper & Wiedenmayer, 1994: 272. MATERIAL. HOLOTYPE: HM (not seen). Fragment of holotype SMF969T: 5 km NW. of Denham, Shark Bay, WA, 25?52'S, 113?28'E, 3m depth, 12.vi.1905, coll. W. Michaelsen & R. Hartmeyer (dredge). HABITAT DISTRIBUTION. Bivalve and worm tubes, sand, coral and Halimeda bed substrata; 3-14m depth; central W coast (WA) (Hentschel, 1911); also Arafura Sea (Hentschel, 1912) (Fig. 182H). DESCRIPTION. Shape. Thinly encrusting. Colour. Live colouration unknown, brown in ethanol. Oscules. Unknown. Texture and surface characteristics. Firm; smooth unornamented surface. Ectosome and subectosome. Star-shaped plumose brushes of intermingled ectosomal and subectosomal auxiliary subtylostyles on surface; most auxiliary spicules perpendicular to surface, with choanosomal principal megascleres protrud- ing through. Choanosome. Hymedesmoid skeletal structure, with choanosomal principal subtylostyles and smaller echinating acanthostyles embedded in and perpendicular to basal spongin; mesohyl matrix moderately heavy, without detritus. Megascleres. Choanosomal principal subtylos- tyles long, fusiform, slightly curved, subtylote, with microspined bases. Length 188-(381.6)- 646um, width 10-(14.6)-191.m. Subectosomal auxiliary subtylostyles long, thin, fusiform, prominently subtylote, lightly microspined, occasionally with smooth bases. Length 307-(403.6)-482,.m, width 3-(4.4)-61.m. Ectosomal auxiliary subtylostyles, short, thin, straight, prominently subtylote, smooth bases, usually polytylote shafts. Length 141-(162.4)- 197m, width 2-(2.9)-4j.m. Acanthostyles long, thin, slightly subtylote, with lightly microspined base and central por- tions, aspinose points and neck regions. Length 96-(108.8)-125p.m, width 3-(6.6)-8 p.m. Microscleres. Isochelae sigmoid (bidentate) anchorate, with small alae attached only at their bases. Length 15-(17.2)-191.m long. Toxas oxhorn or u-shaped, variable in size, relatively thick, gently curved at centre and with reflexed points or only slightly reflexed points. Length 38-(122.6)-239j.m, width 1-(3.5)-6j.m. REMARKS. Hentschel (1911) initially over- looked the presence of toxas in this species, al- though later described by him in specimens from Aru Is, Indonesia (Hentschel, 1912), but these were also seen in the holotype redescribed above. Hentschel (1911, 1912) also overlooked the presence of two categories of auxiliary spicules indicating its assignment in C. (Thalysias) rather than C. (Clathria). Spicule dimensions seen in type material also vary slightly from those pub- lished by Hentschel (1911). This species is well differentiated from other thinly encrusting (hymedesmoid) microcionids in having bidentate sigmoid isochelae, for which de Laubenfels (1936a) created Damoseni. The recognition of de Laubenfels’ genus is not upheld since this feature is homoplastic, also known to occur in other microcionids (e.g., C. (C.) nexus Koltun, with an erect ramose growth form), and other poecilosclerids (e.g., Strongylacidon stel- liderma Carter). Clathria (Thalysias) phorbasiformis sp. nov. (Figs 183-184, Plate 7F-G) MATERIAL. HOLOTYPE: NTMZ2138: Dudley Point, East Point Aquatic Life Reserve, Darwin Har- bour, NT, 12?25.0'S, 130°49.1'E, intertidal, 27.1x.1984, coll. J.N.A. Hooper. PARATYPES: NTMZ2203: same locality, 23.xi.1984. NTMZ2418: same locality, 12?24.5'S, 130?48.0'E, 3m depth, 14.viii.1985, coll. J.N.A. Hooper (snorkel). OTHER MATERIAL: NT - NTMZ2214: same locality as type 358 MEMOIRS OF THE QUEENSLAND MUSEUM er ME Ny e el [ ) f p 100 um e D FIG. 183. Clathria (Thalysias) phorbasiformis sp.nov. (holotype NTMZ2138). A, Choanosomal principal subtylostyles. B, Echinating acanthostyles. C, Subectosomal auxiliary subtylostyle. D, Ectosomal auxiliary subtylostyles. E, Wing-shaped and accolada toxas. F, Palmate isochelae. G, Section through peripheral skeleton, H, Australian distribution. I, NTMZ2237. J, Acanthostyles incorporated into spongin fibres. material, 8.1.1985, QMG300149 (fragment coral boulders or in crevises and pools; intertidal, Dar- NTMZ2223), QMG300150 (fragment NTMZ2237). win Harbour (NT) (Fig. 183H). HABITAT DISTRIBUTION. Encrusting on laterite , : rock, dead coral, exposed at ELWS tides, under dead DESCRIPTION. Shape. Thickly encr usting, 0.7- 1.3cm thick, producing thin cylindrical REVISION OF MICROCIONIDAE 359 FIG. 184. Clathria (Thalysias) phorbasiformis sp.nov. (QMG300150). A, Choanosomal skeleton. B, Fibre characteristics (x400). C, Echinating acanthostyles. D, Acanthostyle spines. E-G, Bases of principal and auxiliary subtylostyles. H, Palmate isochelae. I, Wing-shaped and accolada toxas. 360 stoloniferous digitate non-anastomosing stoloniferous processes, up to 6mm in diameter, which may or may not re-attach to substrate. Colour. Orange-brown to brick-orange alive (Munsell 2.5 YR 7/6-8); grey-brown in ethanol. Oscules. No oscules visible optically alive or preserved, but numerous minute pores, 0.1- 0.25mm diameter, scattered over surface seen at higher magnification. Texture and surface characteristics. Texture firm, compressible; abundant clear mucus produced upon exposure to air; surface optically smooth, microscopically hispid, even, or small ridges and low conules following contours of substrate; digitate surface processes may have more prominent sculpturing superficially resembling C. (T.) reinwardti. Ectosome and subectosome. Usually dense ec- tosomal skeleton, only slightly opaque or pellucid between surface conules, with subectosomal cavities and canals barely visible below surface (alive); ectosome microscopically hispid, with points of choanosomal principal styles protrud- ing, singly or in plumose brushes, with thickest brushes in areas where ultimate choanosomal fibres in peripheral region closest to surface; specialised ectosomal skeleton well developed, with discrete brushes of smaller ectosomal auxiliary subtylostyles forming continuous palisade; relatively thick but variable layer of spongin and detritus also on ectosome; clear regional and structural differentiation between ectosomal and subectosomal skeletons; subec- tosomal region variable in thickness dependent on proximity of peripheral fibres to surface, con- taining plumose columns of larger auxiliary sub- ectosomal subtylostyles, not associated with fibres, but often bound together by collagen; sub- ectosomal skeletal columns originate from ends of choanosomal principal megascleres, which in turn echinate fibres of peripheral skeleton in plumose tufts or singly. Choanosome. Thick growth forms — Skeletal ar- chitecture vaguely plumo-reticulate; spongin fibres relatively light, irregularly anastomosing, fully cored by acanthostyles, lying in rows of 3-5 megascleres abreast (i.e., entirely incorporated into spongin fibres lying in parallel spicule tracts); principal subtylostyles only rarely seen coring fibres, and acanthostyles only rarely echinate fibres (i.e., lie at right angles to fibres); spongin fibres predominantly echinated by prin- cipal subtylostyles, in plumose tufts or singly, particularly abundant at fibre nodes; fibre anas- tomoses form oval or elongate meshes, 250-800j.m MEMOIRS OF THE QUEENSLAND MUSEUM diameter; fibres thicker in deeper areas of choan- some (70-100jp.m diameter) than in periphery (55-80j.m diameter); major portion of branch diameter consists of extra-fibre plumose tracts of subectosomal auxiliary megascleres with choanosomal reticulate skeletal comprising less than half of branch diameter; extra-fibre plumose spicule tracts originate approximately half-way along length of perpendicular choanosomal styles, or in thicker sections they originate at ends of principal megascleres; extra-fibre plumose spicule tracts ascend to, diverge, and pierce ec- tosomal skeleton; mesohyl matrix only lightly pigmented, variable in density, usually heavier near periphery; extra-fibre spicules mostly occur in well defined tracts with few scattered random- ly throughout mesohyl. Thinly encrusting growth forms — hymedes- moid skeletal construction with basal layer spon- gin lying on substrate, uncored but very heavily echinated by both acanthostyles and choanosomal subtylostyles standing perpen- dicular to substrate; subectosomal spicule tracts arise from distal half of erect choanosomal megascleres, diverging and ascending to surface in plumose brushes, surmounted by plumose brushes of ectosomal auxiliary spicules at periphery. Megascleres. Choanosomal principal subtylos- tyles fusiform, tapering to long points, slightly curved near the basal end or occasionally straight, with subtylote, mostly smooth bases, occasional- ly roughened subapically or slightly tubercular. Length 245.2-(425.8)-583.lum, width 10.2- (19.5)-33.8 um. Subectosomal auxiliary subtylostyles long, fusiform, mostly straight, with subtylote microspined or occasionally smooth bases. Length 275-(386.2)-485.3um, width 4.0-(9.6)- 18.8um. Ectosomal auxiliary subtylostyles relatively short, straight or whispy, slender, with subtylote, relatively heavy basal microspination. Length 70.0-(140.5)-261.2jum, width 1.2-(4.0)-6.91.m. Acanthostyles fusiform, straight or slightly curved near base, slightly subtylote, evenly spined but characteristically free of spines at points; spines large, recurved. Length 95.4- (115.4)-132.4.m, width 4.6-(8.4)-12.7 um. Microscleres. Palmate isochelae abundant, sub- divided into 2 size categories, smaller ones some- times contort; lateral alae completely attached to shaft, approximately same length as front ala but completely detached from it. Length I: 10-(14.9)- 2] .9j.m, length II: 2.5-(5.6)-9.21.m. REVISION OF MICROCIONIDAE Toxas moderately abundant, vaguely separated into 2 forms although intermediates occur: smaller wing-shaped toxas relatively thick, generously curved at centre with slightly reflexed, abruptly pointed ends; accolada toxas long, nearly straight, with slight central curvature and slight or no apical reflexion. Length 30- (85.1)-222.91.m, width 0.5-(1.4)-2.5j m. Associations. Growing in dense clumps amongst algae (Gellidium), with stoloniferous branches intertwined, occasionally attached to algae itself; some specimens growing over, or next to other encrusting sponges (Placospongia, Mycale, Antho (Plocamia)), compound ascidians, and coralline algae. ETYMOLOGY. Like Phorbas (Anchinoidae), REMARKS. This species incorporates most echinating acanthostyles into spongin fibres. either together with one or few principal sub- tylostyles enveloped by spongin, or excluding principal megascleres completely. This feature is consistent except for one thinly encrusting specimen that lacks a reticulate fibre skeleton. in which case acanthostyles stand perpendicular to substrate. Principal spicules. are mostly outside fibres, perpendicular to (echinating) fibres. and fibre nodes, and protruding through the surface, This is reminiscent of Hallmamn’s spicata group (see C. (T.) coppingeri). Incorporation of echinaung acanthostyles secondarily into fibres has been observed in some specimens of C. (Dendrocia) dura, C. (D.) imper- fecta, and to a lesser degree C, (D) myxilloides, but these instances are infrequent. inconsistent (Le., seen in some sections of the skeleton hut not in others), and probably aberrant. À similar phenomenon has been described for C. (T. ) orien- talis by Brondsted (1934) but this too is atypical for the species (whereby the larger auxiliary sub- tylostyles usually core fibres). Analogous struc- lures are described in other poecilosclerids, particularly for the families Anchinoidae and Crellidae, but in these species acanthostyles also comprise the ‘principal’ structural spicules. Clathria (Thalysias) phorbasiformis differs from other species in the "phorbasiformis' com- plex in gross morphology and spicule geometry. In live surface features and colouration it some resemblance to thickly encrusting C. (T) rein- wardti, although spicule geometry, spicule size, skeletal architecture and fibre characterisilics are clearly different between them, 361 Clathria (Thalysias) placenta (Lamarck, 1814) (Figs 185-186) Spongia placenta Lamarck, 1814;:374; 1815:356. Wilsonella placenta, Topsent, 1930:24, pl.3, fig.8. Clathria placenta; Hooper & Wiedenmayer, 1994: 273. Not Microciona placenta, de Laubenfels, 1954;146- 147, text-fig.94. MATERIAL. HOLOTYPE: MNHNDTS552: King I Bass Strait, Tas, 39°50'S, 144°00"E, Peron & Lesweur collection. HABITAT DISTRIBUTION. Ecology unkranwn; Bass Strait, Tasmania (Fig. 185G). DESCRIPTION. Shape. Thick, flabellate growth form, 180mm high, 170mm wide, up to 10mm thick, with even margin; probably onginally with basal stalk but now detached. Colour. Grey-brown dry. Oscules. Not seen. Texture and surface characteristics. Harsh, flexible. brittle in dry state; surface relatively even, with longitudinal annular striations running from basal stalk to margin of fan, and raised fibre reticulations forming polygonal pattem. Eclosome and subectosome. Ectosome almost completely detached fromdry type specimen, but where present appears to be sparse, plumose, erect or paratangential palisade of ectosomal styles arising from ascending subectosomil spicule tracts, the latter embedded in peripheral skeleton; choanosomal fibres immediately subec- tosomal. Choanosome. Choanosomal skeleton irregularly reticulate with primary (ascending) and secon- dary (transverse) fibres; primary fibres (105- ViSyum diameter) cored by multispicular tracts of subectosomal auxiliary styles, occupying up to EE fibre diameter, tracts becoming plumose peripherally; secondary fibres (35-88m diam- eter) without coring spicules; all fibres heavily echinated by small acanthostyles sometimes nearly enveloped in spongin; fibre anastomoses farm irregular oval and rectangular meshes ( 145- 510um diameter): mesohyl matrix light, with few loose megascleres dispersed between fibres. Megascleres. Choanosamal principal megascleres absent or completely undifferen- tiated from subectosomal spicules. Subectosomal auxiliary styles thin. straight. slightly curved or slightly sinuous, with smooth rounded bases and sharp fusiform points. Length 175-(237.5)-285p.m. width 5-(6.6)-Sjum. 362 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 185. Clathria (Thalysias) placenta (Lamarck) (holotype MNHNDTSS52). A, Subectosomal auxiliary subtylostyle. B, Ectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Accolada toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype. Acanthostyles short, slender, subtylote, Ectosomal auxiliary styles straight or slightly fusiform pointed, spined only on base and near curved near apical end, with rounded apical end, with smooth regions at "neck" microspined bases, fusiform points. Length 115- (proximal to base) and point. Length 52-(54.4)- (138.1)-156j.m, width 2-(2.6)-4j.m. 58m, width 3.5-(4.2)-6.m. REVISION OF MICROCIONIDAE FIG. 186. Clathria (Thalysias) placenta (Lamarck) (holotype MNHNDT552). A, Skeleton, ectosome detached. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Palmate isochelae. F, Accolada toxas. Microscleres, Palmate isochelae small, with some contort forms, small alae less than 30% of shaft length, lateral alae completely fused to shaft, front ala completely detached Length 8- (10.7)-141um., Toxas accolada, moderately long, thick, with only very slight central curvature and straight points. Length 105-(119,5)-148jum, width 1.0- (1.4)-2-0u m. REMARKS. Lamarck's (1814) holotype from Buss Strait and de Laubenfels'(1954) sample USNM22908 from Truk, Caroline Is are not con- specific; the latter becomes C, (T.) lematolae sp. nov. The Truk sample is only similar to C. placen- tü in having fibres cored by subectosomal auxiliary spicules instead of principal spicules (i.c., the principal and larger auxiliary spicules are undifferentiated in their geometry). In most other details the two species can be readily dis- tinguished (the Chuuk sample has an encrusting growth form, an extremely smooth surface, skele- tal structure is hymedesmoid including posses- sion of a very extensive subectosomal skeleton, occupying almost half of ihe sponge diameter, acanthostyles are about twice the size of those in C. placenta with much more robust spination, toxas are slightly accolada but moreso wing- shaped, megascleres are mostly subtylote, and dimensions of most spicules differ). Topsent (1930) implied that C. (T) placenta) was similar to C, (Wisonella) australiensis (Caner), but this is certainly not true (the two having very different skeletal structures, spicule geometries and absence of foreign detritus in the skeleton of C. (T) placenta). Clathria (T) placen- tais a member of the ‘juniperina’ complex having à reduced spicule skeleton (whereby fibres shed some or all their spicules, in this case only from the secondary fibres), and principal and auxiliary spicule of similar geometry (refer to discussion under C. (T) cactiformis). Clathria (Thalysias) procera (Ridley, 1884) (Figs 187-188, Table 38) Rhaphidophlus procerus Ridley, 1884a:451-452, po. fig k, pl.42, fig.o; Burton, 19312:343, pl.23, ig.2. Clarhieia procera; Dendy, 1922:64, pl.2, figs 6-7; Bur- ton, 193823:28-29; Burton, 195943:243; Lévi, 1963:66; Bergquist, 1967:164-165, text-fig.3; Thomas, 1973:34-35, pl.2, fig.5, pl,7, fig.3; Heraquist, 1977:65; Hooper & Wiedenmayer, | 994: 273, MEMOIRS OF THE QUEENSLAND MUSEUM Tenacie procera; Burton & Rao, 1932:340; Burton, 19344:550; Burton, 1934b:28. Rhaphidophlus spiculosus Dendy, 1889b:75, 86, 87, 59, pt fig.4 [Gulf of Manaar, Ceylon]; Dendy, 1922:64. Clathria spiculosa; Dendy, 1905:171-173, pl.8, hg.2 [Gulf of Manaar, Ceylon]; Hentschel, 1912:363,364; Hallmann, 1912:177; Dendy, 1916a:46, 95, 128-129 |Okhamandal, Kattiawar]. Clathria spiculosa var. ramosa; Hentschel, 1912:363- 364, Not Clathria spiculosa var. macilenta, Hentschel, 1912:364 [Aru L, Arafura Sea]. Echinanema gracilis Ridley, 18843:617, pl.54, fiel; Dendy, 1922:64. Rhaphidophlus gracilis; Ridley & Dendy, 1887:152, 242, 252, Topsent, 1892b:24. Clathria ggacilisi Dendy, 1905:171; Vosmaer, 1935a:634. Not Rhaphidaphlus arborescens Ridley, 18842:450- 451, pl.40, fig.L, pl42, tig.n; Burton & Rau, 1932:340. cf. Microciona prolifera; Vosmaer, 1935a:610, 634, 669. MATERIAL. HOLOTYPE: BMNH1882.2.23. 315: Off East Point, Port Darwin, NT, 12724,5'8. 130?48.0'E, 14-22m depth, coll. HMS ‘Alert’ (dredge), PARATYPE: BMNH18822.23,31]: same locality. HOLOTYPE of R. spleulosus: BMNH 1889, 1.21,5 (fragment BMNH1954. 2.23.101): Gull'of Manaar, Sri Lanka, 8°N, 78°E, PARATYPE of R, spiculosus: BMNH1887. 8.4.31: same locality. HOLOTYPE of E. gracilis BMNH1882,10,17,111: Providence Island, Seychelles Is, Indian Ocean, PIFS, 51°02'E, 48m depth (dredge), HOLOTYPE of C. spiculosa var. ramosa: SMF1698 (fragment MNHNDCL 2304): Straits of Dobo, Aru ly Arafura Sea, Indonesia, 6°S, 134°50°E, 20.11.1908, 40m depth coll. H. Merton (dredge). OTHER MATERIAL: QLD- NTMZ3983, QMG 301032, QMG303514- 0MG304392, QMG304771. NT- NTMZ2604, QMG303582. WA- NTMZ1308. INDIAN OCEAN - BMNH1907.2.1.63, BMNH 1954.2.23.113, BMNHi954.2.23.114, HABITAT DISTRIBUTION. On loose, soft substrates (sand, mud. gravel, shell grit) associated with shallow- waler or deeper offshore reeís; 11-78m depth; widespread throughout the Indian Ocean and Indo- west Pacific: Gulf of Carpentaria, Low Is, Direction Is, Snake Reef, Turtle Is (FNQ); Bynoe Harbour, Darwin Harbour, Cape Wessel, Arafura Sea (NT); Port Hed- land (W A)(Fig. 187H); also Scottburgh, Natal (Burien, 1931a; Lévi. 1963), Cargados Carajos, Seychelles, Amirante, Red Sea and Arabian Sca (Ridley, 18842; Ridiey & Dendy, 1887; Dendy, 1922; Burton & Rao, 1932; Burton, 19593; Thomas, 973b); Tuticorin, Cape Comorin, Palk Straits, and Madras Straits, Gulf of Manaar (Burton & Rao, 1932; Burton, 19383), Aru Is, Indonesia (Hentschel, 1912), Hawaii (Bergquist, 1967; 1977) REVISION OF MICROCIONIDAE 365 100 um FIG. 187. Clathria (Thalysias) procera (Ridley) (NTMZ1308). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Wing-shaped and U-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype BMNH1882.2.23.313. DESCRIPTION. Shape. Long, single or bifurcate cylindrical digits, whip-like, very slightly flat- tened laterally, 230-640mm long; stalk tapers in both directions from thick central region 4-14mm diameter, to woody base 4-6mm diameter, and rounded points 6-13mm diameter; apex with single or no bifurcation is single; point of attach- ment to substrate expanded, rhizomous; gross morphology superficially resembles Junceela gorgonian whip-coral. 366 TABLE 38. Comparison between present and published records of spicule dimensions for Clathria (Thalysias) procera (Ridley). (N=25). Clathria spiculosa var. ramosa SMF1698 Holotype (BMNH1882.2. 3.313) SPICULE 322-(334.2)- 248-(283.2)- Choanosomal| 348x11- 309x16- | 263-288 x 15- styles (13.2)-16 (18.2)-22 18 (rare) (common) (uncommon) | MEMOIRS OF THE QUEENSLAND MUSEUM diverging) towards periphery; fibres imperfect- ly divided into ascending primary fibres (40-75jm diameter) and transverse secondary components (130-210um diameter); primary fibres multi- spicular, cored by subec- tosomal auxiliary subtylostyles occupying up Measurements in pm (N-1) 201-310 x 4-12 318-(334.2)- | 342-(367.1)- Subectosomal| 58,8 (92). | 393x8-(11.3)-| ^52" styles 1i å 9.2-12.5 172-(258.4)- 295 x 3-(5.6)-7 124-(178.2)- 290 x 2-(3.8)-5 112.3-235.6 x 2.2-5.8 Ectosomal styles to 90% of fibre diameter; secondary fibres less heavi- ly cored, occasionally unispicular; fibre reticula- tion producing irregularly 210-294 x 4-8 x 0.8-(1.3)-1.5 | x0.9-(1.2)-1.8 Colour. Pale orange alive (Munsell 5YR 8/6), pale grey in ethanol. Oscules. Not visible in either live or preserved specimens. Texture and surface characteristics. Firm, only very slightly compressible but flexible, with ob- vious stiff axis; basal region woody, more rigid than central or apical regions; surface optically smooth, without conules or other surface proces- ses, microscopically hispid with minute subder- mal canals and grooves. Ectosome and subectosome. Well developed series of erect spicule brushes forming a con- tinuous palisade, composed of ectosomal auxiliary subtylostyles; ectosomal brushes em- bedded on ultimate fibres, with echinating acan- thostyles and choanosomal principal styles protruding through bases of each spicule brush; subectosomal auxiliary subtylostyles form tan- gential or paratangential tracts below ectosomal skeleton; choanosomal principal styles em- bedded in peripheral fibres form diverging brushes contributing to subectosomal skeleton; mesohyl of peripheral skeleton heavier and more darkly pigmented than deeper regions of choanosome; subectosomal region relatively cavernous, occupying up to 50% of sponge diameter (less in basal stalk region). Choanosome. Skeletal architecture distinctly axially compressed, with moderately heavy, yel- low spongin fibres forming tight anastomoses near core, becoming more plumose (or merely Acanthostyles repa ae Ps joy E ger: a 58-75x4-9 | oval or eliptical meshes at EEL DIA RALLY: : core (110-275pm Chelae I 12-(15.3)-18 | 13-(15.6)-19 14.5-19.2 16 12-16 diameter), becoming wider, Chelae IT 4-(6.6)-10 6-(8.3)-10 6-10.5 9 more rectangul ar at dix 18-(60.5)-122 | 31-(101.2)-145 | 65 145 115 5 | 45-56 du periphery (230-425ym diameter); echinating acan- thostyles more heavily con- centrated on peripheral fibres and at fibre nodes; choanosomal principal megascleres uncommon or even rare in some regions of skeleton, absent entirely from the fibre core, mostly found in peripheral skeleton echinating fibres and support- ing ectosomal skeleton; mesohyl matrix relative- ly light in axial region with many loose subectosomal auxiliary megascleres scattered be- tween fibres. Megascleres (Table 38). Choanosomal principal subtylostyles straight or slightly curved at centre, with smooth slightly subtylote or rounded bases, fusiform points; principal subtylostyles differ from auxiliary subtylostyles in relatively thicker diameter with thickest part at centre of spicule, less pronounced basal constrictions, and smooth bases. Subectosomal auxiliary subtylostyles fusiform, relatively long, straight or only slightly curved, tapering to sharp points, with distinct basal constrictions and prominent subtylote swelling; bases predominantly microspined, microspines long. Ectosomal auxiliary subtylostyles similar to larger auxiliary megascleres but relatively short, thin, prominently subtylote, invariably microspined. Echinating acanthostyles large, subtylote, heavily spined bases and central regions, aspinose at points and ‘necks’ proximal to base; spines large, robust, recurved. REVISION OF MICROCIONIDAE FIG. 188. Clathria (Thalysias) procera (Ridley) (QMG300166). A, Choanosomal skeleton. B, Fibre charac- teristics (x294). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Base of subectosomal and ectosomal auxiliary subtylostyles. G-H, Palmate and modified isochelae. I, Wing-shaped and u-shaped toxas. 368 Microscleres (Table 38). Palmate isochelae in- completely divided into 2 size categories, both abundant, larger unmodified, smaller often con- tort (58-72% of spicules); lateral alae completely fused to shaft; front ala shorter and completely detached from lateral alae, Toxas wing-shaped and u-shaped, thin, vari- able in length, with pronounced central curvature, slightly reflexed or straight points. REMARKS. In live colour, surface charac- teristics, texture, gross morphology, spiculation and skeletal architecture this species is quite dis- tinctive. In particular it has sparse choanosomal principal styles found only outside (echinating) peripheral fibres; subectosomal auxiliary megascleres coring fibres; axial compression of central fibres and the diverging, wide-meshed reticulation in the peripheral skeleton; and echinating megascleres are concentrated on peripheral fibres and spongin fibre nodes. This latter feature is also found in C. (7.) cactiformis although the 2 species are not conspecific as supposed by Burton & Rao (1932), where C. (T) cüctiformis has an aspicular secondary fibre skeleton and Jacks any axial compression. The principal megascleres echinating fibres and ab- sence of principal spicules from within the fibre core indicates it belongs to Hallmann's (1912) spicata' group. Records of C. procera subsequent to. Ridley (18842) make no mention of choanosomal prin- cipal spicules echinating peripheral fibres, al- though this feature is characteristic for the species. Conversely, authors following Dendy (1922) note that there are two classes of auxiliary megascleres, both of similar Jength but different thickness, the thicker ones conng fibres and the thinner ones scattered in the mesohyl, but this distinction was nat corroborated from re-ex- amination of any material. Hallmann (1912). Dendy (1922), Burton & Rao (1932), Burton (19382) and subsequent authors included a number of other species as synonyms of C. (T) procera, but most of these synonymics arc not supported here. Clathria spiculosa var. macilenta is certainly different from C. (T) procera and is clearly a synonym of C. (T. ) reist- wordt, In contrast, Clathria spiculosa var. ramosa Hentschel (SMF1693) is conspecific with C. procera, haying closely comparable skeletal structure, spicule geometry and spicule size (Table 38), although growth form differs slightly from typical morphs (being arborescent, with a woody cylindrical stalk and numerous, MEMOIRS OF THE QUEENSLAND MUSEUM thin, evenly cylindrical branches bifurcating but not anastomosing, and bifurcate branch tips su- perficially resembling Seriotopora coral). There is some doubt about the conspecificity of some other specimens identified as C. (T.) spiculosa by Dendy (1889b, 1905, 1922) and C. (T) procera by Burton (19312, 19382) and Thomas (1973b), in particular the clathrous and lamellate morphs. These specimens all differ from typical forms in their skeletal architecture, although their spicule geometries are all fairly similar and for this reason they are retained here in synonymy for the time being. Contrary to Burton & Rao (1932) and Vosmaer (19352) C. (T.) arborescens is a distinct species from C. (T.) procera, both species differing sub- stántislly in their spicule geometry, spicule sizes und skeletal architecture. Clathria reinwardti var. palmate Ridley is conspecific with C. frondifera (= C. (T) vulpina), as suggested by Bergquist & Tizard (1967), and not with C. (T.) procera, as supposed by Burton & Rao (1932). Clathria (Thalysias) topsenti is similar in many respects tu C. (T.) procera, but shows virtually no axial com- pression of the choanosomal skeleton, spicule geometry is clearly different, and the two species are not considered to be synonyms, Clathria (Thalysias) ramusa (Kieschnick, 1896) (Figs 189) Rhaphidophlus ramosus Kieschnick, 1896:533; Kieschnick, 1900:569-570, pl.45, figs 47-50. Clathria ramosa, Hooper & Wiedenmayer, 1994- 273. Not Clathria ramosa Lindgren, 1897:482-483; Lindgren, 1898:308-309, pl-17, fig.9, pl.18, fig. 15, pl.19, fig.16; Hentschel, 1912:367. Not Thalysias ramosa; de Laubenfels, 1936a: 105, Not Collocluthria ramosa Dendy, 1922:74-76. MATERIAL. HOLOTYPE: PMJ Porif.92: Thursday J., Torres Strait, Qld, 10°35°S, 142*13'E, no other delails known (presently missing from collections; Wiedenmayer, pers.comm.) HABITAT DISTRIBUTION, Ecology unknown; known only from Torres Strait, Old, DESCRIPTION. Shape. Arborescent, bushy, with small stalk and small lobate, conical branches, between which stretches a transparent dermal membrane. Colour. Yellow-brown in preserved state. Oscules. Unknown. Texture and surface characteristics. Harsh: rugose. REVISION OF MICROCIONIDAE 100 um FIG. 189. Clathria (Thalysias) ramosa (Kieschnick) (redrawn from Kieschnick, 1900). A, Choanosornal principal subtylostyle. B, Subectosomal auxiliary subtylostyle, C, Ectosomal auxiliary subtylostyle. D, Sinuous toxa. E, Palmate isochelae. Ectosome and subectosome. Ectosomal skeleton composed of discrete brushes of small auxiliary subtylostyles. Choanosome. Choanosomal skeleton irregularly reticulate, with heavy spongin fibres divided into primary and secondary components differing sig- nificantly in diameter; fibres only lightly cored by choanosomal principal subtylostyles within axis of skeleton, occasionally absent; fibres usually more heavily cored towards periphery; echinat- ing acanthostyles abundant, character of soft parts unknown. Megascleres. Choanosomal principal subtylos- tyles prominently subtylote, straight or slightly 369 curved, with basal spination. Length 150-480j.m, width 13-24. Subectosomal auxiliary subtylostyles long, thin, prominently subtylote with microspined bases. Dimensions unknown. Ectosomal auxiliary subtylostyles identical in geometry to larger auxiliary spicules. Dimen- sions unknown. Acanthostyles cylindrical, club-shaped, evenly spined, subtylote. Length up to 150p.m, width 9-13,um. Microscleres, Palmate isochelae in 2 size classes. Length up to 13pm. Toxas thin, sinuous, raphidiform. Dimensions unknown. REMARKS. This species is barely recognisable other than belonging to Clathria and having a specialised ectosomal skeleton (i.e., C. (Thalysias)) which is both implied in Kieschnick's (1900) description and his tacit in- clusion of the species in Rhaphidophlus. Until the presently missing holotype is re-examined, the affinilies of this species remain uncertain, Clathria (Thalysias) reinwardti Vosmaer, 1880 (Figs 190-192, Table 39, Plate 8A-B) Clathria reinwardti Vosmaer, 1880:152; Vosmaer, 1935a:610, 632, 639; Bergquist & Tizard, 1967:184-186, pl.4, fig.2; Bergquist et al., 1971:102-106; Van Soest, 1989:223, fig.34; Hooper & Wiedenmayer, 1994: 273. Clathria reinwardti var, subcylindrica Ridley, 18842:446-448. Rhaphidophlus reinwardti; Kelly Borges & Bergquist, 1988:141-143, figs 3-4, pl.3f. Clathria typica. var. porrecta. Hentschel, 1912:298, 359-360. Tenacia typica var. porrecta Hallmann, 1920:77 1. Clathria spiculosa yar, macilenta Hentschel, 1912;364, Not Clathria reinwardti var. palmata Ridley, 1884a:447. MATERIAL. HOLOTYPE: RMNH(MLB)120 (not seen): Moluccas, Indonesia, no other details known. HOLOTYPE of C. spiculosa var. macilema: SMF1514 (fragments MNHNDCL22432, 2250): Straits of Dobo, Ar I, Arafura Sea, Indonesia, 6°S, 134*30' E, 40m depth, 20.iii, 1908, coll, H. Merton (dredge). LEC- TOTYPE of C reinwardti var. subcylindrica - BMNHI881.10.21.260: Thursday I., Torres Strait, Qld, 10*35' S, 142?13'E, vii. 1881, coll. HMS ‘Alert’ (dredge). PARALECTOTYPE of C. reinwardti var. subcylindrica - BMNH1882.2,23.183: Prince of Wales Channel, Torres Strait, Qld, 10?35'S, 142*13'E, vii.1881 coll. HMS ‘Alert’ (dredge). HOLOTYPE of 370 C pia war. porrecía: SMF1653 (fragment MNHNDCL2228}: Strails of Dabo, Aru L, Arafura Sea, Indonesia, 6°S, 134°50°E, 22.iii.| 908, 12m depth, coli. H. Merton (dredge), OTHER MATERIAL: NT - AMZ3099, AMZ4311 (RRIMP-0917); NTMZI76, NTMZI77, NTMZ270; NTMZ1094, NTMZ2080, NTMZ2121, NTMZ2206, NTMZ 2211, NTMZ2227, NTMZ2232, NTMZ2264, NTMZ2389, NTMZ2423, NTMZ2543, NTMZ 2545, NTMZ2554, QMG300179 (NTMZ2717), NTMZA72, NTMZ228, NTMZ435, NTMZ2174, NTMZ2197, QMG303260, NTMZ3150, NTMZ55. NTMZ77, NTMZ348, NTMZ350, NTMZ359, NTMZ361, NTMZ352, NTMZ363, NTMZ364, NIMZ441, NTMZ1364, NTMZ 1271, NTMZI1378, NTMZ2493. NTMZ2514, NTMZ3299, NTMZ3308, NTMZS70, NTMZ 586, NTMZ574, NTMZI327, NTMZ2502, NTMZ3242, NTMZ3247, NTMZ3251, NTMZ 3254, NTMZ3236, NTMZ3264, NTMZ3271, NTMZ3275, NTMZ3279, NTMZ32BS8, NTMZ 3296. NTMZ3310, NTMZ324, NTMZ333, NTMZ602, NTMZ38, NTMZ40, NTMZS0, QMG300753 (NCLQ66C-4677-Y, fragment NTMZ3906). WA - QMG30I121, QMG301 135, QMG301169, NTMZ3336 [NCIQ66C-I450-C). QLD- QMG300824, QMG304085, NTMZ4018, NTMZ4021, NTMZ4022, QMG303014, NTMZAUA3. INDONESIA - OMG303687 (NCIOCDN-1285-H). SMFI5$89, PNG- NTMZ2561, NTMZ2562, NTMZ2363, NTMZ2564, QMG300371 (NCIQ66C- 4495-A), QMG300375 (NCIQ685C-4516-Y), OMG 300883 (NCIQ66C-4547-F), QMG303104, PHILIP- PINES - QMG300344, QMG300304 (NCIQG6C- 5727-2). MICRONESIA - QMG304835. VIETNAM -QMG300045. HABITAT DISTRIBUTION. Predominantly found on coral rubble and dead coral substrata. fringing coral reefs or lagoon faunas, occasionally growing on live coral on the reef crest; mostly Found in turbid, shallow subtidal-intertidal waters between (- 1m depth, oc- casionally deeper. Speculated that association with dead coral substrates indicates some role in reel hverosion; widely distributed throughout Indo-west Pacific; Darwin Harbour, Parry Shoals, Timor Sca, Port Essington. Orontes Reef, Trepang Bay, Coboung Peninsula, Wessel Is (NT); Hibernia Reel, Carter L, Sahul Shelf. Direction I, (WA); Gulf of Carpentaria, Cockbum Is, Cape York, Shelburne Bay, Blanchard Reef, Adolphus |. (FNQ) (Fig. 190H)Y: also Cebu, Negros Orientale, Philippines (present study), Chuuk, Caroline Islands (presem study), Hon Rai L, Vietnam (present study), Solomon [s (Bergquist et al., 1971), Motupore I, PNG (Kelly Borges & Bergquist, 1988; present study), Aru Is, Sulawesi, Lesser Sumba Is, Indonesia (Vosmaer, 19352; Van Soest, 1989, present study), DESCRIPTION. Shape, Typically simple digitate, stoloniferous, cylindrical or occasional- ly laterally compressed branches (7-25mm diameter), forming meandering digits with mul- MEMOIRS OF THE QUEENSLAND MUSEUM tiple points of attachment lo substrate: no dif- ferentiation between branches and stalk, with branches attaching directly to substrate; branches frequently anastomose with adjacent branches sometimes forming complex intertwined digits; frec branches mostly simple, rarely bifurcate; several thickly encrusting, bulbous specimens also collected, presumably immature growth stages. Colour. Very light orange (Munsell SYR 8/4), orange-hrown (7.5YR 8/244), orange-red-brown (2.5YR 7/8), light brown (JOR 7/4), to grey-white (2.5Y 8/2) mgmentation alive, orange-brown (SYR 8/4) to grey-white (2.5 Y 3/2) in ethanol; ectosomal membrane varies from colourless (opaque), to grey (2.5 Y 8/2); subectosomal and choancsomal regions generally darker than periphery, usually clearly visible through ec- 1osomal membrane when alive. Oscules, Abundant, relatively large (560- 2760j.m diameter), predominant on lateral sides of branches; oscules slightly raised with promunent membrancous lip (often orange pig- mented} surrounding aperture; generally thicker specimens have larger oscules raised further above surface and more prominent subectosomal sculpturing; oscules collapse in wir, Texture and surface characteristics, Soft, come pressible, flexible, moderately easy to tear; sur- face smooth. pellucid, semi-transtucent in life, with preminent stellate subectosomal channels radiating towards oscules, particularly in thicker specimens, more even surface ornamentation in thinner specimens; ectosomal membrane collap- ses upon dessicatiun and preservation, becoming roughened and packed with ridges and cavities. Ectosame and subectosame. Thin layer of smaller ectasomal auxiliary subtylostyles form discrete brushes erect on surface, ina continuous palisade, supported by long nr short subectosomal plumose tracts from peripheral fibre skeleton; ectosomal region generally poorly collagenous but variable in thicker specimens; subectosomal region cav- ernous with lacunae (120-660).m diameter) sup- ported paucispicular plumose tracis of subectosomal auxiliary subtylostyles, no fibres, but moderate quantities of collagen between spicule tracts; plumose tracts of choanosomal principal styles also near periphery in some cases protruding through ectasomal skeleton. Choanosome, Skeletal architecture irregularly or semi-regularly reticulate, with anastomosing spongin fibres (40-120jum diameter) forming curved oval, straight triangular or rectangular meshes (50-470j.m diameter) in choanosome; REVISION OF MICROCIONIDAE E Z 8 AA f A SR CNN i a oe NC J^ i 371 FIG. 190. Clathria (Thalysias) reinwardti Vosmaer (NTMZ2174). A, Choanosomal principal styles. B, Echinat- ing acanthostyles. C, Subectosomal auxiliary styles. D, Ectosomal auxiliary styles. E, Palmate isochelae. F, Larger accolada toxas and juvenile oxhorn-like toxa. G, Section through peripheral skeleton. H, Australian distribution. I, Paralectotype of variety subcylindrica BMNH1882.2.23.183. J, NTMZ77. fibre meshes generally more irregular near core than periphery; no clear distinction between primary and secondary fibres; fibres light, always fully cored by choanosomal principal styles, with dense echinating acanthostyles on surface; abun- dant auxiliary spicules scattered between fibres; mesohyl matrix light, poorly pigmented, sur- rounding ovoid to eliptical choanocyte chambers (130-250pj.m diameter); specimens from turbid, muddy intertidal habitats incorporate moderate amounts of inorganic detritus into mesohyl but not into fibres. Megascleres (refer to Table 39 for dimensions). Choanosomal principal styles slightly curved at centre, short, thick, invariably with smooth rounded bases, hastate or occasionally stron- gylote points. 372 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 39. Comparison between spicule dimensions (in pm) between type specimens and other material of Clathria (Thalysias) reinwardti Vosmaer from different localities (N25). CULE mu A | NIRE NE E E Ed». [od ee UL BF 6 | 7 | s | T s. [9 | Subectosomal auxiliary styles straight or sometimes slightly curved near basal end, rela- tively thick, with fusiform, sharp points, some- times telescoped or mucronate, and rounded or slightly subtylote, usually faintly microspined bases. Ectosomal auxiliary styles morphologically similar to subectosomal spicules, but markedly shorter, thinner, fusiform, sharply pointed or mucronate, straight, with slight subtylote basal swellings and profusely microspined bases. Echinating acanthostyles short, stout, with rounded, bluntened points, slightly subtylote bases, unevenly spined with aspinose ‘neck’ proximal to base; spines heaviest on base and points, spines broad at base, sharp, recurved. Microscleres (refer to Table 39 for dimensions). Palmate isochelae in 2 size classes, both abun- dant, scattered throughout mesohyl and lining choanocyte chambers; long lateral alae complete- ly fused to shaft, completely detached from front ala; front ala entire; some smaller forms with contort shaft; some larger forms with median spikes on interior of shaft. Toxas basically accolada although juvenile forms resemble oxhorns; extremely thin, hair- like, long, slight central curvature, slightly reflexed or with straight points; distributed singly or in trichodragmata throughout mesohyl. Larvae. 2896 of specimens examined contained incubated parenchymella larvae in varying stages Choano- 4 i 3 111-(195.1)- | 131-(201.6)- | 146-(192.9)- | 110-(190.3)- | 116-(195.6)- | 148-(200.5)- somal 15 5 4 pe: ba $ e ej * | 280x 5- d 236x 7- | 259x7- 279x 8 264 x 9- styles (Mon) - d (11.8)-21 (11.1)-16 | (11.4)-21 (15.9)-21 113-(235.5)- | 141-(246.1)-| 141-(219.7)- | 113-(236.1)-| 146-(233.0)- | 137-(238.6)- somal tte 4 ay : ir * | 337x3- 326 x 3- 301 x 3- 337 x 4- 317x 5- 317 x 5- styles (spined) i ; (68-16 | (2611 (5.9)-11 6.7)-14 | (2-3 | (76-13 66-(102.3)- | 85-(106.5)- | 82-(104.1)- | 66-(100.7)- | 69-(102.7)- | 73-(92.8)- Boiosomal m e": X "s X* | wox2- | 162x2- | 157x2- | 166x2- | 170x4 | 139x4- y Senden (3.9)-8 (4.1)-7 (3.6)-7 (3.9)-7 (5.1-8 (3.9)-8 31-(59.1)- | 45-(64.7)- | 42-(60.1)- | 39-(57.9)- | 31-(58.2)- | 35-(60.3)- Ae 50-70x 63 | > po i pons 4 130 x 3-014)- | 78 x 3-(7,4)- | 72 x 3-(6.6)- | 79 x 3-(7.5)-|77 x 5-(7.9)- | 76 x 6-(8.7)- l y 13 12 10 13 13 12 Chelae I 13-19 10-19 12-15 — |10-(14.4)-21 | 10-(14.0)-19 | 10-(14.3)-18 | 10-(14.2)-19 | 10-(14.9)-21 | 11-(13.9)-19 Chelae II EE 4-8 2-(6.1)-9 | 2-(5.4)-9 | 2-(6.0)-9 | 2-(5.8)-9 | 4-(66)-9 | 4-(6.4)9 8-(121.2)- | 12-(119.3)- | 9-(102.5)- | 16-(125.8)- | 12-(124.1)- | 24-(115.6)- Toxas en ip ^ ea * | 237x0.5- | 186x0.5- | 173x0.5- | 236x0.5- | 228x0.5- | 186x0.5- i $21 (12-314. | (1222.5 | (12)2.55 | (13)30 | (12)2. | (11-20 | Source: l. Holotype (Vosmaer, 1880). 2. Lectotype of var. subcylindrica (BMNH1881.10.21.260). 3. Lectotype var. macilenta (SMF 1514). 4. Northern Territory specimens. 5. Fagus New Guinea specimens. 6, Indonesian specimens. 7. Philippines specimens. 8. Micronesian specimen. 9. Vietnam specimen of development; larvae oval to eliptical, 180- 825m long, 80-4001m wide; smaller larvae identical in colouration to adult mesohyl, larger larvae relatively darkly pigmented with larval styles and toxas, mostly at periphery, and mature larvae well differentiated in cellular structure with a layer of cells surrounding periphery; cilia not observed (preserved material). No obvious reproductive period for C. (T.) reinwardti because sexual reproductive products present in samples from May to January (i.e., all seasons except wet season) from Darwin and Cobourg Peninsula regions (Fig. 192); larval size not correlated with seasonality with larvae in various stages of maturity encountered throughout year; apparent absence of reproductive products during wet season probably only due to low number of samples collected during February-April, and possible that this species produces viviparous larvae all year. Associates. Virtually every specimen examined (94% of material) harboured Scyllidae polychaete worms (Typosyllis spongicola), lying between fibre meshes longitudinally within branches. The relationship between C. (T.) rein- wardti and T. spongicola is probably widespread and facultative because specimens from all localities and material collected in 1965 (Bergquist & Tizard, 1967) and 1974 (AMZA311) also contained infestations of this polychaete. REVISION OF MICROCIONIDAE Ww -l Uu FIG. 191. Clathria (Thalysias) reinwardti Vosmaer (specimen QMG303260). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Base of subectosomal and ectosomal auxiliary styles. G, Palmate and modified isochelae. H, Accolada toxas and juvenile oxhorn-like toxa. 374 SAMPLES WITH | SEASON TOTAL SAMPLES LAE — ARVAE PREDE —— 3 1 31] E EE LEN ee FIG. 192. Clathria (Thalysias) reinwardti Vosmaer. Seasonal production of incubated larvae in the NT. Variation. Shape: characteristic, consistent al- though some variability in number of branch bifurcations, thickness of branches, degree of lateral flattening; 3 morphs recognised: thin and evenly cylindrical; laterally flattened; or thickly cylindrical with knobbed and uneven surface; no correlation found between variability in skeletal architecture or spiculation and growth form; see Kelly Borges & Bergquist (1988) for further details on variability in growth form and coloura- tion. Colour: relatively consistent range from grey, orange-brown to red-brown; density of pig- mentation may be related to exposure and water clarity where deeper and more turbid water populations are generally less heavily pigmented and shallow populations are brighter coloured Skeletal structure: Ectosomal skeleton typically dense, continuous, discrete spicule brushes, oc- casionally thin, paratangential ectosomal crust. Subectosomal skeleton cavernous with long plumose, non-echinated spicule tracts (5196) or with choanosomal fibres close to surface (49%). Choanosomal skeleton typically irregularly reticulate, heavy and compact, with fully cored fibres forming rectangular meshes; but 496 of specimens with curved fibres forming oval-elip- tical meshes; 2% with cavernous skeletons throughout and very few thin fibres and spicule tracts; 4% regularly reticulate with ladder-like fibre anastomoses. Spicule skeleton lying outside fibres dense (48%) with abundant loose spicules strewn throughout mesohyl, moderate (32%), or very light (20%) with few extra-fibre spicules. Mesohyl matrix typically light (47% of specimens), moderate (25%), heavy but only lightly pigmented (18%), or heavy, dark brown pigmented (10%). Megascleres: Subectosomal auxiliary subtylostyles typically with microspined bases although most specimens had at least some smooth ones (0-4% of megascleres were entirely smooth (in 4% of specimens), 5- 10% (15%), 11-20% (34%), 21-30% (15%), 31- 40% (15%), up to 64% (17%)). Choanosomal principal styles and smaller auxiliary ectosomal subtylostyles invariably with smooth and MEMOIRS OF THE QUEENSLAND MUSEUM microspined bases, respectively. Echinating acanthostyles dense, very heavily echinating fibres (17%), moderate (43%), light (17%) or very lightly echinating (23%). Microscleres: Modified contort forms of palmate isochelae present in most specimens (although not pre- viously recorded in this species), with proportion of twisted larger isochelae ranging from 0% of spicules (46% of specimens), 1-4% (39%), 5- 10% (13%), up to 20% (2%); in smaller isochelae proportion of twisted forms 0% of spicules (13% of specimens), 1-4% (29%), 5-10% (34%), up to 20% (24%); toxas varied in abundance from very common in specimens (30% of specimens), com- mon (36%), uncommon (26%) or rare (8%). Variability in spicule dimensions: Although some variability recorded in mean spicule dimensions for samples from different localities these were not statistically significant for any spicule categories (P > 0.05); similarly spicule dimen- sions were relatively homogeneous for samples collected during different seasons, although only a small sample size was taken during the wet season. REMARKS. Vosmaer (1880) erected C. (Thalysias) reinwardti for a specimen from the Moluccas incorrectly identified as Spongia can- nabina Esper, but his original diagnosis was in- correct. Ridley (1884a) subsequently described two specimens (as var. subcylindrica) from Tor- res Strait where they were reportedly abundant. Vosmaer (1935a) redescribed the spiculation of the holotype which agreed closely with Ridley’s diagnosis, and hence emended the definition of this species. From present data and published results of Kelly-Borges & Bergquist (1988) it is shown that C. (T.) reinwardti is a dominant species of the intertidal and shallow subtidal fringing reef communities throughout the tropical Indo-west Pacific, particularly prevalent in more turbid waters. Aside from Kelly-Borges & Bergquist (1988) the species has been described as lacking any ectosomal specialisation (viz. Clathria condi- tion), whereas careful histological sectioning shows that it has a classical ectosomal skeleton of two distinct size categories of auxiliary megascleres, localised in the ectosomal and sub- ectosomal regions respectively (viz. Thalysias condition). Bergquist & Tizard (1967) suggested that toxas were not previously recorded because they are very slender and tend to be associated with larvae. However, all specimens examined in this study contained toxas, irrespective of REVISION OF MICROCIONIDAE FIG. 193. Clathria (Thalysias) ridleyi (Lindgren) (fragment of holotype BMNH1929.11.26.20). Section through peripheral skeleton. locality, seasonality or possession of larvae. Toxas are typically most abundant in the mesohyl matrix, occurring as both toxodragmata and as 375 very shallow subectosomal drainage canals radiating from large pores. Irregular and cor- rugated surface features were Observed only in a small proportion of live samples, al- though these surface features are common in dessicated material soon after collection. This species may be confused with C. (T.) erecta, differing only slightly in gross morphol- ogy and surface ornamentation, and having a similar skeletal structure, whereas comparison of spicule geometry and spicule sizes can distinguish the two species. Its spicule geometry is similar to C. (T.) fasciculata, but this has a bushy clathrous growth form and different skeletal structure (e.g., pronounced fascicular columns comprising the main skeletal tracts). In its nearly regular retuculate, rectangular skeletal structure C. (T) reinwardti resembles to some extent C. (T.) vulpina (which has an open reticulate tubular and lamellate growth form (and to which Bergquist & Tizard (1967) referred the variety C. rein- wardti var. palmata)). Clathria reinwardti can be differentiated from all these species by its characteristic acanthostyle morphology, growth form, size and geometry of toxas, and ec- tosomal-subectosomal fea- tures. The species also differs from C. (T.) procera and C. (T.) spiculosa with similar growth form in choanosomal architec- ture, spicule geometry and fibre characteristics. single spicules; they are probably also charac- Clathria (Thalysias) ridleyi teristic of the adult sponge. Bergquist et al. (1971) (Lindgren, 1897) (Fig. 193) suggested that the brick red colour and irregular lumpy surface were characteristic for this species, Rhaphidophlus sp; Ridley, 1884a:452-453. Rhaphidophlus ridleyi Lindgren, 1897:483; Dendy, whereas most specimens seen by the author in situ 1896:44; Lindgren, 1898:283, 310, 311, pl.17, fig.8, were predominantly pale orange-brown, with ^ — pl.18, fig.14, pl.19, fig.17; Hallmann, 1912:187; smooth, turgid, membraneous surface and only Van Soest, 1984b:115. Tenacia ridleyi; Lévi, 1961b:522-524. text-Fig. 14. Clathria ridleyi; Hooper & Wiedenmayer, 1994: 273 cf. Clathria ramosa; Vosmaer, 1935a:61 L, 642, 669. MATERIAL. HOLOTYPE: NHRM (fragment BMNH1929.11.26.20): near Membalong, SW. of Helitung L, Java Sea, Indonesia, 3°09°S, 107?38'E, coll, C. Aurivillius (beach debris). HABITAT DISTRIBUTION. Depth 10-14 m: on dead ar live coral substrate; Torres Strait (FNQ) (Ridley, 18843); also Java Sea (Lindgren, 1897), and Taganak I., Philippines (Lévi, 1561b). DESCRIPTION. Shape. Ramose, with thin cylindrical bifurcating and anastomosing branches, short stalk, Colour, Red alive. Oscules. Unknown. Texture and surface characteristics, Harsh. sur- face highly ornamented and hispid. bearing ridges and conules. Ectosome and subectosome. Ectosome relatively thin, with single layer of plumose spicule brushes, composed of intermingled ectosomal and subec- tosomal auxiliary subtylostyles. Choanosome_. Choanosomal skeleton irregularly reticulate, with light spongin fibres forming rec- tangular meshes and with both primary and secondary components; primary vaguely ascend- ing fibres have moltispicular core of 8-10 rows of choanosomal principal styles, whereas secondary elements less heavily cored; acanthostyles dis- persed evenly over fibres. Megascleres. Choanosomal principal styles slightly curved, with rounded, smooth non-tylote bases. Length 150-300um, width 8-1 5pm. Subectosomal auxiliary subtylostyles straight, usually with microspined bases. Length up to 300m, width up to 12.5j.m. Ectosomal auxiliary subtylostyles are identical in morphology to subectasomal spicules, Length up to 120pm, width up to 44m. Acanthostyles subtylote, with blunt or rounded, profusely microspined points, with aspinose 'necks" proximal to base. Length 68- 75m, width 5-9. 5j.m. Microscleres. Palmate isochelae unmodified, single size category. Length 9-181:m. Toxas not recorded in Ridley's or Lindgren's material, but described as raphidiform by Lévi, with slight central curvature and no apical flexion. Length 80-1 10m. REMARKS. Lindgren (1897, [898) erected this species for Ridley's (18842) unnamed specimen from Torres Strait, differentiating it from other MEMOIRS OF THE QUEENSLAND MUSEUM ramose Clathria (Thalysias) in skeletal architec- ture, fibre characteristics, and Ectyoplasia-like acanthostyles (with recurved spines on the apex of spicules). However, the species is barelv recognisable other than belonging to Clathria (Thalysias). Only a slide preparation of a skeletal section was located in the BMNH (Fig. 193), showing few distinctive characteristics, Acan- thostyles with apical spines have also heen recorded for C. (7.) mutabilis and C. (T) topsenti, and it is possible that this species is related to, or synonymous with one of these. Lévi (1961b) recorded C. (T.) ridleyi from the Philippines, noting some similarities in growth form and sur- face characteristics with C. (T) erecta, although spicule geometry and skeletal arrangement differ between the two species. Clathria (Thalysias) rubra (Lendenfeld, 1888) (Figs 194-195) Echinonema rubra Lendenfeld, 1888;221; Whitelegge, 19022:212. Clathria rubra; Hooper & Wiedenmayer, 1994; 272. Thalassodendron paucispina Lendenfeld, 1888:224- 225; Whitclegge, 1901:86-87, Rhaphidophlus paucispinus; Hallmann, 1912:176- 188, 195, 202, 203, 300, pl.25, figs 1-2, pl.26. lig. T, text-fig. 36; Guiler, 1950:8. Tenacia paucispina; Hallmann, 1920:770. Not Tenacia paucispina; Burton, 1934a:559. Thalassoendran rubens var, dura, in part; Lendenfeld. 1888:224; Whitelegge, 1901:87. Thalassodendron rubens var. lamella, in part; Lenden- feld, 1888:224, pl.7; Whitelegge, 1901:87, Not Thalassodendron rubens Lendenfeld, 1888:223. Clathria mullipora Whitelegge, 1907:496, pl.45, fig.23. ef. Microciona prolifera. Vosmaer, 19353:637. 610, 670. MATERIAL. HOLOTYPE: AMG9048 (presently missing): Port Jackson, NSW, 33°51'S, 151°16'E. PARATYPE of E. rubra: &MG9049: unknown locality (label “Clathria pumila var, rubra Lend, type ?'J. LECTOTYPE of T. paucispina: AMG9121a; Port Jackson, NSW, 33°51°S, 151*I6'E (dry, label ‘Thalysias paucispinus; type’), PARALECTOTYPES of T. paucispina: AMG9121b: same locality (dry, label 'eotype"). BMNHISB7.1.27.1, 1954.2.10 71, 1954,2.12.54 (fragments AMG3557): same locality. AM2Z961 (dry; presently missing): same locality. LEC- TOTYPE of T. rubens var. lamella: AMZAS9: Port Jackson, NSW, 33*51'S, 151"16'E (wet). PARALEC- TOTY PES of T, rubens var. lamella: AMZA61: same locality (wet, fragment from figured specimen), BMNH]887,4.27.124 (1954.2.10.70): same locality (dry). BMNH1887.1.24.28 (fragment ZMB1147): same locality (wel). HOLOTYPE of T. rubens var. dura; AMG9123: same locality (dry). PARATY PES REVISION OF MICROCIONIDAE 377 Wit N NI, g AA Ñ A Y W (lae Wig Ny rN 100 um FIG. 194. Clathria (Thalysias) rubra (Lendenfeld) (A-F, lectotype AMG9121; G, paralectotype BMNH1887.1.27.1). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, U-shaped toxa. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. 1, Lectotype. J, Paralectotype of C. multipora AMZ723. 378 of T. rubens var. dura: BMNH1887.1.24.2 (wet), BMNH1887.4.27.112 (1954.2.12.42): same locality (dry). LECTOTYPE of C. multipora: AMZ722: Off Botany Bay, NSW, 34°S, 151°11°E, 80-92m depth, coll. FIV ‘Thetis’ (dredge; label *Rhaphidophlus paucispinus, var. multiporus’), PARALECTOTY PE of C. multipora: AMZ723: same locality. OTHER MATERIAL: NSW- AMZ458, AMZ117, AMZA809, AMZ814, AM unregistered (label ‘ex. Port Jackson, NSW, coll. A Dendy’). HABITAT DISTRIBUTION. On rock reef, shell-grit or gravel substrates; 20-90m depth; Port Jackson, Botany Bay, Shoalhaven Bight (NSW); Maria I. (Tas) (Fig. 194H). DESCRIPTION. Shape. Thick, flabellate-lamel- late or branching growth forms up to 250mm long, 100mm wide, 30mm thick, usually with short basal stalk up to 60mm long, 25mm diameter; lamellate-flabellate morphs growing in 1 or more planes, even margins; ramose forms with cylindrical tapering digits or with closely anastomosing branches; intermediate flabellate- digitate growth forms with uneven palmate digitate margins. Colour. Live colouration unknown, preserved material dark-brown or grey-brown, usually with paler grey surface crust. Oscules. Moderately large, up to 4mm diameter, confined to areas on lateral or apical margins of branches. Texture and surface characteristics. Harsh, com- pressible, flexible when preserved; surface smooth, with or without small conules, often with well developed subectosomal ridges and oscular areas. Ectosome and subectosome. Surface with dis- ctinct crust or peel; ectosomal skeleton ranges from very dense, well developed continuous palisade of erect plumose brushes, to sparse, with covering of erect discrete brushes dispersed over surface, sometimes on same specimen; subec- tosomal skeleton interdispersed with ectosomal brushes, consisting of plumose tracts of larger subectosomal auxiliary subtylostyles arising from ends of peripheral choanosomal fibres, protrude through and/or lying paratangential to ectosomal layer. - Choanosome. Skeletal architecture more-or-less regularly reticulate, very heavy spongin fibres forming wide elongate-oval meshes (200-550j,.m diameter), imperfectly differentiated into primary (pauci- or multispicular) fibres (up to 160m diameter) and secondary (uni- or paucispicular) fibres (up to 110j.m diameter); fibres substantially heavier and more regularly MEMOIRS OF THE QUEENSLAND MUSEUM reticulate at core, more radial and plumo-reticu- late towards periphery; peripheral fibres ter- minate in plumose tufts of choanosomal and subectosomal megascleres, supporting ectosomal skeleton; fibres cored by choanosomal principal styles occupying only 10-40% of fibre diameter; mesohyl matrix heavy but only lightly pig- mented, containing few loose subectosomal and choanosomal megascleres dispersed between fibres; echinating acanthostyles sparse, confined mostly to larger fibres; choanocyte chambers oval, 50-80j.m diameter. Megascleres. Choanosomal principal styles rela- tively robust, usually slightly curved at centre or near basal end, with rounded or very slightly subtylote, smooth bases, and fusiform points. Length 168-(204.4)-295um, width 9-(10.9)- 14j.m. Subectosomal auxiliary subtylostyles long, slender, straight, with slightly subtylote, smooth or microspined bases, and fusiform points. Length 202-(227.1)-281 um, width 5-(6.6)-8,.m. Ectosomal auxiliary subtylostyles short, slender, straight or slightly curved near basal end, with subtylote spined bases and slightly hastate points. Length 105-(125.4)-154j.m, width 3- (4.4)-5pm. Acanthostyles subtylote, tapering cylindrical, fusiform, with evenly distributed spines or fewer spines at ‘neck’ proximal to base, spines small, straight, erect. Length 75-(84.3)-98um, width 4- (6.8)-8.m. Microscleres. Palmate isochelae of a single size class, large, unmodified, with lateral alae longer than front ala; lateral alae completely attached to shaft but detached from front ala along lateral margin. Length 17-(20.1)-24j.m. Toxas u-shaped, thick, with only slight central curvature and tapering, non-reflexed points. Length 45-(54.7)-82 um, width 1.5-(2.1)-3jum. REMARKS. This species is very similar to C. (T.) cactiformis in having a similar range of variability of growth forms and surface features, and on this basis Vosmaer (1935a) suggested they may be synonymous. However, C. (T.) rubra is substantial different from C. (T.) cactiformis in spicule geometries (particularly principal styles, acanthostyle spination, toxa morphology), spicule sizes, fibre characteristics (where all fibres are cored in this species but only the primary ascending fibres are cored in C. (T.) cactiformis), and the presence of plumose tufts of choanosomal principal styles, projecting through and echinating peripheral spongin fibres in C. (T) REVISION OF MICROCIONIDAE 379 FIG. 195. Clathria (Thalysias) rubra (Lendenfeld) (A-B, AMZ458; C-G, lectotype AMG9121). A, Choanosomal skeleton. B, Fibre characteristics (x283). C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of auxiliary subtylostyles. F, Palmate isochelae. G, U-shaped toxa. 380 cactiformis, whereas in C. (T) rubra these spicules are usually confined entirely within fibres (except on peripheral fibres where they form small bundles). A key character distinguish- ing C. (T) rubra from other species is possession of small, peculiar u-shaped (oxea-like) loxas, consistent in all specimens, very different from accolada toxas in C. (T) cactiformis. The species is a member of the ‘juniperina’ species complex having a reduced skeleton (whereby fibres shed some or all their spicules) (see discussions under C. (T.) cactiformis and C. (T.) hirsuta). Judging from the number of specimens of this species collected by the early trawling expedi- tions (Lendenfeld, Dendy, Whitelegge, Hallmann) it appears to have been a prominant member of the SE. Australian temperate sponge fauna. However, it has not been recollected for many decades, despite recent collections in both shallow and deeper waters off Sydney (EPA and NSW Water Board), and its status is uncertain. Clathria (Thalysias) ef. rubra (Lendenfeld, 1888) (Figs I96) cf. Echinonema rubra Lendenfeld, 1888:221. Tenacia paucispina; Burton, 1934a:559. MATERIAL. SPECIMEN: GREAT BARRIER REEF, QUEENSLAND - BMNH1930.8.13.108: Pen- guin Channel, off Alexander Bay, 16^ 15'S, 145°3 l'E, 20-31m depth, 24.i1,1928, coll. GBR Expedition (dredge). HABITAT DISTRIBUTION. On rock and shell gravel; 20-31m depth; known Australian distribution: Cairns region (FNQ) (Fig. 196H). DESCRIPTION. Shape. Arborescent, cylindrical branches 3-5mm diameter, branches bifurcate and anastomose producing a tangled mass., Colour. Live. colouration unknown, brown in ethanol. Oscules. Unknown, Texture and surface characteristics. Firm, com- pressible; slightly conulose surface. Ectosome and subectosome. Erect, discrete brushes of ectosomal auxiliary subtylostyles forming thin, discontinuous palisade on surface, with sparse, paratangential subectosomal skeleton composed of larger auxiliary subtylos- tyles and long principal subtylostyles protruding from peripheral fibres supporting ectosomal skeleton; principal subtylostyles also occasional- ly protruding through surface; terminal spongin MEMOIRS OF THE QUEENSLAND MUSEUM fibres branch immediately below surface; mesohyl matrix heavy in peripheral skeleton. Choanosome. Skeletal architecture irregularly reticulate; very heavy spongin fibres forming wide, oval or elongate reticulate meshes (150+ 350,.m diameter), more cavernous in periphery than at core; spongin fibres imperfectly divided into primary, mostly ascending, multispicular fibres (60-80pm diameter) and secondary, mostly transverse, paucispicular fibres (25-60j.m diameter); echinating acanthostyles relatively sparse in deeper choanosome, more-or-less con- ceniraled on exterior side of fibres and at fibre nodes; mesohyl matrix heavy, granular, with numerous miücroscleres and auxiliary spicules scattered. throughout mesohyl between fibres; choanocyte chambers oval, up to 80j.m diameter. Megascleres. Choanosomal principal subtylos- tyles slender, straight or slightly curved near base. slightly subtylote smooth or faintly microspined bases, fusiform sharply pointed or occasionally telescoped points. Length 174-(266.7)-346,um, width 5-(7.7)- Yj.m. Subectosomal auxiliary subtylostyles straight, slender, subtylote microspined or smooth bases, fusiform points. Length 219-(262,0)-358jum, width 3-44, 5)-6um. Ectosomal auxiliary subtylostyles very slender, siraight, subtylote smooth or microspined bases, fusiform points. Length 121- (152.8)- 18 E jum, width 2-(3.3)-411 m. Acanthostyles long, slender, subtylote, evenly spined except for partially aspinose area at ‘neck’ proximal to base, spines small, recurved. Length 73-(82.3)-93 pm, width 4-(4.8)-7 um. Microscleres. Palmate isochelae large, un modified. Length 15-(16.8)- 191.m. Toxas intermediate between wing-shaped and u-shaped, relatively thick, with gently rounded central curvature and reflexed points. Length 18- (43,6)-106pm, width 0.8-(1,8)-4.01.m. REMARKS. Burton's (1934a) Tenacia paucispina from the Great Barrier Reef is similar to C. (T.) rubra but there is some doubt about its conspecificity. Burton (19342) indicated that il was most similar to Clathria multipora Whitelegge, subsequently demoted to a variety (or subspecies) of C. (T.) rubra by Hallmann (1912), but com- pansen between Burton's specimen and type material (see C. (T.) rubra; Figs 194-195) shows differences in the geometry of particular spicules (toxas, acanthostyles, bases of principal styles) and in some spicule sizes. Burton (1934) did not describe his specimen, and simply noted that his 381 REVISION OF MICROCIONIDAE 50 um FIG. 196. Clathria (Thalysias) cf. rubra (Lendenfeld) (BMNH1930.8.13.108). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyle. E, Palmate isochela. F, Wing-shaped toxas. G, Section through peripheral skeleton. H, Australian distribution. 382 specimen consisted of a 'tangled, anastomosing mass of angular ... nodulose branches, each 3- 5mm diameter'. Even with a fragment of Burton's specimen it is uncertain whether it is rubra or a new taxon. Clathria (Thalysias) spinifera (Lindgren, 1897) (Figs 197-198, Table 40) Rhaphidophlus — filifer var. spinifera Lindgren, 1897:483; Lindgren, 1898:311, pl.17, fig.7, pl.19, fig.18. Rhaphidophlus spinifer, Thiele, 1903a:958, pl.28, fig. 23; Hallmann, 1912:177. Clathria spinifera; Hooper & Wiedenmayer, 1994: 274. Not Clathria spinifera Sarà, 1978:67-70, text-figs 41- 43; Desqueyroux-Faundez & Moyano, 1987:50 . cf. Microciona prolifera; V osmaer, 1935a:611. MATERIAL. LECTOTYPE: ZMUU (not seen) (frag- ments BMNH1929.11.26.6, NHNDCL 2427): specific locality unknown, Java Sea, Indonesia, no other details known. PARALECTOTYPES: ZMUU (not seen): Off Phan Thiet, Vietnam, South China Sea, 11?05'N, 108?50'E, 45m depth, coll. Capt. Svensson (dredge). OTHER MATERIAL: INDONESIA - SMF1815 (fragment MNHNDCL2378). WA- NTMZ1750.(frag- ment QMG300493). HABITAT DISTRIBUTION. Rocky reef and as- sociated sand, shell-grit and gravel beds; 45-84m depth; Port Hedland (WA) (Fig. 197H); S China Sea and Java Sea (Lindgren, 1897), Moluccas (Thiele, 1903a). DESCRIPTION. Shape. Arborescent, digitate, stalked sponge, 225mm long, 230mm wide, with slightly flattened cylindrical branches, 5-]2mm diameter (although junctions of anastomoses usually thicker), usually fused and anastomosing except at distal end; basal stalk cylindrical, woody, 55mm long, 15mm diameter. Colour. Light red-brown to grey-brown alive (Munsell 5YR 7/4) with olive-brown mottle (2.5YR 7/4), and flecks of black and copper- green on surface (possibly due to oxidation of pigments after collection); dark chocholate brown in ethanol. Oscules. Infrequently seen, scattered, not localised to any particular region, flush with sur- face, 0.5-2mm diameter. Texture and surface characteristics. Basal stalk firm, almost rigid, branches firm, compressible, highly flexible; surface minutely rugose with dis- tinct, shiny surface crust with numerous ir- regularly distributed microconules, dissected by minute ridges and canals. MEMOIRS OF THE QUEENSLAND MUSEUM Ectosome and subectosome. Moderately well developed ectosomal skeleton, with more-or-less continuous palisade of discrete spicule brushes composed of smaller ectosomal auxiliary sub- tylostyles; some detritus on ectosomal skeleton and collagen heavier and more darkly pigmented in periphery than in core; subectosomal skeleton mostly erect, plumose, occasionally tangential or paratangential to surface crust, with individual megascleres arising from subectosomal brushes invariably protruding through surface, composed of larger auxiliary megascleres arising from plumose brushes of principal styles on ultimate choanosomal fibres; ectosome and subectosomal regions together comprise only small proportion of total branch diameter; principal styles and acanthostyles echinating peripheral fibres extend close to ectosomal crust but rarely protrude beyond ectosome. Choanosome. Skeletal architecture contains both plumo-reticulate spongin fibres and plumose spicule tracts outside fibres; no differentiation of axial and extra-axial regions of choanosome al- though peripheral skeleton predominantly plumose; spongin fibres moderately heavy, 58- 1102m diameter (heavier in Indonesian specimen); fibres form oval to elongate meshes, 95-460j.m diameter, and fibre skeleton becomes increasingly plumose towards periphery; fibres indistinctly divisible into primary and secondary systems, both approximately equal diameter demarkated only by coring spicules; neither category of fibre cored by spicules for more than 60% of fibre diameter; primary ascending fibres multispicular with 2-6 principal styles per tract, many protruding through fibres at acute angles (pseudo-echinating) for less than half their length, forming plumose structures; secondary connecting fibres often transverse, rarely with more than 2 spicules per tract contained entirely within fibres; echinating acanthostyles heavy (lighter in Indonesian specimen), particularly abundant on fibre nodes (together with protrud- ing principal styles) forming characteristic stel- late-plumose echinations; towards periphery plumose brushes of principal styles protrude through fibres completely, forming multispicular tracts, and from midway along these brushes or at their points arise ascending tracts of subec- tosomal auxiliary megascleres; mesohyl matrix heavy, moderately lightly pigmented, containing few loose auxiliary megascleres. Megascleres (refer to Table 40 for dimensions). Choanosomal principal styles robust, short, thick, usually curved at centre, slightly hastate (abrupt- REVISION OF MICROCIONIDAE 383 v E ^81. | by sA 100 um FIG. 197. Clathria (Thalysias) spinifera (Lindgren) (SMF1815). A, Choanosomal principal style. B, Subec- tosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Echinating acanthostyle. E, Accolada toxa. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, NTMZ1750. TABLE 40. Comparison between spicule dimensions for type and other material of Clathria (Thalysias) spinifera (Lindgren). Measurements in jum, and cited as range (and mean) (N25). Feigmént of | holotype (BMNH 1929.11.26.6 153-(196.9)- Choanosomal styles 242x [? ^a 4)- NTMZ1750 (NW Australia) (N=1) (Thiele, 183-(206.0)- | 155-(187.2)- 242x12- | 200.x 8-(16.2)- (16.1)-18 20 172-(196.4 Subectosomal| 203-(233.8)- | 168-(258.)- | 41, K i pe styles 256 x 5-(6.8)-8 | 291 x 5-(6.9)-9 Ectosomal 97-(132,0)- | 92-(115.5) | 96- TOv styles 171 x 3-(3.8)-5 158 x 2-(4.5)-6 151 x 3-(4.9)-6 Acanthostyles| 75-(84.8)-93 x | 69-(72.3)-77x | 102-(108.6)- 6-(7.6)-9 7-(9.1)-11 | 114x 5-(8.6)-9 Chelae 14-(14.7-16 | 13-(15.2)-17 | 11-(13.9)-16 add 50-(149.0)- | ., g Toxas 26-(79.7)-124 | "gi rog. | 32-(84.2)-119 x 0.5-(0.9)-1.5 x 0.5-(0.8)-1.2 (12-1.5. ly) pointed, with rounded bases, rarely subtylote, mostly smooth (holotype and Indonesian specimen) or 50% spined (WA specimen). Subectosomal auxiliary subtylostyles long, thick, straight, tapering fusiform pointed, with slightly subtylote bases, usually minutely microspined, occasionally smooth. Ectosomal auxiliary subtylostyles short, straight or slightly curved near base, with more pronounced subtylote bases, invariably microspined, fusiform points. Acanthostyles are relatively long, slender, sharply fusiform pointed, with subtylote bases, evenly spined except for ‘neck’ proximal to base and extreme point; spines large, recurved. Microscleres (refer to Table 40 for dimensions). Palmate isochelae abundant, relatively small, un- modified, of a single size category; Indonesian specimen and holotype have typical palmate isochelae (relatively large front and lateral alae, front ala spatulate) whereas WA specimen has poorly silicified chelae, with poorly defined alae, superficially resemble small sigmas (lateral alae completely fused to shaft appearing virtually only as a ridge, front ala narrow, recurved at tip). Toxas accolada, very thin, slight central curva- ture, long slightly curved arms, little or no apical reflexion. REMARKS. The Port Hedland specimen, described above, shows some differences from both Lindgren's and Thiele's material: spongin MEMOIRS OF THE QUEENSLAND MUSEUM fibres are lighter; echinating acanthostyles are heavier on fibres; isochelae are poorly silicified, slightly sigmoid and have ill-defined alae; acan- thostyles are slightly longer; principal styles are shorter and up to 5096 have spined bases (Table 40). Conversely, skeletal structure, fibre charac- teristics, the distribution of spicules throughout fibres and spicule geometries are virtually identi- cal. Clathria (Thalysias) spinifera has unusual fibre characteristics with only bases of principal styles enclosed in primary spongin fibres (forming mul- tispicular ascending plumose tracts), and fully enclosed in secondary fibres (forming paucispicular transverse connecting tracts). Together these fibres produce an irregular renieroid-reticulation. Points of principal styles, especially in ascending spicule tracts, usually protrude through spongin fibres emphasising a plumose skeletal structure. This feature is more prominent in the Australian specimen than in Indonesian material and is reminiscent of Hallmann's (1912) ‘spicata’ group (see remarks for C. (T.) lendenfeldi), and the ‘coccinea’ group (e.g., M. coccinea Bergquist (1961a:38), M. rubens Bergquist (1961a:38), M. scotti Dendy (1924a:352), and M. parthena de Laubenfels (1930:27)). Both the ‘spicata’ and ‘coccinea’ groups of species have extra-fibre tracts com- posed of choanosomal principal megascleres. In the ‘spicata’ group those tracts occur exclusively outside fibres, and usually ascend to the ec- tosomal region, whereas in the ‘coccinea’ group tufts of principal styles congregate around fibre nodes, and they do not usually protrude beyond that region: C. (T.) spinifera shows a condition intermediate to both groups. Clathria (Thalysias) styloprothesis sp. nov. (Figs 199-200) |Echinonema typicum] Carter, nudum). Not Echinonema typicum Carter, 1881a:362. Unidentified sponge-algae associate, 'possibly undescribed’; Scott et al., 1984:291-293. 1878:163 (nomen MATERIAL. HOLOTYPE: WAM649-81(1) (frag- ment NTMZ1729): Goss Passage, off Beacon Is, Wal- labi Group, Houtman Abrolhos, WA, 28?28'S, 113?46'E, 30m depth, 7.iv.1978, coll. B.R. Wilson (trawl). PARATYPE: PIBOC-04-345 (fragment QMG300043): N. edge of Pelsart Is, Houtman Abrol- hos, WA, 28?47.2'S, 113?58.5' E, 22m depth, 10.vii.1987, coll. V.B. Krasochin, USSR RV ‘Akademik Oparin’ (SCUBA), REVISION OF MICROCIONIDAE 385 FIG. 198. Clathria (Thalysias) spinifera (Lindgren) (A-G,J, NTMZ1750; I, fragment. of holotype BMNH1929,11,26.6). A, Choanosomal skeleton. B, Fibre characteristics (x389). C, Echinating acanthostyles. D, Acanthostyle spines. E-G, Bases of choanosomal and auxiliary styles. H, Accolada toxa. LJ. Palmate and modified isochelae. 386 HABITAT DISTRIBUTION. Rock, sand and coralline substrata; 22-30m depth; Houtman Abrolhos and SW. coast (WA) (Fig. 199G). DESCRIPTION. Shape. Flabellate, irregularly vasiform, with relatively long lamellae, up to 130mm high, 125mm maximum breadth, and cylindrical basal stalk, 18mm long, 6mm diameter, lamellae moderately thin, up ta Smm maximum thickness, with rounded or uneven, bifurcate margins. Colour Live colouration unknown, yellow- brown in ethanol. Oscules, Sparse, scattered over extemal surface, up to 2mm diameter, exhalant pores minute, dis- persed over entire surface, giving eclosome slightly reticulate appearance, Texture and surface characteristics. Texture rub- bery, compressible: surface optically smooth, slightly uneven, with subdermal striations and grooves visible only near margins of lamellac. Ectusome and subectosome. Thin, disorganised ectosomal crust composed of acanthostyles erect on peripheral fibres, intermingled with paratan- gential or erect plumose brushes of auxiliary styles of 2 sizes (larger ones less common than smaller ones). together forming nearly con- tinuous palisade of erect spicule brushes on sur- face; subectosomal skeleton absent entirely: choanosomal skeleton immediately subdermal. Choanosome, Skeleton structure irregularly reticulate, with very thick ‘fibres’ formed almost exclusively by Codiaphyllum algal filaments, 70- 156m diameter, with only a superficial layer of spongin covering surface of algal filaments: coring spicules excluded entirely from within ‘fibres’, although some auxiliary subtylostyles lic on surface of “fibre” and many echinating acan- thostyles embedded within surface and erect on ‘fibre’; ‘fibre’ (algal filament) meshes usually form large nodes, and in penpheral skeleton nodes usually have tangential Jayer of subec- tosomal subtylostyles lying on surface; ‘fibres’ sinuous, extending into peripheral skeleton, with ectosomal crust perched over ‘fibre’ ends; mesohyl matrix very light, with few microscleres scattered between meshes, and smal! oval choanocyte chambers, 49-86j.m diameter. Megascleres, Choanosomal principal styles ab- sent. Subectosomal auxiliary subtylostyles relative- ly uncommon, straight, relatively thick, fusiform, with slightly constricted, smooth bases. Length 211(253.8)-292um. width 345.2)-8 jun. Ectosomal auxiliary subtylostyles most com- mon, straight or slightly curved at centre, thick, MEMOIRS OF THE QUEENSLAND MUSEUM fusiform, with smooth subtylote bases. Length 92-(128.5)-14 8p m, width 4-(6.2)-7.51.m. Acanthostyles extremely abundant, short, thick, with slightly swollen subtylote bases, even- ly spined except for aspinose points; some modified to acanthostrongyles; spines large, slightly recurved at point. Length 48-(56.4)- 63um, width 3.5-(7.2)-9j.m. Microscleres. Isochelae abundant, palmate, un- modified, of a single size category; lateral alae entirely fused to shaft, often reduced to small ‘wings’. front ala completely detached from lateral alae. Length 10.5-(13.2)-1610m. Toxas wing-shaped, short, relatively thick, slight angular central curves, slightly reflexed arms, Length 8-(43.5)-96jum, width 1.0-(1.8)- 2.5um. Associates. Probable obligatory symbiotic relationship. with red algae (Halymeniaceae), possibly Codiephyllum (identified from a super- ficial comparison with published descriptions and figures in Scott etal., 1984); algal filaments replace spongin fibres entirely, or alternatively, sponge parasitic on algae, penetrating into deepest layers of cortex of blades; association well documented (Scott et al., 1984), with similar associations known for some other microcionids (Aniho opuntioides (Lamarck) and A. frondiferu (Lam.): Topsent, 1929). ETYMOLOGY. Greek proiheco- from Topsent's (1929) lerm 'styloprathese' referring to the incorpora- tion of algal filaments into the skeleton, displacing spongin fibres, REMARKS. This species is similar to C. (T) cactiformis in acanthustyle geometry, and in fact a specimen of this species mentioned by Carter (1878) from the "west coast of Australia^ was originally named Echinonema typicum (a junior synonym of C. (T.) cacriformis). Carter's material has not been found in BMNH collections, but his description mentions the sponge-algae relation- ship as being a ‘pseudomorph’ of the free living sponge. Carter's (1878) original report nf Echinonema rypicum is not valid (nomen nudum), and his subsequent description of the species (Carer, I88la) is based on different material from his [878 notice of the species. Hence the name 'rvpiciam' is not available far this species. The external morphology of Clathria (Thalysias) styloprothesis appears to be identical to the red algae Codlophyllum flabelliforine (Sonder), redescribed and figured in detail by Scott et al. (1984) from the WA coast (lat. 28°- 3578, 5-21m depth), particularly in the thickness REVISION OF MICROCIONIDAE 387 100 um i is Woes FIG. 199, Clathria (Thalysias) styloprothesis sp.nov. (holotype WAM649-81(1)). A, Subectosomal auxiliary subtylostyle. B, Ectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton (a=algal filament), G, Australian distribution. H, Holotype. I, Paratype PIBOC04-345 (fragment QMG300043). J, Cross-section through algal lamella (pale area) showing spicules and collagen (darker areas), MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 200. Clathria (Thalysias) styloprothesis sp.nov. (paratype PIBOC04-345 (QMG300043)). A, Algal fila- ments and sponge spicules at periphery. B, Spicules embedded in filaments. C, Echinating acanthostyle. D, Acanthostyle spines. E, Palmate isochelae. F, Wing-shaped toxas. REVISION OF MICROCIONIDAE 389 100 um SETS ISSN C I VAS ARN N FE S SYN 25um A Sedit as FIG. 201. Clathria (Thalysias) tingens sp.nov. (holotype NTMZ2202). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyle. D, Accolada toxas. E, Echinating acanthostyle. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Incrusting holotype in situ growing next to Jotrochota baculifera. 390 and cortication of algal filaments. Scott et al. (1984) consider that this relationship involves an algae which has been invaded by à sponge, in which case C. (T) styloprothesis is merely a thin- ly encrusting, excavating and opportunistic sponge species. By comparison, Topsent (1929) suggested thal, in à similar case, the algal fila- ments are incorporated into the sponge (‘styloprothése’). In his material the algal fila- ments were much thinner and acanthostyles varied from forms with rounded points (siron- gyloids), to Endectyon-like (Raspailiidae) forms with relatively smooth bases and spined points (see Antho opuntioides and A. frondifera below, respectively), both of which formed a secondary renieroid reticulate skeleton, Clathria (Thalysias) styloprothesis differs from C. (T.) cactiformis in the geometry and dimen- sions of its spicules, particularly microscleres, as well as incorporating algal filaments into fibres. In this latter respect it is easily differentiated fram other microcionids, although this siatement is based on the assumption that the algal-sponge relationship is species specific, and that flabello- digitate sponges (or algae) with short squat acan- thostyles, simular in geometry to those found in C. (T) cactiformis, are conspecific. Clathria (Thalysias) tingens sp. nov. (Figs 201-203, Plate SC-D) MATERIAL. HOLOTYPE: NTMZ2202: Dudley Point, East Point Aquatic Life Reserve, Darwin, NT, 12?25.0'S, 130°49.1°E, intertidal, 23.xi.1984, coll. J.N.A. Hooper. PARATYPES - NTMZ2231: same locality, 8.113. 1985, NTMZ2530: Orontes Reef, mouth of Port Essington, Cobourg Peninsula, NT, 11*03.6' S. 132?05.4E, 11m depth, 17,ix.1985, coll. J,N.A. Hooper (SCUBA). OTHER MATERIAL: WA- QMG301154. NT- QMG300141 (fragment NTMZ2111), QLD - QMG303826, QM unreg. HABITAT DISTRIBUTION, Intertidal laterite rock, dead coral reef flats, in rock pools. encrusting on un- derside of dead faviid coral boulders, and on coral reef slopes in deeper waters, sheltered on the sides of faviid coral boulders ot Acropora thickets; 0-32m depth: Known only from Australia: Darwin Harbour, Port Essington, Cobourg Peninsula (NT); Hibernia Reef, Sahul Shelf (WA); Raine 1. (FNQ), Hook Reef (MEQ) (Fig. 201H). DESCRIPTION. Shape. Thickly encrusting, 5- 30mm thick, gelatinous lobate, following con- tours of substrate with prominent surface folding and ridge-like sculpturing. Colour. Ectosome pale pink and white mottle io pale red-orange alive (Munsell SRP 8/4-2.5R MEMOIRS OF THE QUEENSLAND MUSEUM 8/4); pigmentation below surface dark or bright red-brown (Munsell 2.5R 5/6-5/8); superficial pigmentation easily abraded from surface leaving a sponge resembling a bleeding wound; even grey-brown in ethanol. Oscules. Pores not visible in either Jive or preserved specimens. Texture and surface characteristics. Compres- sible, gelatinous, slimy in si/u, producing abun- dant clear mucous upon exposure to air; surface optically smooth, lobate, with occasional folds and minute subectosomal canals and ridges, al- though strialions not prominent; mottled or speckled external appearance superficially resembles a compound ascidian. Ectosome and subectosome. Minutely hispid, light, poorly differentiated series of ectosamal and subectosomal auxiliary spicule brushes pierce surface; choanosomal principal megascleres protrude through ectosome in thin encrustations but not in thicker sections; both ectosomal and subectosomal spicule brushes form plumose or stellate bouquets beluw peripheral skeleton but development variahle, ranging from a dense continuous palisade in thicker regions to sparse, irregular paratangential bundles of spicules in thinner sections; thinly encrusting paratype (NTMZ2530) with simple tangential ectosomal skeleton composed of bath sizes Of auxiliary spicules; subectosomal skeleton with larger auxiliary subtylostyles forming exten- sive, plumose, and discrete paralangential tracts originating close to substrate, ultimately diverg- ing and piercing ectosomal skeleton; individual subectosomal auxiliary subtylostyles also scat- tered throughout mesohyl; mesohy! matrix in peripheral skeleton heavy, granular, containing numerous pigmented spherulous cells with granular inclusions concentrated in periphery, together with occasional calcareous and siliceous foreign particles. Choonosome. Skeletal architecture hymedes- moki, with spongin fibres reduced to a basal layer, up to 30m thick, lying on coralline sub- stratum, with bases of choanosomal principal styles und acanthostyles embedded and perpen- dicular to substrate; no folding of basal spongin or fibre nodes observed; choanosomal skeleton occupies only small percentage of sponge thick- ness with major portion being extensive plumose subectosomal and ectosomal skeletons in periphery of sections; mesohy! substantially lighter in deeper choanosomal skeleton, closer to substrate, than in peripheral skeleton; choanosama| chambers $2-65y.m diameter. REVISION OF MICROCIONIDAE 391 FIG. 202. Clathria (Thalysias) tingens sp.nov. (QMG301154, Indian Ocean). A, Hymedesmoid skeleton. B, Spicules embedded in detritus and collagen. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of choanosomal and auxiliary styles. G, Accolada toxas. H, Palmate isochelae. 392 MEMOIRS OF THE QUEENSLAND MUSEUM © FIG. 203. Clathria (Thalysias) tingens sp.nov. (QMG303826, Pacific Ocean). A, Hymedesmoid skeleton. B, Erect echinating and principal spicules embedded in basal fibres and coralline substratum. C, Echinating acanthostyle. D, Acanthostyle spines, E, Base of auxiliary subtylostyle. F, Palmate isochelae. G, Accolada toxas. REVISION OF MICROCIONIDAE Megascleres Choanosomal principal megascleres long, thick subtylostyles, with dis- tinctive curvature towards basal end, prominent tylote bases abundantly microspined, tapering towards fusiform point. Length 198.5-(374.3)- 524.0jcm, width 8-(12.9)-19.1um, Subectosomal auxiliary subtylostyles long, relatively thin, mostly straight, fusiform pointed, prominent tylote bases usually microspined but also with variable proportion of smooth or polytylote bases (4-16% of spicules). Length 259.3-(377. 4-502. Sum, width | 4.5-(7.2)- 10,5y.m. Ectosomal auxiliary subtylostyles short, slender, straight. fusiform, with less prominent tylote bases, usually microspined. occasionally smooth. Length 117.9-(170.0)-208.6,um, width 2.3-(3.8)-5.6u m. Acanthostyles long, slender, straight, sub- ty]ote, fusiform pointed, evenly spined, spines small, sharp, slightly recurved. Length 79.9- (108,3)- 150,9 rm, width 4.6-(8.0}-12.6nm,. Microscleres. Palmate isochelae abundant, un- modified, with lateral and front alae approximate- ly smae length, front ala completely detached along lateral margin, lateral alae fused to shall. Length 9.7-(15.3)-20.7 jum. Toxas common or abundant. accolada, short or long, invariably thin, with very slight to moderate central curvature and straight or very slightly reflexed points. Length 37.4-(183.4)-341.7m, width 0.8-(1.9)-3,.2pm, Associations. Growing next to, or covering other encnisting sponges (Jotrochora, Placospongia, Ulosa, Mycale). compound and simple ascidians, and coralline algae. ETYMOLOGY Latin ringens, refering to its superfi- cially tinted pink live colouration. REMARKS. Clathria (Thalysias) tingens is similar to C. (7!) toxifera m spicule geometry and spicule dimensions, but differs in the geometry of its acanthostyles (evenly spined), choanosomal principal styles (markedly curved basal region, bearing spination on the base and ‘neck’ region proximal to the base), and toxa morphology (pos- session of accolada toxas). In addition, live colour, differential pigmentation berween ec- tosomal and choanosomal regions, and surface sculpturing are also diagnostic for this species, In having an easily abraded dermal pigment, the present species is reminiscent of the pink C. (Wil- sonella) tuberosa, and yellow morphs of C. (7) abietina, both from NW Australia, and the yellow C. (T.) venosa from the West Indies. Clathria 393 (Thalysias) tingens should also be compared with C, (Microciona) hymedetmivides Van Soest from Curacao in secondary colouration, hymedesmoid skeletal architecture (seen in thinly encrusting portions of C. (T.) ningens), and similarities in the morphology of some of their spicule categories. De Laubenfels (1954:135) recorded a specimen of C. (T.) cervicornis from the Marshall Is, which also had differentiated ectosomal (pale orange- brown) and choanosomal pigmentation (brilliant vermillion), although these two species differ in most other respects (e.g, growth form. skeletal architecture and spicule geometry). There are some geometric differences in spicules between Indian Ocean and Pacific Ocean specimens (Figs 202-203), but these are minor. Clathria (Thalysias) toxifera (Hentschel) (Figs 204-205, Plate SE) Hymeraphia toxifera Hentschel, 1912:382-383, p.20, fig.40, Microcimna toxifera; Burton, 1938a:31, pl.5, fig. 30; Vacelet & Vasseur, 1977:116. Clathria toxifera; Hooper & Wiedenmayer, 1994- 274, Not Strylostichon toxiferum Topsent, 1913a:621. MATERIAL. HOLOTYPE: SMF967T: Bei Minen, Aru J., Arafura Sea, Indonesia, 6°S, 134*50'E, 15m depth, 8.1v.1908, coll. H. Merton (dredge). OTHER MATERIAL: NT-NTMZ2136, NTMZ2198, NTMZ2204, NTMZ2213, NTMZ2219 (fragment QMG300506), NTMZ2222, NTMZ2233 (fragment QMG300149), NTMZ2421, NTMZ2544, NTMZ2555, QMG303296, NTMZ2217, NTMZ2074b, NTMZ2173, NTMZ2504, NTMZ1348, NTMZ3909. WA- QMG301186. THAILAND - NTM Z36681. HABITAT DISTRIBUTION. Intertidal laterite rock, dead coral reel Mats, in rock pools, on sublittoral faviid coral heads, usually exposed at ELWS tides; usually encrosting under dead faviid coral boulders, in cavities, or on metal debris (aluminium and steel) scattered over coral reefs; 0-20m depth; common intertidal encrusting sponge throughout Indo-Pacific; Darwin Harbour, Port Essington, Cobourg Peninsula, Wessel Is (NT); Hiber- nia Reef, Sahul Shelf (WA) (Fig, 204H); also Aru Is, Indonesia (Hentschel, 1912), Ko Samui, Gulf of Thailand (present study), Madras (Burton, 19383). DESCRIPTION. Shape. Thinly encrusting, l- Smm thick, often covering substantial areas of subjrala, Colour. Pale orange-brown to darker red-brown alive (Munsell SYR 7/6 - 2.5R 4/10), with whitish stellate subdermal drainage canals running over surface: colourations darkens upon exposure to air, brown to beige-grey in ethanol. 394 100 um 100 um 25um Q 6 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 204. Clathria (Thalysias) toxifera (Hentschel) (NTMZ2213). A, Choanosomal principal subtylostyles. B, Echinating acanthostyles. C, Subectosomal auxiliary subtylostyle. D, Ectosomal auxiliary subtylostyles. E, Wing-shaped toxas. F, Palmate isochelae. G, Section through hymedesmoid skeleton. H, Australian distribution. I, NTMZ2198 in situ. Oscules. Oscules minute, 150-450j.m diameter, scattered over surface, raised slightly above sur- face (on ends of conulose) or flush with surface (at nodes of drainage canals). Texture and surface characteristics. Firm, hispid, compressible in thicker regions; surface uneven, roughened, usually following contours of sub- strate, with raised projections, meandering ridges and cavities in thicker growths, or more even and with only slightly sculptured surface in thinner growths; surface with prominent subectosomal drainage canals radiating towards oscules, but these collapse, and stellate surface sculpturing disappears upon preservation; sponges produce slight clear mucous when exposed to air. Ectosome and subectosome. Ectosome slightly translucent or opaque in life, minutely hispid; spicule brushes paratangential or erect, com- posed of 2 layers: outer layer with smaller ec- tosomal subtylostyles, inner layer of larger subectosomal auxiliary subtylostyles; both layers appear intermingled but closer examination shows brushes of larger spicules originate deeper within mesohyl, whereas layer of smaller brushes occur only near periphery; points of choanosomal principal subtylostyles also protrude through sur- face brushes, up to 300pm in thin sections, REVISION OF MICROCIONIDAE whereas in thicker sections principal megascleres barely pierce surface, surrounded at their points by ectosomal spicule brushes in classical Thalysias architecture; development of ec- tosomal skeleton variable, ranging from con tinuous dense palisade of spicule brushes in thicker growths, to sparse, irregularly paratan- gential, discrete brushes in thinner sections; foreign debris sometimes incorporated into ec- tosomal skeleton with particles surrounded by spicule brushes. In subeciosamal region are also in longitudinal bands, forming dense tracts, composed of subectosomal auxiliary suhtylos- tyles usually congregated around erect principal spicules, running tangential or paratangential to ectosome; subectosomal tracts diverge near periphery to form plumose subectosomal brushes underlying ectosomal skeleton; in thin sections plumose tracts originale approximately halfway along length of pnncipal spicules whereas in thicker sections subectosomal brushes do not diverge until peripheral skeleton; subectosomil region 0.5-3mm thick containing abundant, moderately heavily pigmented. Choanosome. Choanosomal skeletal hymedes- moid in thinner sections with single megascleres embedded in basal spongin lying flat on sub- stratum, or microcionid in thicker regions with basal spongin slightly raised nodes (—'fibres ): peripheral skeletal architecture distincily plumose; choanosomal principal megascleres and echinating acanthostyles perpendicular to substrate with bases embedded in basal spangin or in erect fibre nodes where present; basal spon- gin moderately heavy, yellow-brown, lying directly on calcareous substrate, 8-20jum thick in hymedesmoid sections, up to 45pm in microcionid sections; few choanocyte chambers observed only in thicker sections, 13-55j.m diameter, usually lined by toxas and/or isochelae. Megascleres, Choanosomal principal subtylos- tyles range greatly in length, thickest near base, usually slightly curved at centre, all with prominently swollen bases, most heavily microspined or granular, rarely completely smooth (0-16% of spicules in individual specimens), all with fusiform tapering points. Length 194.0-(368.7)-685.1,.m, width 5.1- (14.1)-25.5j.m. Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, with distinctly swollen tylote bases, usually lightly microspined, less fre- quently smooth ((0- 1066 of spicules in individual specimens), Length 225.2-(354.3)-494.3j.m. width 2.0-(6.5)-12.91. m. 395 Ectosomal auxiliary subtylostyles short, straight, fusiform, less markedly tylote than larger auxiliary subtylostvles, with smooth or basal spines. Length 123.3-(171.1)-229. 61m, width 1.5-(3.6]-6.65.m. Acanthostyles thick, slightly curved towards basal end, with subrylote bases, long tapering points, mostly evenly, lightly spined except for bare ‘neck’ proximal to base, spines large. recurved, sharply pointed, Length 121.9-(154,5)- 208 Op m. width 3.0-(7.9)-14.01.m. Microscleres. Palmate isochelae usually ubun- dant (uncommon in 14% of samples), moderately large, of a single size, unmodified (although more heavily silicified in 15% of specimens), with front ala detached from and generally shorter than lateral alae, lateral alae completely fused to shaft. Length 10-422.6)-301.m. Toxas verging on oxhom, very abundant, rela- lively thick but variable in length. with wide central curvature and straight or only slightly refiexed points. Length 16-(89.8}241 jum, width 0.8-(2.9}-3 um. Associations. Growing over or in proximity to other encrusting sponges (Desmanthus, Mycale, Ulosa, Haliclona, Petrosia, and other microcionids), compound ascidians, coralline algae, Plarygyra and faviid corals, barnacles (Chthamalus). and metal debris (aluminium cans) on the reef flat (with some evidence of etching on the metal substrate). Tis probable that ihe species plays an active role in coral bioerosion. REMARKS. This species is perfectly recog- nisable from Hentschel’s (1912) original descrip- tion and matenal described here from Nonhern Territory, Western Australia and Thai waters do not differ markedly from the Indonesian popula- tion. Conversely. we do not know any details of Burton's (19382) specimen from Madras since he merely repeated verbatim the original descrip- lion, spicule measurements and figures from Hentschel (1912). Live colour, surface features, spicule size and spicule ornamentation differentiate C. (T) toxifera from other encrusting Clathria (Thalysias) but there are no unique structural differences. The older literature mostly concerns preserved material, only rarely describing species’ in situ characters, and most encrusting microcionids have few unique morphological features. Consequently. differentiation between encrusting microcionids relies mainly on details 396 MEMOIRS OF THE QUEENSLAND MUSEUM cO FIG. 205. Clathria (Thalysias) toxifera (Hentschel) (QMG301186). A, Hymedesmoid skeleton. B, Spicules embedded in basal spongin and coralline substratum (x435). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of auxiliary subtylostyles. G, Palmate isochelae. H, Wing-shaped toxas and juvenile oxhorn-like toxa. REVISION OF MICROCIONIDAE of the mineral skeleton, particularly spicule geometnes. The possession of stellate subectosomal surface sculpturing (i.e., subectosomal drainage canals radiating towards oscules) is known for C. (7.) venosus, and to a lesser extent C. (T-) virgultosa (sensu Wiedenmayer, 1977:143), both from the Caribbean. But this feature cannot be given too much taxonornic importance given that it repre- sents an ecological adaptation: viz. in thinly encrusting sponges subectosomal canals repte- sent the primary means of water circulation whereas in thicker specimens the primary aquiferous system is predominantly internal. In any case C. (T) roxifera differs from the Carih- bean species in spicule geometry and ornamenta- lion and spicule sizes. Clathria (Thalysias) toxifera should also be compared with other encrusting microcionids from the Indo-Malay and Indo-Pacific region which have hymedesmoid architecture. These in- clude numerous species from the Arafura Sea: C. (T.) aruensis (Hentschel, 1912:381), C. (T) calochela (Hentschel, 1912), C. (T) distincta (Thiele, 1903a), C. (T.) longiroxa (Hentschel, 1912), C. (Microciona) | rhopalophora (Hentschel, 1912), C. (M.) hentscheli nom. nov., C. (M.) similis (Thiele, 19032), C. (M.) terrasrvla (Hentschel, 1912) and C. (M.) thielei (Hentschel, 1912) (all of which lack isochelac). Species from other localities are: C. (T.) michaelseni (Hentschel, 1911) from Shark Bay, WA (with sigmoid anchorate-like (bidentate) isochelac); C. (M.) aceratoobtusa (Carter. L886g) from the Mergui Archipelago, Burma, and from Shark Bay, WA (Hentschel, 1911) (with smooth echinat- ing megascleres); C. (M.) affinis (Carter, 1880a), C. (M.) bulboretarta (Carter, 18803), C. (M.) fascispiculifera (Carter, 1880a) (with sigmoid palmate isochelae), and C. (M.) quadriradiata (Carter, 18802), all from the Gulf of Manaar, Sri Lanka; C. (T) dubia (Kirkpatrick, 1900a) from Christmas l., Indian Ocean (with anchorate-like isochelae); C. (C.) pellicula Whitelegge (1897) from the Ellice Is, Pacific Ocean; C. (T.) eurypa (de Laubenfels, 1954) from Palau Is, and Suva, Fiji (Tendal, 1969:40) (with 2 categories of isochelae). All those taxa differ from the present species in various details of spicule size, spicule diversity, geometry and ornamentation. Some of these species are redescribed in the present work whereas others will be redescribed 1n a forthcom- ing monograph on the Indo-Malay microcionids (Hooper et al., in prep.). From re-examination of the relevant type material of each of these species, 397 and from a detailed précis of the literature, it is clear that there are many encrusting species still undescribed, and that many of those already described are in urgent need of revision, Clathria (Thalysias) vulpina (Lamarck, 1814) (Figs 206-209, Tables 41-42, Plate BF) Smingia vulpine Lamarck, 1814:449; Lamarck, 1814-376 Rħaphidophlus vulpe; Ridley, 1884a:615. ia at all vulpinus; Topsent, 1932:110. pl.5. 1g.3. Clathria vulpine, Hooper & Lévi, 1993a:1246-1250, figs 1-12, table 6; Hooper & Wiedenmayer, 1994: 274. Halichondria frondifera Bowerbank, 1875:288-289 Amphilectus frondiferu; Vosmaer, 1880:115. Clathria Jrondifera, Ridley, 1884a:448-449, 612, pl.42, fig.i, p.53, fig.j; Ridley & Dendy, 1887:149, 160, 178, 246, 254; Topsent, 1892b:23, pl. I, fig.4; Lindgren, 1897:480, 483; Lindgren, 1898:309- 310; Dendy, 19162:128; Dendy, 1922:65; Hentschel, 1912:360-361; Raw, 1911:382, 389, 396; Burton, 1938a:27-28, pl.3, fig.21; Burton, 1959:243; Lévi, 1961c:21-22; Thomas, 1970b:206-207, text-fig.11; Thomas, 1973:33-34, pl.2. fig.6, pl.8, fig.4; Tanaka et al., 1976:801-805; Tanaka et al., 1977:767-772; Tanaka et al,, 1978:1283; Thomas, 197953:26-?7, pl.2, fig. 1; Hoshino, 1981:161; Liaaen-Jensen etal., 1982:171. Clathria frondifera vat. sera-tubulosa; Wilson, 1925:439. Ciara Jrondifera var. dichela, Hentschel, 19]12;360- 361. Tenacia frondifera var. dichela; Hallmann, 1920:771. Tenacia frondifera; Burton. & Ray, 1932:337-339; Burton, 19343:559; Lévi, 1961h:521-322, text-figs 12,13, Rhaphidophlus frondifera; Thiele, 1903a:958, test- [1g.23. Thalysias frondifera; de Laubenfels, 1954; 138-139, text-fig. 88. Clathria dichela; Vacelet et al., 1976:71, pl.3, fig.h; Vacelei & Vasseur. 1977:114. Rhaphidophlus dichela, Van Soest, 1984b:115. Clathria corallitincia Dendy, 1R89b:85, pl4, lig. S; 1916b:128; Dendy, 1922:65. Rhaphidophlus seriatus Thiele, 1899:14, pl.l, lige, pls, fig.7. Clathria reimwurdti var. palma; Ridley, 18842-447- Bergquist & Tizard, 1967; 186, ? Clathria rypica; Vacelet & Vasseur, 1971:94. Not Rhaphidophlus. filifer var. spinifera, Lindgren, 1898:311-312, pl.17, fig.7, pl.19, fig.18; Thiele, 1903a:958, Not Clathria nuda; Hentschel, 1912:298, 359, 364- 365, pI.19, fip.28. Not Clathria frondifera var. major; Hentschel, 1912:361. cf. Micraciona prolifera, Vosmaer, 19353:609, 629. 398 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 206, Clathria (Thalysias) vulpina (Lamarck) (holotype MNHNDT639). A, Choanosomal principal styles. B, Subectosomal auxiliary styles. C, Ectosomal auxiliary styles. D, Echinating acanthostyle. E, Accolada - wing-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. MATERIAL. HOLOTYPE: MNHNDT639: "? Australia", Peron & Lesueur collection. Fragments of HOLOTYPE H. frondifera: BMNH1877.5.21.1351-2: Gaspar Straits, off Belitung T., Indonesia, 3°10'S, 107*15'E. HOLOTYPE and PARATYPE of C. fron- difera var. setotubulosa: USNM21256, 21257: specific locality unknown, Philippines. HOLOTYPE of T. fron- difera var, dichela) SMF1673 (fragment MNHNDCL2230): Straits of Dobo, Aru I., Indonesia, 6°S, 134*50' E, 16m depth, 20.711.1908, coll. H. Merton (dredge), HOLOTYPE and PARATY PE ofR. seriatus: NMBI6, 17 (fragments BMNH1908.9.24.165-166, ZMB2897): Kema, off Minahassa, Celebes (Sulawesi), Indonesia, 2°50°S, 123^30'E, 30m depth, 1895, coll. P. & F. Sarasin (dredge). HOLOTYPE of C. reinwardti var, palmata: BMNH1881.10.21.264: Bird 1., Torres Strait, Old, 11°42'S, 143?05'E, coll. HMS ‘Alert’ (dredge), HOLOTYPE of C. corallitincta: BMNH1889.1.21.17 - Gulf of Manaar, Ceylon (Sri Lanka), 8*5("N, 79?40' E, coll. E. Thurston (dredge). OTHER MATERIAL (Hooper & Lévi, 19933 for ad- ditional list); INDONESIA - QMG303689 (NCIOCDN-1388-S), QMG303682 (NCIOCDN- 1252-U). PHILIPPINES - QMG300298 (NCIQ66C- 5715-C), QMG300310 (NCIQ66C-5716-C), QMG300321, QMG300332. PNG- QMG300368 (NCIQ66C-4446-X). VIETNAM - PIBOCOS5-19] (fragment QMG300047). MADAGASCAR - PIBOCBI2-200 (fragment QMG300056). NT- QMG303587, QMG303323, QMG303378, NTMZ3918, NTMZ3931, QMG300760 (NCIQ66C- 4776-1), QMG300560 (NCIQ66C-4825-L). WA - REVISION OF MICROCIONIDAE 399 FIG. 207. Clathria (Thalysias) vulpina (Lamarck) typical growth forms. A, Holotype MNHNDT639. B, Ridley's (1884) "Alert" specimen of C. frondifera BMNH1881.10.21.288. C, Holotype of C. reinwardti var. palmata BMNH1881.10.21.264. D, Holotype C. corallitincta BMNH1889.1.21.17. E, Holotype R. seriatus NMB16. F, Holotype C. frondifera var. setotubulosa USNM21257. G. Holotype C. frondifera var. dichela SMF1673. H, Hentschel's (1912) Aru I. specimen SMF1699B. 1, NTMZ1810, deeper water, NW Australia . J, QMG300047, shallow water, Gulf of Thailand . 400 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 208. Clathria (Thalysias) vulpina (Lamarck) (NTMZ2691). A, Choanosomal skeleton. B, Fibre charac- teristics (x260). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of auxiliary subtylostyles. G, Palmate isochelae. H, Accolada - wing-shaped toxas. REVISION OF MICROCIONIDAE SAMPLES WITH DESI TOTALSAMPLES| “LARVAE | Jaway | | a || o | vemm |. ad espe S2 | sepremper | 5 | 9 | | OCTOBER | — 19 | 9 [ Novemmer | 2 | 0 [ pecewpR | — 6 | — 0 jJ FIG. 209, Clathria (Thalysias) vulpina (Lamarck) In- cidence of larval production in NT populations. PIBOCO04-457 (fragment QMG300053). QLD- QMG303522, QMG300861, QMG303038, QMG304758. QMG304409, OMG30437D, QMG303898. NSW- QMG301376, QMG301385, QMG301405. HABITAT DISTRIBUTION. Mostly on rock or dead coral reefs, exposed to currents or shellered between coral heads; 0-80m depth; widely distributed throughout Indo-west Pacific; Bynoe Harbour, Darwin Harbour, Orontes reef, Melville J., Wessel Is (NT): Shark Bay, Camaryon, Barrow [,, Ex mouth Gall, Part Hedland, Monte Bello Is, Amphinome Shoals (WA); Gulf of Carpentaria, Thursday L, Bird L, Home Is, Green la Frankland Is, Low Is, Cook Reef, Shelburnc Bay, Howick Is, Lizard I. (FNQ); Hook Reef (MEQ); Byron Bay (N. NSW) (Fig. 206H); also Mozambique (Thomas, 19793), Madagascar (Vacelet et al., 1971, 1976, 1977; present study), Amirante Js (Ridley & Dendy, 1887), Seychelle [s (Ridley & Dendy, 1887; Thomas, 1979a), Aldabra Is (Lévi, 1961c), Red Sea (Topsent, 1892b; Burton, 19593), Kattiawar W. coast of India, Madras, Gulf of Manaar and Sri Lanka (Dendy, 1889b, 1916b; Lindgren, 1897; Burton, 19383; Thomas, 1970b), Mergui Archipelago and An- daman Is (Burton & Rao, 1932), Straits of Malacca, Malaysia and Gaspar Straits (Bowerhank, 1875), Aru Is, Arafura Sea, Java Sea and Sulewasi, Indonesia (Thiele, 1889; Lindgren, 1898; Hentschel, 1912; present study), Hon Rai T., Vietnam (present study }, Negros Orientale, Bohol Sea, Mindinao, and S. Philip: pines (Wilson, 1925; Lévi. 1961b; present study), Guam, Micronesia (de Laubenfels, 1954), S. Japan (Hoshino, 1981), New Caledonia (Hooper & Lévi, 19933), Madang, Papua New Guinea (present study) DESCRIPTION. Shape. Tubulo-digitate, vari- able in size ranging from small single digits (80mm high) to massive multiple digitate lobes 401 attached on a common base (450mm high); digits cavernous, insubstantial, composed of tight or loosely anastomosing sub-branches (trabeculae). Colour. Deep mauve-red (Munsell 2.5R 6/4) to pinkish red alive (2.5R 8/4) in shallow water specimens; pigmentation usually absentin deeper water specimens, beige brown alive (7.5YR 8/4). Oscules. Oscules scattered between surface con- ules, not confined to any particular region, 2- 6mm diameter, raised slightly above surface with distinct membrancous lip; ostia 0.4-2.2mm diameter flush with surface. Texture und surface characteristics, Soft, mib- bery, compressible: surface usually with small tapering digitate or spiny processes arising from free branches: surface also ornamented by minute grooves and striations, irregularly folded and cav- ernous; when intact ectosomial membrane stretched across adjacent branches, through which suhectosomal canals can be seen. Ectesome and subectosome. Conspicuous palisade of tangential, paratangential and erect spicule brushes composed of smaller auxiliary sublylostyles. with choanosomal principal megascleres protruding through surface, overlay- ing prominent plumose subectosomal spicule tracts; ectosomal skeletal density variable even within a single specimen. Choanosome. Choanosomal skeleton regular or subrenieroid reticulation of ascending, primary spongin fibres (60-li5um diameter) and transverse connecting fibres (25-39um diameter); fibre anastomoses form rectangular t square meshes lined by oval choanocyte cham- bers (22-58p m): fibres heavy, slightly heavier in axis than towards periphery; primary fibres cored by multispicular tracts of both choanosomal prin- cipal and subectosomal auxiliary megascleres, and conng spicules occupy less than 66% of primary fibre diameter; secondary fibres paucispicular, and coring spicules comprise only S0% of fibre diameter; echinating acanthostyles usually abundant, and choanosomal principal styles also echinate fibre endings in peripheral skeleton, individually or forming plumose brushes, supporting plumose tracts of subec- tosomal auxiliary spicules and ultimately ec- losomal brushes above; auxiliary megascleres also scattered between fibres. Megascleres. Choanosomal principal styles curved near base, thick, hastate or fusiform pointed, occasionally mucronate, with rounded, non-tylote, smooth bases. Length 113.3-(184.4)- 253.0um. width 3.6-(11.2)-19.4jm (holotype 156.5-(173.8)- 193.3 x 10.1-(13.1)-15.91.m). 402 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 41. Summary of results of one-way ANOY's (model 1), testing for variability in spicule lengths and widths of Clathria (Thalysias) vulpina, between groups subdivided on the basis of locality, season and bathymetric distribution, | soue | SUDO ae | ex» | 1.23 P24.05 | styles [m 1.27 P«0,40005 P2003 [AcanthosiviesL | io | søs | (850) 1.18 Chelae 1 L Number of groups — — — — Season o ene — | Choanosom E "HERE DIJEXESESES EXESES Cee [a Tram om a T ee | wm cpm (800) vas m i Emu m eie e uim Paois PARAE (600) (850) E ICTTNENUNEUME SN NNUNETSN NNCNEES (600) ce CHE CUN HY 3 CENEEN CHETNETEN HEUNE SN [| S rim [me [om ow [re I Fani ae | 029 | meus | (em moss | iem) ziz 1.7 locality mus (Northwest Shelf, Darwin, Cobour: j gb Greal Barrier Reef, Indonesia, Philippines, Guam). 2. Aem groups (wet (FMA), pre-dry (MJJ), dry ( SO), pre-wet (NDJ), for Darwin & Cobourg Peninsula material only). f depth groups (0-4m, 4-10m, 10-40m, 40m depth). Subectosomal auxiliary subtylostyles mostly straight, less often slightly curved near base, only slightly subtylote, frequently microspined, less often smooth, fusiform pointed, Length 123.6- (2214)-310.9pm, width 1.0-(4.7)-10.4um {holotype 148.2-(195.8)-231.8 x 3.6-(6.1)- 8.8m). Ectosomal auxiliary subtylostyles styles steaight, short, thin, subtylote, invariably microspined, fusiform pointed. Length. 64.2- (99,1)-161.]jym, width 1.0-(3.3)-6.94m (holotype 65.9-(87.8)-116.4 x 1.1-(3.4)-6.1 jum). Acanthostyles subtylote, fusiform sharply pointed, spined on base and midsection of shaft but aspinose on ‘neck’ proximal to base and point; spines large, recurved, sharp. Length 51.8-(73.1)- 94 7,,m, width 1.1-(6.6)- 11.6]. m (holotype 59.7- (74,3)-86.6 x 4.5-(7.8)-10.7,.m). Microscleres. Palmate isochelae abundant, scat- tered throughout the mesohyl, in 2 size classes with very few intermediate sizes; contort forms extremely rare seen in few specimens; chelae relatively poorly silicified, lateral and front alae approximately same length; front ala completely detached from lateral alae, which are fused com- pletely to shaft. Length T: 7.1-(12.4)-17.5p. m (holotype 9.1-(12.8)-17.1j.m), length 1L: 1.1- (3.8)-6.91.m (holotype 1.1-(3.7)-6.1 jum). Toxas common to uncommon, distributed sing- ly or in toxodragmata throughout mesohyl, seen particularly surrounding choanocyte chambers; toxas commonly accolada (long, thin, rhaphidiform, with little or no central curvature or apical flexion), to less commonly wing-shaped (short, thin, widely curved centrally, without flexed points). Length 7,2-(97.9)-199,5pm, width 0.5-(1.2)-3.24m (holotype 16.7-(55.8)- 94.6 x 0.6-(1.7)-2.3j.m). Larvae and reproductive periodiciry, 28% of all specimens examined (including material reported by Hooper & Lévi, 1993a) contained incubated (viviparous) parenchymella larvae in various stages of development; young larvae spherical, more mature larvae oval-elangate, 170-4201.m diameter; cilia not observed (preserved material); undifferentiated forms (170-340pm diameter) contain whispy juvenile megascleres in axis, with mesohyl matrix similar in colouration to adult; more advanced larvae (230-4204 m diameter) contain juvenile styles and toxas at centre, isochelae towards periphery, and distinct ectosomal layer of differentiated cells. In Northem Territory populations sexual REVISION OF MICROCIONIDAE 403 TABLE 42. Latitudinal gradients in spicule dimensions for populations of Clathria (Thalysias) vulpina. Measurements (in pm) are mean lengths (L) and widths (W) ISE. _SPICULE_ Choanosomal LOCATION/LATITUDE NCAL 22°s | NWS 19°s | GBR 16's cp ues | INDO6'S | PHIL 10°N (N) Q75) (250) (300) Marianas & Caroline slands, Micronesia. reproduction was distinctly seasonal, with in- cubated larvae only seen in samples collected during September-October (dry season) (Fig. 209). Associations. 24% of all specimens sampled had polychaete infestations by Typosyllis spongicola (with at least one worm in mesohyl); other epiphytic and epizootic associations not ob- served. Variation. Two morphs differentiated - 83% with spinous (rounded or sharply pointed) surface processes, and 17% without surface processes superficially resembling Hyattella intestinalis (Lamarck) (Dictyoceratida) - but growth form apparently unrelated to water depth, season or geography of samples. Presence or absence of pigmentation directly related to water depth. Variable ectosomal development, with 52% of specimens having paratangential-tangential skeletons of intermingled ectosomal and subec- tosomal auxiliary spicules and choanosomal styles protruding through ectosome; 31% of specimens had tangential skeletons in some sec- tions of ectosome and erect spicule brushes in other areas (e.g., on points of surface processes); 17% had a continuous, erect palisade of plumose spicule brushes (i.e., true Thalysias condition). 58% of specimens had thin paratangential subec- tosomal skeleton of larger auxiliary megascleres arising directly from ultimate choanosomal fibres; 28% had distinctly plumose tracts of sub- ectosomal spicules, outside of fibres, supporting ectosomal skeleton; 14% had long plumose sub- ectosomal spicule brushes and cavernous peripheral skeleton. 64% of specimens had regular choanosomal skeleton forming square Location: NCAL = New Caledonia; NWS = Northwest Shelf, WA; GBR = northern Great Barrier Rech OU: DAR = MICRO SA region,NT; CP = Cobourg Peninsula region,NT; INDO = Southeast Indonesia; PHIL = southern Philippines; MICRO 14°N (300) (175) (125) (125) styles L 156.84+9.2 167.6410.9 | 169.7+10.9 185.849.7 197.9410.1 196,94+13.5 wW 6.240.9 11.5413 | 114413 | 107409 | 120814 | 7.9408 Acanthostyles L| 51.1428 W 3,541.0 66408 | 67415 | 6.6408 5,940.4 Chelae L : 129409 | 123409 | 119811 | 126507 | 13.3207 Toxas L 79.3*17.2 79.1421.8 66,8+19.3 94,1422.2 111.7418.2 | 96.4*18.6 104.4*26.2 129.8*23.9 wW 06:02 | 12:02 | 12402 | 12402 12302 | 12:02 (renieroid) or sometimes triangular (isodictyal) meshes at core, more irregular in periphery, whereas 3696 were regularly (sub)renieroid- reticulate throughout skeleton, forming cavern- ous meshes. Fibre characteristics consistent; fibres heavy (59%) or moderately heavy (41% of specimens); primary fibres ascending (radial) (17%), transverse or longitudinal through branches (35%), or without any apparent pattern (48% of specimens); mesohyl matrix lightly pig- mented (41%), abundant, moderately heavily pigmented (17%), or heavily pigmented (28% of specimens). Choanosomal principal styles rare in 3% of specimens, largely replaced by subec- tosomal auxiliary spicules in fibres; proportion of subectosomal styles with smooth bases ranged from 0-4% of spicules sampled (7% of specimens), 5-10% (20%), 11-20% (24%), 21- 30% (21%), 31-40% (14%), up to 76% of spicules (14% of specimens); acanthostyles heavy echinating (71%) or sparsely echinating (24% of specimens). Contort isochelae abundant (7%), common (90%) or rare (3% of specimens); larger contort chelae seen in only 1046 of specimens (between 4-896 of spicules sampled); smaller contort chelae in 7% of specimens (2-10% of spicules); toxas very common (28%), common (45%), uncommon (24%) or rare (3% of specimens). Variability in spicule dimensions: Intraspecific variability was relatively high for most spicule categories, but variability was not attributed to seasonal or bathymetric distributions of samples (for all northwest Australian populations com- bined) (i.e., variation was equally consistent within and between groups). Conversely, popula- tions collected from various localities (Northwest Shelf, Darwin, Cobourg Peninsula, Great Barrier Reef, Indonesia, Philippines, Guam, New Caledonia) showed significant statistical. dif- ferences in dimensions of some spicules (choanosomal styles, acanthostyles, larger isochelae, toxas) (Table 42). Analysis of mean spicule size versus latitude (Table 41) showed certain trends in latitudinal gradients whereby spicule sizes diminish at higher latitudes. This is similar to the trend observed by Hooper & Bergquist (1992) for Cymbastela (Axinellidae), REMARKS. Clathria (Thalysias) vulpina, better known under its junior synonym of Clathria jron- difera, is a widely distributed, predominantly In- than Ocean species and a major component of the tropical macrobenthos. The cavernous, insub- stantial tubulo-digitate growth form, regularly rectangular skeletal construction and deep red colouration are characteristic for the species, al- though spicule geometry 1s unremarkable. The holotype is identical to NW. Australian material, and it is probable that onginal matenal collected by Perron & Lesseur was obtained from WA (although not specified by Lamarck, 1814). Neither Ridley (18842). Wilson (1925) nor Top- sent (1932) recorded toxas in the holotype, but these spicules are defintely present, relatively abundant albiet thin. Topsent (1932) compared the species with C. (T) clathraia (Schmidt) and suggested that although spiculation was closely comparable, the two species could be differen- tiated by the presence nf basal spination on both categories of auxiliary megascleres and the ab- sence of toxasin C. (7.) vulpina. These characters are shown here to be of little systematie impor- tance and erroneous, respectively, but these two species are otherwise differentiated by their skeletal construction and growth form. Wilson (1925) also noted that C. (7!) vulpina showed considerable intraspecific variation in growth form (and size), but his stitement over- emphasises this apparent variation. From the numerous published records of this species frem Australasian and Indo-Malay regions in par- licular (most appearing under the name of C. frondifera), itis apparent that this species has a number nf consistent and characteristic features including ils cavernous growth form, regular Isub)renieroid choanosomal skeletal structure and individual spicule geometries, whereas ec- tosomal development is much more variable. To illustrate this variability, of the previously pub- lished material. Wilson's (1925) variety of C. MEMOIRS OF THE QUEENSLAND MUSEUM setatubulosa trom the Philippines and de Laubenfels'(1954) specimen of C. frondifera from Guam have well developed Thalysias spe- cial ectosomal skeletons (i.e., distinctly smaller auxiliary subtylostyles producing an erect palisade), By comparison, Dendy's (1905) material from Sri Lanka lacks any specialised ectosomal structure, although there are two sizes of auxiliary spicules dispersed throughout the mesohyl and lying tangential to the surface, Using this species as a case-in-point, Wilson's (1925) presented pertinent arguments on the dif- ficultv in clearly differentiating Clathria und Thalysias species based on this variability in ec- tosomal development, given that the formal definition of the two taxa rests on this feature, and these arguments are supported here in demoting Thalysias to subgenus status. This species was briefly redescnbed from New Caledonian specimens (Hooper & Lévi, 19933), but no details on synonymy, population variability or living populations were given. The New Caledonian population is the most easterly recorded population and present material differs slightly from that described by Hooper & Lévi (1993a). The synonymy presented above is most- ly new, corroborated by re-examination of the relevant type material, although Clathria coral- litineta Dendy (18896) was already merged with C. frondifera by Dendy (1905), who also com- mented on the fact that ectosomal skeletal development varied substantially in the Gulf of Manaar population. Similarly, Clathria rein- wardri var. palmata Ridley was synonymised wath this species by Bergquist & Tizard (1967), and that decision is supported in the present stud y. Rhaphidophlus seriatus Thiele (1899) from Sulewasi has lighter spongin and more fully cored fibres than typical populations of C, (T.) vulpina, whereas the two species are identical in most other respects (spicule geometry, skeletal architecture, growth form), and there is no jus- tification in maintaining the two species separate- ly. Conversely, Rhaphidophlus — filifer war- spinifera Lindgren, proposed as a synonym of C. Jrondifera by Thiele (1903a), is rejected here given that they differ substantially in most char- acters and C. (T) spinifera is maintained as a good species (see description above). Burton's (1938a) assertion that Hentschel's (1912) Clathria nuda from the Arafura Sea was also a synonym of this species is not upheld here, It has an almost regular radial (extra-axial) construc- tion, nearly completely lacks echinating acan- REVISION OF MICROCIONIDAE thostyles, has only a single size of isochelae, and has substantially thicker toxas with different geometry than those seen in C. (T.) vulpina. Clathria (Thalysias) wesselensis sp. nov. (Figs 210-211, Plate 9A) MATERIAL. HOLOTYPE - NTMZ3952 (fragment QMG300702): S. of W. Rimbija L, Cape Wessel, NT, 11°01.0°S, 136?44.2'E, 15m depth, 17.xi.1990, coll. J.N.A. Hooper (SCUBA). PARATYPE QMG300361 (NCIQ66C-4761-Q): N. side Cumberland Strait, Wes- sel Is, Gove Peninsula, NT, 11?27.6' S, 136?28.T E, 13m depth, 14.xi.1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION, Fringing coral reefs, gentle slope, coral rubble, rich sponge beds, high sedimentation and turbidity; 13-15m depth; known only from Australia: Wessel Is (NT) (Fig. 210H). DESCRIPTION. Shape. Massive, club-shaped, up to 145mm high, with or without short, cylindrical basal stalk, 35mm long, 25mm diameter, expanded at apex, up to 75mm diameter, slightly flattened on upper surface. Colour. Pale red-orange alive (Munsell 5R 7/6), khaki brown in ethanol. Oscules, Large, up to 12mm diameter in life but contracted in ethanol, on apex of upper surface conules, surrounded by large, orange, transparent, membraneous lips up to 30mm high, collapsing in air. Texture and surface characteristics. Compres- sible, fibrous, basal skeleton firm, flexible. Ectosome and subectosome. Thick crust of ec- tosomal auxiliary subtylostyles erect on surfaec, in discrete plumose bundles, forming continuous palisade, incorporating some scattered sand grains, and abundant, granular, dark brown col- lagen in peripheral skeleton; subectosomal skeleton thick, plumose brushes of large subec- tosomal auxiliary subtylostyles supporting ec- tosomal brushes, arising from ends of peripheral choanosomal fibres; subectosomal region exten- sive; ectosomal and subectosomal skeletons detachable, up to 250jum thick. Choanosome. Skeletal structure irregularly reticulate, slightly more compressed at core than in periphery; spongin fibres moderately light, 65-130j.m diameter, predominantly longitudinal in sections (radial in sponge), not clearly divisible into primary or secondary elements except in axial region; primary fibres in peripheral skeleton long, close-set and parallel towards core, widely spaced and diverging near surface, becoming plumose at periphery, infrequently anastomosing along length but frequently bifurcating, especial- 405 ly closer to surface; primary fibres in axial region of skeleton close-set, short, irregularly intercon- nected by short secondary fibres, 30-50um diameter; all fibres cored by choanosomal prin- cipal subtylostyles (but barely different in mor- phology from auxiliary spicules); primary longitudinal fibres cored by multispicular tracts of principal spicules, up to 25 abreast, occupying less than 6096 fibre diameter; secondary fibres cored by uni- or paucispicular tracts of principal spicules, occupying up to 4046 of fibre diameter; all primary fibres heavily echinated by small acanthostyles but sparse on secondary fibres; mesohyl matrix heavy, granular, lightly pig- mented, with abundant microscleres and some auxiliary megascleres scattered between fibres; choanocyte chambers large, oval, 50-75jum diameter, lined by isochelae; some detritus incor- porated into mesohyl but mainly in peripheral skeleton. Megascleres. Choanosomal principal subtylos- tyles long, slender, straight, with subtylote bases, usually microspined, tapering fusiform points. Length 307-(358.3)-395j.m, width 4-(5.4)-7j.m. Subectosomal auxiliary styles very similar in geometry but slightly shorter, more slender than principal spicules; long, slender, straight, sub- tylote smooth or microspined bases, fusiform points. Length 207-(248.8)-2654m, width 2- (2.7)-4um. Ectosomal auxiliary subtylostyles short, slender, straight, subtylote smooth or occasional- ly microspined bases, fusiform points. Length 151-(172.8)-186um, width 1.5-(2.1)-3 um. Echinating acanthostyles short, slender, straight or slightly curved near distal end, sub- tylote, sharply pointed or blunt, more-or-less evenly spined; spines long, slender, prominently recurved. Length 78-(84.4)-95 jum, width 3-(3.9)- 4.5j.m. Microscleres. Palmate isochelae very abundant, moderately large, single size class, front and lateral alae approximately same length, lateral alae completely fused to shaft, front ala detached along lateral margin. Length 14-(17.2)-20jum. Toxas very abundant, very slender; longer ac- colada toxas with slight angular curvature at centre, straight arms Length I: 146-(306.8)- 415pm, width 0.5-(1.2)-2.0j.m.; shorter wing- shaped toxas moderately rounded at centre, reflexed arms. Length II: 33-(52.9)-72jum, width 0.5-(0.8)-1.5j.m. Reproductive products. Numerous, small, elipti- cal embryos present in holotype, 150-185j.m 406 100 um MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 210. Clathria (Thalysias) wesselensis sp.nov. (holotype NTMZ3952). A, Choanosomal principal subtylos- tyles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles, D, Accolada and wing- shaped toxas. E, Echinating acanthostyle. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. long, with extensive cellular differentiation but no observable larval spicules. REMARKS. This species is similar to others in the ‘juniperina’ species complex having principal spicules barely differentiated from auxiliary spicules (see remarks for C. (T.) cactiformis above). Clathria (T.) wesselensis sp. nov. differs from most of these species in having all its fibres cored. By comparison, C. (T.) juniperina has fibres cored mainly by subectosomal auxiliary spicules, largely (but not completely) replacing principals as the primary coring spicules. In other species of the ‘juniperina’ group this character is further developed whereby principal spicules are completely excluded from some or all fibres, such as in C. (T.) cervicornis in which principal spicules have been lost completely, or C. (T.) cactiformis and C. (T) placenta where only primary fibres are cored by principal spicules and REVISION OF MICROCIONIDAE 407 © r= © © FIG. 211. Clathria (Thalysias) wesselensis sp.nov. (paratype QMG300361). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-G, Bases of principal and auxiliary styles. H, Palmate isochela. I, Wing-shaped and accolada toxas. 408 secondary fibres are completely clear. In this respect the present species is most similar to C. (T.) arborescens, differing substantially in gross morphology, geometry of acanthostyles and toxas, most spicule dimensions, and having prin- cipal spicules longer than auxiliary spicules. Both these occupy the base of the ‘juniperina’ species group (i.e, principal spicules are present but reduced in primary fibres), whereas species such as C. (T.) cervicornis are most derived having lost their principal spicules completely. OTHER SPECIES OF CLATHRIA (THALYSIAS) Clathria (Thalysias) amabilis (Thiele, 1905) Stylotellopsis. amabilis Thiele, 1905:456-457, text- fig.72a-d [Punta Arenas]; Burton, 1932a:326 [Falkland Is]; Burton, 1940:115 [Argentina]; Kol- tun, 19643:66 [Antarctica]; Sarà, 1978:64-66 [Tier- ra del Fuego]. Rhaphidophlus | amabilis; Van Soest, 1984b:129 [generic synonymy]. MATERIAL. HOLOTYPE: ZMB3309. SE. Pacific rim, SW. Atlantic, Antarchea, Clathria (Thalysias) anomala (Burton, 1933) Rhaphidophlus artomalus Button, 1933:252-253, fig.3 [Natal]; Lévi, 1963:67 [Natal]. MATERIAL. Holotype: NM1410. South Africa. Clathria (Thalysias) araiosa Hooper & Lévi, 1993 Clathria (Thalysias) araiosa Hooper & Lévi. 1993a:1256-1259, figs 17-18, table 9 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300694 (fragments NTMZ3886, ORSTOMR 1370). SW Pacific. Clathria (Thalysias) arteria (de Laubenfels, 1954) Axociella arreria de Laubenfels, 1954:148-149, tèxt- fig.96 [NW. Ponape}. MATERIAL. HOLOTYPE: USNM22876, central W, Pacific. Clathria (Thalysias) arvensis (Hentschel, 1912) Hymeraphia arvensis Hentschel, 1912:381, pl.20, fig. 38 [Aru I., Arafura Sea]. Eurypon (Hymeraphia) aruensis; Lévi, 1958:27. MATERIAL. HOLOTYPE: SMF955T. Indonesia. Clathria (Thalysias) basiarenacea (Boury-Es- nault, 1973) Rhaphidophlus bastarenacea Boury-Esnaull, 1973:287, fig.49 [San Antonio Bay, Brazil] MATERIAL. HOLOTYPE: MNHNDNBE972. SW Atlantic, Clathria (Thalysias) bitoxifera (Koltun, 1970) MEMOIRS OF THE QUEENSLAND MUSEUM Axociella bitoxifera Koltun, 1970:202-204, text- fig.29, pl.8, figs 1-2 [Kurile-Kamchatka Trench, NW. Pacific]. MATERIAL. HOLOTYPE: ZIL. NW Pacific. Clathria (Thalysias) calochela (Hentschel, 1912) Hymeraphia | calochela Hentschel, 1912:383-385, pl.20, fig.41 [Aru L, Arafura Sea]. MATERIAL. HOLOTYPE: SMF1679, Indonesia. Clathria (Thalysias) coriocrassus (Bergquist & Fromont, 1988) Rhaphidophlus coriocrassus Bergquist & Fromont, 1988:111-112, pl.51, figs e-f, pl.52, figs a-b; Daw- son, 1993:39 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPORII3. New Zealand. Clathria (Thalysias) corneolia Hooper & Lévi, 1993 Clathria (Thalysias) corneolia Hooper & Lévi, 1993a:1253-1256, figs 15-16, table 8 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300691 (fragments NTMZ3877, ORSTOMRS527). SW. Pacilic. Clathria (Thalysias) eratitia (Esper, 1797) Spongia cratitia Esper, 1797:195, 196, 221, pl.53 | East Indies’ ]. Rhaphidophlus cratitius; Ehlers, 1870: 18-19, 31; Rid- ley, 1884a:450-45]; Ridley & Dendy, 1887:151- 152; Noli, 1888:51; Thiele, 1899:13; Thiele, 1903a:957-959 [Ternate, Moluccas}; Hallmann, 1912:177, 187. Thalysias cratita; de Laubenfels, 1954:137-138, text- fig.87 [NW. Ponape, Caroline Is]. Microcionu cralitia, Hartman, 1955:176-177. Desmavidon cratitia, Vosmaer, 1880:159. MATERIAL. HOLOTYPE: Unknown. Indonesia, central W. Pacific. Clathria (Thalysias) cullingworthi Burton, 1931 Clathria cullingworthi Burton, 19314:345, pl.23, figs 3-4, text-fig.4 [Natal coast]; Lévi, 1963:66., Thalysias eullingworthi; de Laubenfels, 19363: 105. MATERIAL. HOLOTYPE: NMI270 (fragments BMNH1935,7.4.68-70), South Africa. Clathria (Thalysias) delaubenfelsi (Levi, 1963) Rhaphidophlus delaubenfelsi Lévi, 1963:60-62, text- fig,70 [Cape of Good Hope]. MATERIAL. HOLOTYPE: MNHNDCLO6IS8. South Africa. Clathria (Thalysias) amirantiensis sp. nov. Colloclathria ramosa Dendy, 1922:74-76, pl.7, fig.2, pl.14, fig.4 [Amirante, Coetivy and Seychelles, In- dian Ocean]. Rhaphidaphlus ramosus; Van Soest, I984b:99, 115. REVISION OF MICROCIONIDAE Not Rhaphidophlus ramosus Kieschnick, 1896:533; Kieschnick, 1900:569-570, pl.45, figs 47-50, Not Clathria ramosa Lindgren. 1897:482-483. MATERIAL. HOLOTYPE: BMNH1921.11.7. 64.Note: C. m) ramosa (Kieschnick, 1896) has priority, NW, Indian cean. Clathria (Thalysias) distincta (Thiele, 1903) Hymeraphia distineta Thiele, 1903a:956-957, fig. 21 [Ternate, Moluccas]; Hentschel, 1912:378-379. MATERIAL. Holotype: SMF789T, Indonesia. Clathria (Thalysias) encrosta Kumar, 1925 Clathria encrusta Kumar, 1925:221, fig.4 [India]. Thalysias encrusta; de Laubenfels, 1936a:105. MATERIAL. HOLOTYPE: IMP196/1. India. Clathria (Thalysias) eurypa (de Laubenfels. 1954) Dictyociona eurypa de Laubenfels, 1954:143, fig.9! [Palau Is]. Microciona eurypa; Bergquist, 1965:164, 165, figs 2]a-b [Palau Is]; Tendal, 1969:40-41 [Suva, Fiji]. Rhaphidophlus eurypa Van Soest, 1984b:115, MATERIAL. HOLOTYPE: USNM22922. Central SW and NW Pacific, Clathria (Thalysias) fascicularis Topsent, 1889 Clathria fascicularis Topsent, 1889:35-37, fig.3 [Banc de Campeche] Pseudanchinoe 1936a: 109, Rhaphidophlus fascicularis; Van Soest, 1984b: 108, 111, 122, table 4. Clathria dentata Topsent, 1889;37-38, fig.4A [Banc dc Campeche]. MATERIAL. HOLOTYPE: MNHN missing (Van Soest, 1984b:108). NE Atlantic. Clathria (Thalysias) fasciculata Wilson, 1925 Clathria fasciculara Wilson,1925:442, pl.42, fig.6, pl.49, figs 7-8 [Sulawesi, Indonesia]; de Laubenfels, 1954:140-141, text-Fig, 89 [Truk, Caroline Is]: Tanita, 1963:124; Tanita, 1964:21; Bergquist, 1965:167-168 | Palau Is]: Tanita, 1968:47; Hoshino, 1971:24; Hoshino, 1981:161; Caberoy, 1981:20-2 1; Van Soest, 1989b:1-2, fig. 47. Thalysias fasciculata; de Laubenfels, 1936a: 105. MATERIAL. HOLOTYPE: USNM21326. Indonesia, Philip- pines, central W Pacific, Japan. Clathria (Thalysias) filifera (Ridley & Dendy, 1886) Rhaphidophlus filifer Ridley & Dendy, 1886:475; Rid- ley & Dendy, 1887:152, 247, 255, p.28, fig.2, pl.45. fig.9 [Masbate, Philippines]; Thiele, 1899:13-15; Kirkpatrick, 1900a:136; Thiele, 1903a:958; Drag- newitsch, 1905:3, 16-17; Dragnewitsch, 1906:441 [Singapore]; Whitelegge, 1907:503: Weltner, 19102:33; Ferrer Hernández, 1914:4, 42, Jascicularis: de Laubenfels, 404 Not Rhaphidophlus filifer, Topsent, 1897b:425, 447; Desqueyroux-Faundez, 1981:758, table2. Not Rhaphidophlus filifer, Lindgren, 1898:283, 311, 312, p.17, fig.7, pI.19. fig.17a-c". Not Rhaphidophlus filifer var. spinifera; Lindgren, 1897:483; Hallmann, 1912:187; Lévi, 1960a:55. Not Rhaphidophius filifer var. mutabilis; Topsent, 1897b:447-448, pl.20, fig.24, pl.21, fig.33. Noi Rhaphidophlus filifer var. cantabrica; Orueta, 1901:331-335, text-figs 1-5, pls 3-4. cf, Microciona prolifera, Vosmaer, 19353:637. MATERIAL. HOLOTYPE: BMNH1887.5.2. 104. Philip- pines, Indo-Malay region. Clathria (Thalysias) flabellata (Burton, 1936) Rhaphidophlus flabellata Burton, 1936:145, fig.4 {Oudekraal, South Africa]; Lévi, 1963:67 [note]. MATERIAL. HOLOTYPE: BMNH1935.10. 21.3. South Africa. Clathria (Thalysias) flabellifera Hooper & Lévi, 1993 Clarhria (Thalysias) flabellifera Hooper & Lévi, 19933:1250-1253, figs 13-14, table 7 [New Caledonia |. MATERIAL. HOLOTYPE; QMG300693 (fragments NTMZ3884, ORSTOMR 1416). SW Pacific. Clathria (Thalysias) hartmani (Simpson, 1966) Axocielita harimani Simpson, 1966:2393; Simpson, 19682:63-65, pl,15, text-figs 6-7, table 24 [San Juan L. Washington]. MATERIAL. HOLOTYPE: PMNH. NE Pacific. Clathria (Thalysias) hechteli sp. nov. Microciona microchela Hechtel, 1965:41-32, text- fg. 7 [Port Royal, Jamaica]; Wintermann-Kibun & Kilian, 1984:134 [Colombia]. Nol Dictyociona — microchela; de Laubenfels, 19532:528. cf. Rhaphidophlus schoenus, Van Soest, 1984b:122. MATERIAL. HOLOTYPE: PMNH 5040, PARATYPE USNM?24498. Carihbean. C. (C) oucrachela (Stephens, 1916) has priority. Clathria (Thalysias) isodictyoides (Van Soest, 1984) Rhaphidophlus isodictyoides Van Soest, 1984b:1 18- 120, pl.8, fig.6, text-lig.47, table 4 | Curacao]. MATERIAL. HOLOTYPE: ZMAPORJ781. Caribbean Clathria (Thalysias) jolicoeuri (Topsent, 1892) Rhaphidophlus jølicoeuri Topsent, 1892c:25 [Banyuls, Mediterranean]: Topsent, 1893d:446; Topsent, 1894a:19; Loisel, 1898:38; Topsent & Olivier, 1943:2 [Monaco]; Topsent, 1925:658-660, lext-fig.14 [Gulf of Naples]; Lévi, 1960b:55,65 [Dakar, N. Atlantic, Mediterranean, Naples and Monaco]; Boury-Esnault, 1971:327 [Banyuls]; 410 Pulitzer-Finali. 1983:610; Pansini & Pronzato, 1985:5 [Mediterranean]. Tenavia jolicoeuri; Lévi, 1959:133-134, text-fig.26 {Sao Tome, Gulf of Guinea]. ef. Micraciona prolifera, Vosmaer, 1935a:64 1. MATERIAL. HOLOTYPE: MOM (fragment BMNH1953.11.9.42). NW Atlantic, Mediterranean. Clathria (Thalysias) kilauea (de Laubenfels, 1951) Axocielita kilauea de Laubenfels, 1951a:262-263, text- fig.9 [Coconut I., Hawaii]. Axociella kilauea; Hechtel, 1965:43-44 | note]. MATERIAL. HOLOTYPE; USNM22779, Central Pacific. Clathria (Thalysias) lambda (Lévi, 1958) Leptoclathria lambda Lévi, 1958:38, text-fig.35 [Mar- mar, Red Sea]. MATERIAL. HOLOTYPE: MNHN missing. Red Sea. Clathria (Thalysias) lematolae sp. nov. Microciona placenta; de Laubenfels, 1954:146-147, text-fig.94. Not Spongia placenta Lamarck, 1814:374, MATERIAL. HOLOTYPE: USNM22908. Central west Pacific. C. placenta (Lamarck, 1814) has seniority. ETYMOLOGY: For the type locality, Clathria (Thalysias) linda (de Laubenfels, 1954) Axocielita linda de Laubenfels, 1954:156-158, text- fig.102 [Ailing-lap-lap, Truk]. Axociella linda; Hechtel, 1965:43-44 [wow]. MATERIAL. HOLOTYPE: USNM22860. NW ceniral Pacific. Clathria (Thalysias) lissoclada (Burton, 1934) Khaphidophlus lissocladus Burton, 1934b:32-33, 51- 52, pl.4, g.l, text-figs 4-5, 16 (Falkland Is}; Lévi, 1963:62, pL9, figs HJ, text-fig.7) |Cape of Good Hope, South Africa]. MATERIAL. HOLOTYPE: ZRS955 (fragment BMNH1933 3.17.1765). PARATYPES BMNH 1931 3.17.30, 31, 32, 38, 154, South Africa, SW Atlantic, Clathria (Thalysias) longitoxa (Hentschel, 1912) Hymeraphia longitoxa. Hentschel, 1912:381, pl.20, fig.39 [Aru 1., Arafura Sea]. Micraciona longiroxa; Burton, 1938a:30-31, pls, fig.29 [Madras, India]; Burton, 1959a:248 [Gulf of Aden]. MATERIAL. HOLOTYPE: SMF1683. NE Indian Ocean, Indonesia, Arabian Gulf Clathria (Thalysias) maunaloa (de Laubenfels, 1951) Microciona maunaloa de Laubenfels, 1951a:260-261, text-lig.6 [Coconut and Hawaii Is, Hawaii]; de Laubenfels, 1957:240 [Oahu, Hawaii}; Bergquist, 1977:65 [Hawaii]. MEMOIRS OF THE QUEENSLAND MUSEUM MATERIAL. HOLOTYPE: USNM22775. Central Pacific. Clathria (Thalysias) membranacea (Thiele, 1905) Ophlitaspongia membranacea Thiele, 1905:450-451, figs 67, 105 [Juan Fernandez Is}; Burton, 1932a:321-322 [Falkland Is]; Burton, 1940:112 [Uruguay]; Desqueyroux-Faundez & Moyano, 1987:49 [Chile, Juan Fernandez Is, Falkland Is]. ? Clathria membranacea; Hallmann, 1912:253, Axociella membranacea; de Laubenfels, 19364:113 [note]; Hechtel, 1965:43 [note]. MATERIAL. HOLOTYPE: ZMB3303 (paratypes ZMB3304, BMNH 1930.1 1.28.21). SW Atlantic, SE Pacific. Clathria (Thalysias) micropunctata (Burton & Rao, 1932) Tenacia micropunctata Burton & Rao, 1932:340-341, Lext-fig.9 [ Tuticorin, India], Thalyseurypon — micropunctata; de Laubenfels, 1936a:107 [note]. Eurypon micropunctata; de Laubenfels, 1953a;526. MATERIAL. HOLOTYPE; IMP788/1. India. Clathria (Thalysias) minuta (Van Soest, 1984) Rhaphidophlus minutus Van Soest, 1984b:115-116, texi-fig.45, lable 4 [Curaçao]; Kobluk & Van Soest, 1989:1216; Meesters. et al, 1991:195 [Curacao, Bonaire]: Muncy et al., 1991:1187 [SE. Brazil]. MATERIAL. HOLOTYPE: ZMAPOR4796. Caribbean Clathria (Thalysias) mutabilis (Topsent, 1897) Khaphidophlus filifer var. mutabilis Topsent, 1897b:447, pl.20, fig.24, pl.21, fig.33 | Ambon, Banda Sea]. Rhaphidophlus | mutabilis; Desqueyroux-Faundez, 1981:743, figs 49-54, 116, MATERIAL. HOLOTYPE; MHNGC-12/27 (fragment MNHNDT1834), Indonesia, Clathria (Thalysias) naikaiensis (Hoshino, 1981) Eurypon naikaiensis Hoshino, 1981:153-155, pi.6, fig.8, text-fig.8 [Sasajima, Japan]. MATERIAL, HOLOTYPE; MMBSSISO090-4-a. Japan. Clathria (Thalysias) nervosa (Lévi, 1963) Axociella nervosa Lévi, 1963:65-66, pl.9E, text-fig.75 [South Africa]. Rhaphidophlus nervosus, Van Soest, 1984b:115 [generic synonymy], MATERIAL. HOLOTYPE: MNHNDCL623. South Africa, Clathria (Thalysias) nuda Hentschel, 1912 Clathria nuda Hemschel, 1912:298, 359, 364-365, pl:19, fig.28 [Aru L, Arafura Sea]. Tenacia nuda; Hallmann, 1920:771. Thalysias nuda; de Laubenfels, 19363:105. cf. Microciona prolifera tropus senta; Vosmaer, 1935a:649, REVISION OF MICROCIONIDAE MATERIAL. HOLOTYPE: SMFI576 (fragment MNHNDCL2278). Indonesia. Clathria (Thalysias) ongulensis (Hoshino, 1977) Axociella ongulensis Hoshino, 1977a:45, text-fig.3, pl. L, fig.3 [fossil demosponge; W. Ongul I., Lutzow- Holm Bay, Antarctica]. MATERIAL. HOLOTYPE: MMBS, Antarcica. Clathria (Thalysias) orientalis (Brondsted, 1934) Rhaphidophlus orientalis Brondsted, 1934:20-22, text- figs 20-22 [Aru l., Arafura Sea]. MATERIAL. HOLOTYPE: Unknown. Indonesia. Clathria (Thalysias) originalis (de Laubenfels, 1930) Esperiopsis originalis de Laubenfels, 1930:27; de Laubenfels, 1932:70-72. text-fig.38 [California]. Axocielila originalis; Lee & Gilchrist, 1985:24-32 [biochemistry]; Sim & Bakus, 1986:11 [California]; Bakus & Green, 1987:71 [S California]. MATERIAL. HOLOTYPE: USNM2144], paratype BMNH1929,8.22.54, NE Pacific. Clathria (Thalysias) oxeota (Van Soest, 1984) Rhaphidophlus oxeotus Van Soest, 1984b:120- 122, text-fig.48, table 4 [Curacao]. MATERIAL. HOLOTYPE: ZMAPOR4880. Caribbean. Clathria (Thalysias) oxitoxa Lévi, 1963 Clathria | oxitoxa Lévi, 1963:54-56, text-fig.62 [Humansdorp, South Africa]. Rhaphidophlus oxiroxa; Van Soest, 1984b: 115, 122, MATERIAL. HOLOTYPE: MNHNDCL610. South Africa. Clathria (Thalysias) pachyania (Lévi, 1960) Axociella pachyaxia Lévi, 1960b:763-764, text-fig. 16, [Senegal, W. Africa]. MATERIAL. HOLOTYPE: MNHNDCL787. NW Africa. Clathria (Thalysias) robusta (Dendy, 1922) Microctona strepsiroxa var. robusta Dendy, 1922:60- 61 [Amirante f]. Tenacia robusta; Burton & Rao, 1932:339-340 |Sin- gapore]. Not Clathria robusta Koltun, 1959:186, pi.25, fig.5, text-fig.147; Van Soest & Stone, 1986:47. MATERIAL. HOLOTYPE: BMNH1921.11. 7.49. W Indian Ocean, Indo-Malay region. Clathria (Thalysias) schoenus (de Laubenfels, 1936) Clathria copiosa var. curacaoensis Amdt, 1927:148, pl.1, fig.3, iext-fig.9 [Curacao]. Aulospongus schoenus de Laubenfels, 1936a:100, pl.13, fig.3 [Dry Tortugas, Florida]. Thalysias schoenus, Simpson, 1968a:56, pls 13-14, text-fig. 5 [Florida]; Randall & Hartman, 1968:223 [West Indies]. Alcolado, 1980:4 [Cuba]. 411 Rhaphidophlus schoenus, Van Soest, 1984b:112-113, pL&, figs 1-4, text-fig.44, table 4 | Curacao, Bonaire, Puerto Rico]; Chen & Mok, 1993: 278 [probable misicentification, Taiwan]. Not Microciona microchela Hechtel, 1965:41, text hg.7 [Curacao, Bonaire, Puerto Rico, Jamaica]. MATERIAL. HOLOTYPE: USNM22404, Carribean, Clathria (Thalysias) tener Carter, 1887 Thalysias tener Carter, 1887a:70 [Mergui Ar- chipelagn]. MATERIAL. HOLOTYPE: IMFN14 (Reniera Fibrosu' ) (fragment BMNH1887.6.1.9). Andaman Sea. Imperfecily known Clathria (Thalysias) topsenti (Thiele, 1899) Rhaphidophlus filifer, in part, Topsent, 1897b:425, 447, pl.20, fig.22 [Ambon, Indonesia]; Desqueyroux-Faundez, 1981:758, table 2. Not Rhaphidophlus filifer Ridley & Dendy, 1886:475. Rhaphidophlus topsenti Thiele, 1899:15, pl.2, fig.3 |Sulawesi, Indonesia]; Whitelegge, 1907:503; Hallmann, 1912:177. cf, Microciona prolifera, Vosmaer, |935a:611, 643 MATERIAL. HOLOTYPE: NMB20 (dry) (fragments ZMB2903, BMNH1908.9.14.167). Indonesia. Clathria (Thalysias) tricurvatifera (Carter, 1876) Thalysias tricurvatifera Carter, 1876:311-312 [Cupe St. Vincent, Hebrides). MATERIAL. HOLOTYPE: unknown, (fragment BMNH1954.3,9.244). NE Atlantic, Imperfectly known Clathria (Thalysias) venosa (Alcolado, 1984) Microciona venosa Alcolado, 1984:6 [Cuba]; Koblak & Van Soest, 1989:1216. Rhaphidophlas venosus; Meesters et al., 1991:194-195 [Curagao, Bonaire]. Rhaphidophlus raraechelae Van Soest, 1984b:116- 118. pl. 8, fig.5, texi-fig.46, table 4 | Curacao]; Pulit- zer-Finali, 1986:151 [West Indies]. MATERIAL. HOLOTYPE; Cuba. Holotype of raraechelae: ZMAPORA874, Caribbean Clathria (Thalysias) virgultosa (Lamarck, 1814) Spongia virgultosa Lamarck,1814; Duchassaing & Michelotti, 1864:86, pl.23, fig.3. Thalysias virgultosa; Duchassaing & Michelot- ti, 1864:86, pl.23,fig.a [St. Thomas, Caribbean]; Tortonese,1962:3; de Laubenfels, 1936a: 104,106; Hartman, 1955-173; Lévi, 1960a:52. Microciena plana Carter, 1876:238, 472. Clathria copiosa Topsem, 1889:40-4 |, fig.6 ; Topsent, 1894b:30, 36; Hentschel, 1912:367. Thalvsias copiosa; de Laubenfels, 1936a:106. Clathria jugosa; Wilson, 1902:37. Tenacia clathrata Schmidt, 1870:56, 80 [Aniilles, Caribbean]; Carter, 1875:195; Hallmann, 1920:769; de Laubenfels, 1936a:106; Desqueyroux-Faundez & Stone, 1992: 73 [list]. 412 Clathria clathrata, Vosmaer, 1880:153; Ridley & Dendy, 1887:147; Wilson, 1902:397; Alcolado, 1976:5. Not Rhaphidophlus clathratus, Hallmann, 1912:209; Topsent, 1920b:17-18; Topsent, 1932:97, pl.5, fig.6, text-fig.3. Pandaros juniperina, Duchassaing & Michelotti, 1864:90, pl.19, fig.3; de Laubenfels, 1936a:106. Thalysias juniperina, de Laubenfels, 1936a:105-107; Hartman, 1955:171-177; Lévi, 1960a:52; Simpson, 1968a:47, 98, pls 11-12, text-fig.4, tables 18-20, 43; Randall & Hartman, 1968:218,223; Wiedenmayer, 1977:140, 142-143, 255, pl.29, figs 3-5, pl.30, figs 1-3, text-figs 146-147; Carballeira, Shalabi & Mal- donado, 1990: 235. Microciona juniperina; Hartman, 1955:171; [?] Wells et al, 1960:216-217, text-figs 13,28; Alcolado, 1980:10; Storr, 1964:42; Wintermann-Kilian & Kilian, 1984:135. Rhaphidophlus juniperinus, Van Soest, 1984b:109- 111, pl.7, fig.11, text-fig.43, table 4; Meesters et al., 1991:195. Not Spongia juniperina Lamarck, 1814:444; Lamarck, 1816:373. Not Microciona clathrata Whitelegge, 1907:493 [see C. biclathrata]. Microciona prolifera; Pearse & Williams, 1951: 135. cf. Microciona prolifera; Vosmaer, 1935a:608-61 1, 627, 628, 667, 630, 638, 644. MATERIAL. HOLOTYPE: Fragments of holotype of S. vir- gultosa: MNHNDNBE1344, 1338, BMNH1928.11.12.50, BMNH1928.11. 12.85, BMNH1954.2.20.67, USNM31049, TMPOR70, 87, IZUGCE38.766. Fragments of holotype of C. (T.) clathrata: BMNH 1870.5.3.156, 39. Caribbean, NE At- lantic. TRANSFERS List of other species described in Thalysias but now transferred to another genus. Spongia carbonaria Lamarck, 1814:375; 1816:357. Thalysias carbonaria; Duchassaing & Michelotti, 1864:83, pl.17, fig, pl.19, fig.2 [St.Thomas]; Carter, 1882a:282, pl.11, fig.11 [Antigua, West Indies]; Tortonese, 1962:3. Pellina carbonaria; Bergquist, 1965:157. Adocia carbonaria; Wiedenmayer, 1977:255, 257, tables 50-51; Van Soest et al., 1983:198. MATERIAL. HOLOTYPE: MNHNDNBE1340 (fragment: MNHNDNBEI1324, BMNH1928. 11.12.44, 56). Referred to Haplosclerida, Chalinidae, Haliclona. Thalysias coccinea Duchassaing & Michelotti, 1864:84, pl.18, fig.5 [St Thomas]. Spirastrella coccinea; Wiedenmayer, 1977:255, table 50; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: BMNH1928. 11.12.46. PARALECTOTYPE ZMAPOR2076. Referred to Hadromerida, Spirastrellidae. MEMOIRS OF THE QUEENSLAND MUSEUM Thalysias hians Duchassaing & Michelotti, 1864:86, pl.16, fig.1 [St. Thomas, Caribbean; originally designated as hyano, but corrected in erratum]. MATERIAL. HOLOTYPE: missing (Van Soest et al., 1983:203).] Unrecognisable. Thalysias ignis Duchassaing & Michelotti, 1864:83, pl.18, figs 1-2 [St.Thomas, Caribbean]. Tedania ignis, Wiedenmayer, 1977:255, table 50; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: TM POR 72 (fragment BMNH1928. 11.12.437), paralectotype ZMAPOR2373 (frag- ment MNHNDNBE1341). Referred to Tedaniidae. Thalysias massalis Carter, 1886a:50 [Port Phillip, Vic]. Reniera massalis, Dendy, 1895:236. MATERIAL. HOLOTYPE: BMNH1886.12. Referred to Haplosclerida, Chalinidae. 15.433. Thalysias proxima Duchassaing & Michelotti, 1864:84, pl.18, fig.3 [Antilles, Caribbean]. Neofibularia proxima; Wiedenmayer, 1977:255, table 50. Xestospongia proxima; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: TMPOR74 (fragments BMNH1928.11.12.45, USNM31047, MNHNDNBE1342). Referred to Haplosclerida, Petrosiidae. Thalysias repens Duchassaing & Michelotti, 1864 [St. Thomas, Virgin Is]; Carter, 1882a:282, pl.11, text-fig.10 [Puerto Cabello and Antigua, West Indies]. Xestospongia subtriangularis, Wiedenmayer, 1977:257, table 51. MATERIAL. SYNTYPES: BMNH (3 specimens, un- registered). Referred to Haplosclerida, Petrosiidae. Thalysias rugosa Duchassaing & Michelotti, 1864:84, pl.18, fig.4 [St. Thomas, Virgin Is]; Wiedenmayer, 1977:251, 253, tables 48, 49 [note]. Xestospongia subtriangularis; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: ZMAPOR2372. Referred to Haplosclerida, Petrosiidae. | Thalysias [as Thalisias] saxicava Duchassaing & Michelotti, 1864:87 [St. Thomas, Virgin Is, Caribbean]. MATERIAL. HOLOTY PE: noextant type material (Wieden- mayer, 1977; Van Soest et al., 1983)]. Unrecognisable. Dictyocylindrus sessilis Carter, 1880a:38, pl.4, fig.2 [Gulf of Manaar, Ceylon]. Aulospongus sessilis; Dendy, 1905:176 [note]. MATERIAL, HOLOTYPE: LFM destroyed. Unrecog- nisable. Thalysias subtriangularis Duchassaing, 1850; Duchassaing & Michelotti, 1864:85, pl.17, fig.1 [St. Thomas, Carib- bean]; Carter, 1879:287 [Kerguelen Is]; Carter, 1885c:196; Dendy, 1889a:58. ? Isodictya mirabilis Bowerbank; Carter, 1878:159. ? Schmidtia aulopora; Ridley, 1881:127-129 [SW Chile]. Xestospongia subtriangularis, Wiedenmayer, 1977:255, table 50; Van Soest et al., 1983:199, 204, MATERIAL. LECTOTYPE: BMNH1928.11. 12.47; lec- totype of var. lyriformis: ZMAPOR2375, paralectotype of REVISION OF MICROCIONIDAE var. lyriformis: ZMAPOR2376.Referred to Haplosclerida, Petrosiidae. Haliphysema tubulatum Bowerbank, 1873c:29 [India]. Aulospongus tubulatus; Norman, 1878:267; Dendy, 1905:176, text-fig.5 [Ceylon]; Dendy, 1922:61; Burton & Rao, 1932:347 [Tuticorin, India]; de Laubenfels, 1936a:101 [note]. Axinella tubulata; Dendy, 1889b:89, pl.5, fig.2. MATERIAL. HOLOTYPE: BMNH not found (poorly preserved fragments BMNH1887.5.21. 1331, 1332). Uncer- tain placement; possible Raspailiidae. Thalysias varians Duchassaing & Michelotti, 1864:86, pl.13, fig.6 [St. Thomas, Caribbean]. Anthosigmella varians, Topsent, 1918:557; de Laubenfels, 1957:242-243; Pang, 1973:47-50, text- fig.14 [Jamaica; plus synonymy]; Wiedenmayer, 1977:255, table 50; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: TM POR 71 (fragments USNM31048, MNHNDNBE1343); paralectotype of var. varians: ZMAPOR2377 (fragment BMNH1928.11.12.49); lectotype of var. encrustans: BMNH1928.11.12.48. Referred to Hadromerida, Spirastrellidae. Echinonema vasiplicata Carter, 1882b:114 [Swan River, WA]; Dendy, 1889a:44. Echinodictyum mesenterinum; Ridley, 1884b:185. Echinodictyum bilamellatum; Dendy & Frederick, 1924:504. MATERIAL. HOLOTYPE: BMNH1887.5.21.1853. referred to Raspailiidae, synonym of Echinodictyum mesenterinum (Lamarck), SYNTYPES: MNHNDT661, 3425, 3427. Referred to Hadromerida, Spirastrellidae. Antho Gray, 1867 Refer to subgenera for synonymy. TYPE SPECIES. Myxilla involvens Schmidt, 1864: 37 (by monotypy). DEFINITION. Two distinct skeletal components: (1) primary (basal or axial) renieroid (rectan- gular) or isodictyal (triangular) choanosomal skeleton composed of acanthostyles and/or acan- thostrongyles; (2) secondary (extra-axial, subec- tosomal) skeleton composed of smooth choanosomal styles forming dendritic, plumose, subisodictyal or plumoreticulate tracts, or simply echinating main spicule tracts; secondary skeleton usually arising from nodes of renieroid skeleton, or ascending upwards from basal spon- gin fibres, with or without axial compression; spongin fibres relatively poorly developed; addi- tional category of echinating acanthostyles present or absent; ectosomal skeleton tangential, paratangential or plumose tracts of 1 or 2 categories of auxiliary styles; microscleres diverse forms of isochelae and toxas. 413 REMARKS. Under Van Soest & Stone's (1986) system all microcionids having a renieroid (and/or isodictyal) basal (or axial) skeleton com- posed of acanthose megascleres are grouped in Antho. This system is supported here with sub- genera recognised on structure and composition of the renieroid skeleton (1) A. (Antho) (with predominantly (acantho)styles forming the renieroid skeleton, less often acanthostrongyles, without echinating acanthostyles); (2) A. (Plocamia) (with predominantly (acantho)stron- gyles forming the renieroid skeleton, less often acanthostyles, and a special category of echinat- ing acanthostyles overlap the main skeleton); (3) A. (Isopenectya) (with an axially compressed and extra-axially renieroid reticulate skeleton com- posed of 2 forms of choanosomal spicules inside spongin fibres, overlaid by a second extra-axial plumose skeleton. /sopenectya (s.s.) could also be included in Echinoclathria, given the close resemblance in growth form and renieroid skele- tal structure with E. leporina, but in A. (Isopenec- tya) the renieroid skeleton is of sparsely spined principal styles (differentiated from the larger smooth styles of the extra-axial skeleton), with differentiated axial (compressed) and an extra- axial (renieroid) regions, overlaid by a second extra-axial (plumose) skeleton composed of larger, smooth principal styles. In Echinoclathria, megascleres of the renieroid skeleton are ex- clusively smooth, and the larger, smooth prin- cipal styles which form a radial skeleton are only found on the surface, embedded in peripheral fibres. Antho and Echinoclathria, differ from other microcionids in having a renieroid skeleton and it is possible that Echinoclathria is a highly derived form of Antho (loss of spinated principal spicules, loss of extra-fibre skeleton, loss of spined acanthostyles (geometrically different from principal spicules)). Antho (Antho) Gray, 1867 Antho Gray, 1867: 524; Lévi, 1960a: 57. Anomoclathria Topsent, 1929: 26 (not Topsent, 1932: 103). Anthoarcuata Bakus, 1966: 431. Dictyoclathria Topsent, 1920b: 18. Dyctioclathria Ferrer-Hernandez, 1921: 172 [lapsus]. Isociona Hallmann, 1920: 768. Jia de Laubenfels, 1930: 28. Plocamilla; in part, Burton, 1935a: 402; Pulitzer- Finali, 1973: 40 (not Topsent, 19282: 63). Quizciona de Laubenfels, 1936a: 111. TYPE SPECIES. Myxilla involvens Schmidt, 1864: 37 (by monotypy). 414 MEMOIRS OF THE QUEENSLAND MUSEUM 500 um FIG. 212. Antho (Antho) opuntioides (Lamarck) (lectotype MNHNDT654). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyles. C, Acanthostyles of renieroid skeleton. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Known Australian distribution. H, Lectotype. I, Paralec- totype MNHNDT3418. REVISION OF MICROCIONIDAE DEFINITION. Primary basal renieroid (and/or isodictyal) choanosomal skeleton composed of acanthostyles and/or acanthostrongyles; secon- dary extra-axial (subectosomal) skeleton plumose, plumoreticulate, or simply composed of choanosomal styles echinaüng (project from) basal renieroid skeleton; spongin fibres poorly developed; special category of echinating acan- thostyles absent; ectosomal skeleton with tangen- tial, paratangential, or plumose tracts of I size of auxiltary styles or subtylostyles; microscleres in- clude diverse forms of isochelae and toxas. REMARKS. Twenty one species have been in- cluded 1n, or referred to, Antho (Antha), although only 1! are widely accepted, 2 of which are known from Australia. Antho (Antho) opuntioides (Lamarck, 1815) (Figs 212-213) Unidentified sponge; Turgot, 1758: pl.24, fig e. Alcyonium opuntioides Lamarck, 1815: 164, Anomoclathria opuntioides; Topsent, 1929: 21-26, lext- figs 1-9. a opuntioides; Hooper & Wiedenmayer, 1994: 255. Not Anomoclathria opuntioides var. frondifera; Top- sent, 1929: 26-29, text-figs 10-14; Topsent, 1932: 103, pl.1, figs 6-7. MATERIAL. LECTOTYPE: MNHNDT654: Precise locality unknown, SW. Australia, Peron & Lesucur collection. PARALECTOTYPES - MNHNDT3416, 3418: same data, HABITAT DISTRIBUTION. Unknown, known only from type locality (Fig. 212G). DESCRIPTION. Shape. Lobate, digitate growth form, up to 255mm high, 375mm wide, 230mm thick, bifurcating cylindrical or slightly flattened branches, up to 120mm long, 18mm maximum diameter, occasionally anastomosing, slightly bulbous branch nodes, tapering or rounded branch tips, Colour. Live colouration unknown, grey in dry State. Oscules. Small pores up to 2mm diameter, pos- sibly oscules, scattered over sides of branches. Texture and surface characteristics. Harsh, brittle in dry state, even, unornàmented surface. Ectosome and subectosome. No ectosomal membrane intact (dry material), although rem- nants of sparse, tangential and paratangential skeleton composed of subectosomal auxiliary subtylostyles scattered near periphery; points of rchoanosomal styles and acanthostyles in peripheral skeleton protrude through surface; en- tire peri skeleton dense, virtually undif- ferentiated from deeper choanosomal skeleton (although spicule tracts with more sparse spongin component in periphery than at core), clearly dominated bv close-set renieroid reticulation of acanthostyles. Choanasome. Skeleton with two distinct com- ponents: plumose extra-axial skeleton composed of multispicular or paucispicular continuous tracts of Jarge choanosomal principal styles ex- tending from centre of skeleton to ectosome; renieroid skeleton regular, tight meshed, rectan- gular and triangular meshes, 80-150,.m diameter, even mesh size throughout skeleton; spongin fibres heavier, slightly more compressed at centre of skeleton than in periphery, with oval meshes 60-135j.m diameter; echinating spicules absent; mesohyl matrix light, with scattered microscleres; choanocyte chambers not seen (dry specimens). Megascleres. Smooth choanosomal principal styles of plumose skeleton robust, short, thick, slightly curved at centre, with rounded smooth or occasionally very faintly microspined bases, fusiform points, Length 84-(98.7)- 308 pm, width 4-(11.2)- 14, m. Acanthose choanosomal styles of reniernid skeleton straight or slightly curved ai centre, evenly spinose except for aspinose points and semetimes aspinose base; spines large, recurved, sharply pointed. Length 93-(104.5)-1124.m, width 8-(13.3)- 6pm. Subectosomal auxihary subtylostyles long, slender, straight, smooth or microspined bascs, fusiform ort occasionally with telescoped points. Length 102-(118.4)-152u m, width 2-(3.4)- 4.5um. Micrascleres, Palmate isochelae large, un- modified, with lateral and front alae approximate- ly equal length, lateral alae completely used to shaft, front ala detached along entire length. Length 16-(19.4)-21,.m. Toxas wing-shaped, generously curved at centre, with only slightly reflexed points. Length 36-(68.9)- 148 um, width 1.542.1)-3.01.m, REMARES. I initially thought this species con- specific with C. (7.) styloprothesis (see above), based on Topsent’s (1929) description of 'siyloprothése', whereby spongin fibres are replaced by algal filaments, but the two species differ in spicule geometry and skeletal architec- ture (see also A. (F) frondifera below). It differs from the allied A. (A.) tuberosa in growth form. 416 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 213. Antho (Antho) opuntioides (Lamarck) (paralectotype MNHNDT3418). A, Choanosomal skeleton. B, Characteristics of fibre and renieroid skeleton. C, Acanthostyles of renieroid skeleton. D, Acanthostyle spines. E-F, Bases of principal and auxiliary styles. G, Palmate isochelae. H, Wing-shaped toxas. REVISION OF MICROCIONIDAE 417 S, d 7 x £ AN Z E E UL JE S E NS Sf — f D/ Nou YE 2 X Lu = ^ 52 =) i PK 25um ll Sw Ss Je z/ — FIG. 214. Antho (Antho) tuberosa (Hentschel) (fragment of holotype ZMB4417). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Acanthostyles of renieroid skeleton. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, WAM648- 81(1). 418 spicule geometry (particularly its acanthoslyle morphology), and spicule sizes (refer to discus- sion in A, (A.) tuberosa). Antho (Antho) tuberosa (Hentschel, 1911) (Figs 214-215, Plate 9B-C) Lissodendoryx tuberosa Hentschel, 1911; 326-328, text-fig.21- fsoctona teberosa, Hallmann, 1920: 768; Burton & Rao, 1932: 341-342. Antho tuberosa: Hooper & Wiedenmayer, 1994: 255. Claihria harimeyeri Hentschel, 1911; 379-381, text- fig.50. Thalysias harimeyeri; de Laubenfels, 1936a: 105. cf. Microciona prolifera, Vosmaer, 1935a: 611, 648, 664. MATERIAL. HOLOTYPE: HM (fragment ZMB4A417): 4km SW. of Denham, Shark Bay, WA, 25756,5'S, 113*30.D' E, 3m depth, 10.vi.1905, coll. W. Michaelsen & R. Hartmeyer (dredge). OTHER MATERIAL: WA- WAM645-81(1) (fragment NTMZ1722) QMG300203 (fragment NTMZ2958), NTMZ3214 (fragments — PIBOCO4-352, QMG300044), QMG300678 (NCIQ66C-4229-N, NTMZI466. S AUST- SAMTS4050 [fragment NTMZ1637). HABITAT DISTRIBUTION. Encrusting on macro- phytes, coralline algae, corals, orother spenges; 3-4ürm depth; Shark Bay, Straggler Rocks, Houtman Ahrol- hos, Port Hedland (WA): Port Noarlunga (SA) (Fig. 214G); Gonjam, Madras, India (Burton & Rao, 1932). DESCRIPTION. Shape. Variable, thinly encrust- ing, up to 3mm thick. lobo-digitate, up to 25mm high, 42mm wide, 28mm thick, with rounded margins, or lobate, flabellate, with several flat- tened digits joined to a common base, and with curved, even or sinuous margins, without a basal stalk, 79mm high, 120mm maximum width, lobes between 18-32mm wide, up to 15mm thick. Colour. Bright red (Munsell 2.5R 5/10), dark red (SR 4/10) or slightly yellow-red alive (2.5 YR 8/8); yellow-brown or light brown in ethanol, Oscules. Indetectable in thinly enerusting specimen, scattered on lateral and upper surfaces of lobate specimens, up to 2.5mm diameter; sur- face minutely porous on lobate-flabellate specimen, pores 0.5-1.2mm diameter. Texture and surface characteristics. Texture compressible; surface optically even (encrusting- lobate specimens) or stated and pitted, with ridges most prominent near margins (Habellate specimen). Ectosome and subectoseme. Hispid, with tracts of smooth choanosomal styles from plumose skeleton protruding singly or in brushes; smaller MEMOIRS OF THE QUEENSLAND MUSEUM subectosomal auxiliary subtylostyles form tan- gential tracts (encrusting specimen), paratangen- tial tracts (lobate specimen), or distinct plumose brushes on ectosome, surrounding protruding choanosomal styles (flabellate specimen); subec- tosomal region undifferentiated from choanosomal skeleton; smaller acanthose choanosomal styles of renieroid skeleton ter- minate in uni- or paucispicular brushes just helow surface; mesohvl matrix in peripheral region light, unpigmented. Choanosome. Two distinct skeletal components: renieroid skeleton regular (encrusting-lobate specimens) or irregular (flabelliform specimen), rectangular and triangular meshes, tight meshed with mesh size 42-125,1.m diameter; meshes more open in lobate specimens (92-197 um} and flabellate specimens (97-208pm maximum diameter); yaguely differentiated primary and secondary components of skeleton, besi developed in flabellate specimens, with pnmary ascending (multispicular) and secondary transverse (uni- or bispicular) tracts of smaller acanthose choanosomal megascleres; encrusting and lobate specimens mesh size decreases and skeleton more compact, with poorer differentia- tion of primary and secondary lines; plumose skeleton of larger, smooth, choanosomal prin- cipal styles forming pauci- or multispicular tracts; plumose tracts continuous, originating from basal attachment extending 1o peripheral skeleton in encrusting-lobate material, but not obviously continuous in flabellate specimens, prominent only near periphery; true echinating acanthostyles absent; mesohyl matrix light, con- taining few loose extra-fibre spicules; spongin fibres 45-I08j.m diameter relatively light, only barely differentiated from mesohyl matrix in flabellate specimens (with only primary ascend- ing elements and small interconnecting secon- dary fibre components obvious); spicule skeleton only minimally associated with fibre skeleton, each forming more-or-less independent support systems; choanocyte chambers relatively large, 155-6521.m diameter, ovoid, often paired, lined by isochelae and rarer toxas. Megascleres. Smooth chpanosomal principal subtylostyles of plumose skeleton slightly curved, fusiform, with tapering, rounded or sub- tylote smooth bases, occasionally slightly microspined: very variable in length. Length 164- (213.0-337um, width 6.5-(9.9)- 1 3. üjum (holotype 126-295 x 6.5-9,um). Acanthose choanosomal styles of renicroid skeleton slightly curved or straight, subtytote, REVISION OF MICROCIONIDAE FIG. 215. Antho (Antho) tuberosa (Hentschel) (NTMZ3214). A, Choanosomal skeleton. B, Renieroid reticulate secondary skeleton. C, Acanthostyle of renieroid skeleton. D, Acanthostyle spines. E-F, Bases of principal and auxiliary styles. G, Palmate isochelae. H, Wing-shaped toxas. 420 fusiform, with evenly distributed small spines; acanthose choanosomal spicules include inter- mediates between larger smooth megascleres of plumose skeleton and entirely spined spicules of renieroid skeleton. Length 86-(114.9)-136,um, width 3.5-(6.3)-10.0j.m (holotype 63-128 x 2.5- 6pm). Subectosomal auxiliary subtylostyles of peripheral skeleton straight or slightly curved, single size category, thin, fusiform, with rounded or subtylote bases, usually microspined. Length 100-(177.4)-274um, width 1.8-(3.1)-4.5um (holotype 129-214 x 2.5-A4jum). Microscleres. Palmate isochelae common, un- modified, variable in size between specimens; lateral and front alae approximately equal length, lateral alae entirely fused to shaft, front ala only partially detached from lateral alae. Length 6- (13.5)-18j.m (holotype 10-15pm). Toxas wing-shaped, relatively common, rang- ing from small thin forms to long relatively thick forms, all with evenly rounded, wide central cur- vature, and straight or very slightly reflexed points. Length 37-(94.5)-232,um, width 0.8- (1.8)-3.6,.m (holotype 6-58 x 0.8-1.8,.m). Larvae. Larger flabellate-lobate specimen (WAM645-81(1)) contained ovoid-elongate, in- cubated embryos, 210-240j.m diameter, with some cellular differentiation and few larval megascleres. Associations. Several specimens had parasitic zooanthids on surface. REMARKS. This species is relatively polymor- phic in growth form, the extent to which the renieroid skeleton is compacted and size of isochelae, but more consistent in choanosomal and ectosomal skeletal architecture, spicule geometry, and to some extent spicule dimensions. However, there is no doubt that all specimens examined are conspecific, their similarities far more obvious than their apparent differences, especially in comparison to its sibling species A. (A.) opuntioides (from SW. Australia) and A. (A.) lithophoenix (from NW. Pacific). The lobate- encrusting specimen from SA shows the greatest departure from the holotype, particularly in spicule dimensions. Hentschel (1911) described this species with a renieroid skeleton enclosed, to a greater or lesser extent, within spongin fibres, but the holotype does not substantiate this. In flabellate specimens fibres are more obviously associated with plumose skeletal tracts cored by smooth choanosomal styles, whereas the renieroid MEMOIRS OF THE QUEENSLAND MUSEUM meshes appear independent of spongin fibres. Similarly, Hentschel did not describe toxas from the holotype but they are present being most abundant in SW WA flabellate-lobate specimens. Burton & Rao (1932) suggested that A. (A.) tuberosa was probably conspecific with Clathria hartmeyeri, indicating that it differed only in the encrusting growth form, lighter mesohyl matrix and in supposedly lacking smooth subtylostyles. Although the latter assertion is incorrect the holotype of C. hartmeyeri has not yet been dis- covered in any museum and this synonymy can- not be corroborated. From Hentschel’s (1911) description of C. hartmeyeri it appears to be identical to the flabellate-lobate form of A. (A.) tuberosa and this synonymy is accepted. Antho (A.) tuberosa is the type species of Isociona Hallmann's (1920), which Van Soest (1984b) merged with Antho. In the strict sense (i.e., comparing A. (A.) tuberosa and A. (A.) in- volvens) the two genera are close in skeletal ar- chitecture, but A. (A.) tuberosa has entirely monactinal megascleres, whereas A. (A.) invol- vens has basal (renieroid) monactinal and/or diac- tinal spicules (acanthostyles, acanthostrongyles). Also included in /sociona is Plocamia lithophoenix de Laubenfels (1927) which is very similar to A. (A.) tuberosa in skeletal structure, spicule diversity and spicule geometry, differing only in the basal spines on choanosomal styles plumose skeleton and specific spicule dimen- sions (smooth choanosomal styles 152-238x11- I5àm, acanthose choanosomal styles 129-166x8-14jum, subectosomal subtylostyles 133-293x3-6j1.m, palmate isochelae 18-26jum, and toxas 18-178x0.8-3,um). Both these and A. (A.) opuntioides are sibling species showing many skeletal and spicule similarities. OTHER SPECIES OF ANTHO (ANTHO) Antho (Antho) brattegardi Van Soest & Stone, 1986 Antho brattegardi Van Soest & Stone, 1986: 42-44, figs 1-3 [Norway] MATERIAL. HOLOTYPE: ZMAPORS190. PARATYPE BMNH1982.9.6.1. NE. Atlantic. species of Jia de Laubenfels. Antho (Antho) brondstedi Fromont, 1988 Antho brondstedi Bergquist & Fromont, 1988: 97, pls 46d-f, 47a-c [New Zealand]; Dawson, 1993: 44 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPORI11. New Zealand. Bergquist & Antho (Antho) dichotoma (Esper, 1794) REVISION OF MICROCIONIDAE Spongia dichotoma Esper, 1794: 202, pl.10 [Norway]; Ehlers, 1870: 8-9 [re-examination of type-material]. Not Spongia dichotoma; Johnston, 1842: 97; Topsent, 1920b: 21. Raspailia dichotoma; Ehlers, 1870: 8. Clathria dichotoma; Arnesen, 1903: 21-22, pl.3, fig.4, pl.6, fig.8 [Norway]; Thiele, 1903b: 394; Koltun, 1959: 184, pl.29, fig.1, text-fig.145 [USSR]; Stephens, 1916: 234; Stephens, 1921: 21 [Ireland]. Not Spongia dichotoma Lamarck, 1814: 448 [missing Topsent, 1933: 46, 56]. Dictyoclathria dichotoma; Topsent, 1920b: 21-22 [Horns Riff, Denmark]; Burton, 1930a: 501. Raspailia moebii; Schmidt, 1875: 120 [Norway]; Thiele, 1903b: 394. Dictyocylindrus abyssorum Carter, 1876: 232, pl.12, fig.3, pl.15, fig.25a-b [N. of Scotland]. Clathria abyssorum; Vosmaer, 1880: 154 [Faroe Is, N.Atlantic]; Arndt, 1913: 119. Raspailia abyssorum; Fristedt, 1885: 48, pl.4, fig.1. Antho dichotoma; Alander, 1942: 63 [Sweden]; Van Soest & Stone, 1986: 44 [Norway]. MATERIAL. HOLOTYPE: unknown; holotype of D. abys- sorum: BMNH1898.5.7.39. NE. Atlantic. Antho (Antho) graceae (Bakus, 1966) Li his le lacunosa; de Laubenfels, 1961: 195- 197. Not Myxilla lacunosa Lambe, 1892: 70-71. Anthoarcuata graceae Bakus, 1966: 431-432, text- fig.3 [San Juan Archipelago, Washington]; Ristau, 1978: 5737 [California]; Lee & Gilchrist, 1985: 24-32 [biochemistry]. Antho graceae; Van Soest, 1984b: 7 [generic synonymy]. MATERIAL. HOLOTYPE: USNM161848. NE. Pacific. Antho (Antho) hallezi (Topsent, 1904) Heteroclathria hallezi Topsent, 1904b: 94; Burton, 1935a: 403. Plocamia hallezi; de Laubenfels, 19362: 78. MATERIAL. HOLOTYPE: MOM (fragment MNHNDT1884). NE Atlantic. Antho (Antho) heterospiculata(Brondsted, 1924) Microciona heterospiculata Brondsted, 1924: 465, text-fig.20 [Colville Channel, NZ]. Quizciona heterospiculata; de Laubenfels, 19362: 111. Not Microciona heterospiculata; Bergquist, 1961a: 39 [probably = Clathria mortensenii Brondsted]. MATERIAL. HOLOTYPE: UZM (not found) (fragment BMNH1901.12.26.13). New Zealand. Antho (Antho) involvens (Schmidt, 1864) Myxilla involvens Schmidt, 1864: 37, 45, pl.4, fig.6 [Adriatic]; Heller, 1864: 48. Hymedesmia involvens; Schmidt, 1866: 16. Antho involvens, Gray, 1867: 524; Topsent, 1928a: 11; Topsent & Olivier, 1943: 2 [Monaco]; Burton, 1956: 133 [W. Africa]; Lévi, 1960a: 57, 76-80, text-figs 42] 19-22 [var. inconstans; Atlantic, Mediterranean]; Vacelet, 1960: 267 [Mediterranean]; Vacelet, 1961: 4] [Corsica, Mediterranean]; Sarà, 1961: 48 [Adriatic]; Sarà & Siribelli, 1962: 10, 36, 48; Lévi, 1963: 62-63, text-fig.72 [Mossel Bay, South Africa]; Sarà, 1964: 228-229 [Ligurian Sea, Mediterranean]; Poggiano, 1965: 3, 7; Rützler, 1965: 33-34 [Adriatic Sea]; Borojevic et al., 1968: 25; Descatoire, 1969: 196; Vacelet, 1969: 206 [Mediterranean]; Boury-Esnault, 1971: 326; Riedl, 1971: 1139 [ecology]; Pulitzer-Finali, 1977: 63 [Bay of Naples]; Rodriguez Solórzano & Rodriguez Babid, 1979: 56-58, text-fig.13 [var. inconstans; Galicia, Spain]; Pulitzer-Finali, 1983: 567-568, 610 [Mediterranean]; Boury-Esnault & Lopes, 1985: 195-196, fig.44 [Azores]; Pansini, 1987: 170 [AI- boran Sea]; Uriz et al., 1992: 104 [Balearic Is]; Solórzano et al., 1991: 177 [Galicia, Spain]; Ackers, Moss & Picton, 1992: 139 [Ireland]. Desmacodes involvens; Vosmaer, 1880: 108; Vos- maer, 1885: 235. Myxilla banyulensis, in part; Topsent, 1892b: 23; Top- sent, 1902: 351, 363, 366; Cotte, 1903: 423. Clathria morisca Schmidt, 1864: 37, 45 [Adriatic]; Schmidt, 1868: 9, 41, 43, pl.2, fig.7 [Mediter- ranean]; Vosmaer, 1880: 150-151 [Algiers]; Top- sent, 1902: 329. Dictyoclathria morisca; Topsent, 1920b: 18-21; Top- sent, 1928a: 301-302, pl.3, fig.3 [Porto Santos, Azores]; Lévi, 1959: 134, text-fig.27, pl.5, fig.1 [Rio de Oro, Gulf of Guinea]; Lévi, 1960b: 761-762, text-fig.15 [var. anisotyla; SW. Cape of Naze, W. Africa]; Sarà, 1960a: 462 [Ischia, Mediterranean]; Desqueyroux-Faundez & Stone, 1992: 35 [index]. Plocamia inconstans Topsent, 1925: 661-664, text- fig.15 [Gulf of Naples]; Topsent, 1939: 6; Pulitzer- Finali, 1983: 610 [list]. Plocamilla inconstans; Burton, 1935a: 402. Holoplocamia inconstans; de Laubenfels, 1936a: 75. Antho inconstans; Ackers et al., 1992: 140 [Ireland]. Isodictya beani Bowerbank, 1866: 274, 334, 335 [Britain]; Gray, 1868: 164; Schmidt, 1870: 77; Bowerbank, 1874: 147, p.58, figs 1-6. Dictyoclathria beanii, Arndt, 1935: 81. Amphilectus beanii, Vosmaer, 1880: 115. Clathria beanii, Ridley, 1881: 485, 486; Bowerbank, 1882: 13, 23, 150; Topsent, 1890c: 203. Myxilla beanii, Topsent, 1892c: 23; Topsent, 1894a: 8, 9, 25; Hanitsch, 1894: 179. Artemisina mediterranea Babic, 1921: 87 [Adriatic]; Babic, 1922: 258-259, text-fig.B; Burton, 1930a: 528; Lévi, 1960a: 57, 76-80; Maldonado, 1992: 1154 [possible synonym of A. (P.) novizelanica]. Microciona virgula Sarà & Siribelli, 1960: 77-79, text- fig.22 [Bay of Naples]; Siribelli, 1960: 16-17, text- fig.7A [Naples]; Sarà, 1964: 228-229 [Mediterranean]. ? Artemisina paradoxa Babic, 1921: 87; Babic, 1922: 260-261, pl.8, fig.6, text-fig.c [Adriatic]; Topsent, 1925: 660; Lévi, 1960a: 85-86 [Adriatic]; Ristau, 1978: 585-586 [note on affinities]. Clathria paradoxa; Burton, 19302: 528. Antho paradoxa, Pulitzer-Finali, 1983: 610. 7 Raspailia incrustans Svarcevskij, 1906: 52, pl.5, fig.6, p}-7, fig-1. MATERIAL: HOLOTYPE: LMJG {fragment BMNH1867.3.11.92), fragments of holotype of C, morisca: MNHNDT2170, BMNH1868.3.2.21. NW. Atlantic. Mediterranean, NW Africa, South Africa. This is probably a species complex (Ackers et al., 1992) contesting the synonymy between irvolvens and inconstans. Antho (Antho) oxeifera(Ferrer-Hernandez, 1921) Clathria oxeifera Ferrer-Hernández, 1921: t71, pl.l [Mediterranean]; Lévi, 1960a: 84-85 [incertae sedis; Mediterranean]; Pulitzer-Finali, 1983; 610 [list]. Labacea oxeifera; de Laubenfels, 1936a; 125 [7]. Anrho oxeifera: Uriz & Maldonado, 1993: 359-362, figs 6-9 ['W. Mediterranean]. MATERIAL. HOLOTYPE: Madrid. Mediterrancan. Antho (Antho) paucispinaSarà & Siribelli, 1962 Anrho paucispina Sarà & Siribelli. 1962; 48-51, text- fig.12 [Mediterranean]; Pulitzer-Finali, 1983; 610, MATERIAL. Holotype: IZUG. Mediterranean. Antho (Plocamia) Schmidt, [870 Plocamia Schmidt, 1870: 62. Dirrhopalum Ridley, in Ridley & Duncan, 1881: 477. Plocamiopsis Topsent, 1904a: 155. Wereroclathria Topsent; 1904b- 93. Lissoplocamia Brondsted, 1924: 470. Plocamilla Topsent, 1928a: 63; Lévi, 19602: 80. Atnomoclarhria; in part, Topsent, 1932: 103 (not Top- sent, 1929: 26), Holoplocamia de Laubenfels, 1936a: 75. TYPE SPECIES, Plocamia gymnazusa Schmidt, 1870; 62 (by subsequent designation of Burton, 19354: 401 )). DEFINITION. Regular basal or axial renieroid (and/or isodictyal) skeleton of acanthostrongyles (less frequently acanthostyles), with or without spongin fibres; renieroid tracts may be echinated bv acanthostyles at spongin fibre nodes; basal renieroid skeleton overlays leptoclathriid or microcionid main skeleton composed of echinat- ing (acantho-)styles and/or choanosomal styles, standing perpendicular to base or axis, joining with echinating megascleres to produce ascend- ing plumose skeletal tracts; extra-axial (subec- tosomal) skeleton plumose, dendritic, or subisodictyal, composed of choanosomal styles, nriginating from substrate or simply confined to periphery, forming tangential, paratangential or plumose extra-axial tracts; eclosomal skeleton MEMOIRS OF THE QUEENSLAND MUSEUM with or without specialised spiculation (1 or 2 categories of auxiliary styles); microscleres in- clude diverse forms of isochelae and toxas. REMARKS. Twenty one species have been referred to Plocamia or one of its synonyms: all are valid. However, A. (F) erecta is poorly known, and other species may eventually merge, particularly the 5 Indian Ocean species (Table 43). Only 2 species are known from Australasia. Antho (Plocamia) frondifera (Lamarck, 1814) (Figs 216-217, Table 43) Spongia frendifera Lamarck, 1814: 445; Lamvarek, 1816; 374, Anamoclathria frandifera; de Laubenfels, 19363- 108. Aurho frandifera, Hooper & Wiedenmayer, 1994: 256. Anomoclathria opuntioides var. frondifera, Topsent, 1929: 26-29, texr-figs 10-14; Topsent, 1932: 103, pl. H figs 6-7. l Hymeniacidon clifton? Bowerbank, 1862a: 773, pl.30, hig.¥; Bowerbank, 1864: 276, figs 70, 291. Acamia cliftoni: Gray, 1867; 515. Nut Alcyonium opuntioides Lamarck, 1815: 164. MATERIAL. LECTOTYPE: MNHNDTS565: Precise locality unknown, suspected to be SW. Australia (Tur- got collection). PARALECTOTYPE: MNHN- DT3356: same details. HOLOTYPE of H. cliftoni: BMNH!877.5.21.608 (fragments BMNHIR77.5.21. 616, 1185, 218): Precise locality unknown. SW Australia, coll. H, Clifton, HABITAT DISTRIBUTION. Ecology unknown; SW Australia (WA) (Fig. 216G). DESCRIPTION. Shape. Lobate, thickly flabel- late, digitate fans, 95-160mm long, up to 25mm thick, with uneven, digitate margins and irregular lobate surface (‘macroconules’) up to 6mm thick. No stalk remaining, if initially present. Colour, Grey or grey-brown in dry state, Oscules. Large, up to 4mm diameter, scattered evenly over surface and lateral margins of digits, with remnants of stellate drainage canals con- verging on each oscule. Texture and surface characteristics. Harsh, brittle in dry state. Uneven, lumpy surface with distinct collagenous crust. Ectosome and subectosome. Ectosomal skeleton membraneous, heavily collagenous in places (al- though rarely intact in dry specimens). with some embedded detritus and sparse tangential and paratangential tracts or single auxiliary subtylos- tyles scattered near periphery. sometimes form- ing bundles protruding through ectosome; points of (smooth) choanosomal principal styles from ascending plumose tracts protrude only slightly REVISION OF MICROCIONIDAE 50 um a a y 200 um FIG. 216. Antho (Plocamia) frondifera (Lamarck) (lectotype MNHNDTS65). A, Choanosomal principal sub- tylostyles. B, Acanthostrongyles of renieroid skeleton. C, Subectosomal auxiliary subtylostyles. D, Wing- shaped toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Paralectotype MNHNDT3356. through collagenous surface membrane; subec- tosomal skeleton virtually undifferentiated from choanosome, although peripheral choanosomal styles of plumose skeleton slightly more dense, diverging, than tracts in skeletal core. Choanosome. Skeleton with 2 distinct com- ponents: ascending plumose and basal/axial renieroid (in some places isodictyal) skeletons; plumose skeleton witth pauci- or multispicular tracts of smooth choanosomal principal styles ascending to surface, rarely branching or anas- tomosing; tracts associated with, but not neces- sarily coring, heavy, dark brown, spongin-coated algal filaments (ostensibly Ficus (Topsent, 1932)), which dominates skeleton; filaments up to 250j1.m diameter, 300-400j.m apart, branch- ing, diverging from base of sponge through sponge surface; renieroid skeleton composed of 1 or 2 acanthostrongyles abreast forming square or triangular meshes up to 120j.m diameter, even mesh size throughout skeleton, overlaying plumose skeleton; some detritus scattered be- tween renieroid skeletal meshes, usually coated with spongin; mesohyl not intact although some granular collagen containing microscleres scat- tered between spicule meshes; choanocytes not observed. Megascleres. Choanosomal principal styles en- tirely smooth, short, robust, slightly curved at centre, with rounded or slightly subtylote bases, fusiform points. Length 88-(103.4)-118jum, width 4-(8.1)- 131.m. Acanthostrongyles of the renieroid skeleton thick or thin, rounded or slightly subtylote at both 424 MEMOIRS OF THE QUEENSLAND MUSEUM Ch 58um A e | —— 43 asm DE A Huc yt = A = E - 9x Xung FIG. 217. Antho (Plocamia) frondifera (Lamarck) (paralectotype MNHNDT3356). A, Choanosomal skeleton. B, Fibre characteristics. C, Acanthostrongyles of renieroid skeleton. D, Acanthostrongyle spines. E-F, Bases of principal and auxiliary subtylostyles. G, Wing-shaped toxas. H, Palmate isochela. REVISION OF MICROCIONIDAE 425 TABLE 43. Comparison between species of Antho (Plocamia). Measuremenis in jim. ACP.) ridleyi eel m Sn TE Choanosornal 212-388 x ecd B8-118x | 120-660x styles 14-20 4-13 11-164 E 62-M75 739 | | (strongyles) Renieroid 92-104 x = a nieroi 6.5-10 4.5-15 spicules (strongyles | (strongyles Ectosomal 129- [uses x rrt x spicules i [uses | aw | - | er 2.5-13 107-248 x Subectosomal | 298-388 x spicules 47 Echinating | 170-202 x oo mo 13 75-158 x ETH mcr EH | 1520 | 20 via 46 x E UE MM 40-116x " aT x | 03- SFF 190-230 x 2. spined) m I Supe: 23-1n (spined) spined) Ridley & Duncan (1881; 481); Dendy (1922: 76); Lévi (19604: 81). 2 1960b: 760); Pulitzer-Finali Ty am Lévi a; 81). 6. Lévi (1952: 53) ends, heavily spined particularly at points, spines large. conical or slightly recurved, sharply pointed. Length 85-(95.2)- 1031.m, width 3-(7.8)- 14um. Subectosomal auxiliary subtylostyles long. very slender, curved at centre or sinuous, sub- tylote usually microspined bases, fusiform or oc- casionally telescoped points, Length 120-(135.9)-184pm, width 1-(1-7)-2.54m. Microscleres. Palmate isochelae large, un- modified, with front and lateral alàe approximate- ly same length, lateral alae entirely fused to shaft, front ala detached along lateral margin. Length 15-(17.4)-20].m. Toxas wing-shaped, short, moderately thick, with large central curvature and slightly reflexed points. Length 40-(67.6)- 11 6,.m, width 1-(1.3)- 2um. REMARKS. Spongin fibres are excluded from the skeleton and replaced entirely by algal fila- ments (‘styloprothesis’; Topsent, 1929), although each filament contains a thin cover of collagen on its surface with embedded spicules. Of the 3 microcionid species demonstrating this sym- biosis A. (A.) opuntioides, and A. (P.) frondifera MM 184 y | 425-430 x 1:25 27 160-550 x 82-15 70-120 x 3-10 al trong Tl 47153206. | 500x25 HUM NS 130-170x | 75-100 4 7 177 x 15.8 Aa [ 10 (styloid) | (strongyles) ERER = 190-260 x 140-1803 || tia xos | 2724x174 5-1] present’ iao | w» | 1: | w | nus | -W0 x " 5-75 x0.5- te ots &i3x2i |2s-70x& | 579305 {spined) 85-130x3 (spined) . Ridiey & Dendy (1957: 158); Dendy (1922 1973: 35). 3, À Ridley & Diwan (1881: hig 4. Ridley & Duncan (1861: 483). 5, are the major structural partners in the symbiotic relationship, whereas C, (T.) styloprothesis is probably a cryptic, invasive sponge and the algal symbiont provides the growth form structure. Antho (P.) frondifera is most similar to A. ÇA.) opuntioides, differing in spicule geometry (acan- thostrongyles versus acanthostyles in the renieroid skeleton), and spicule dimensions (Table 43). Neither species has been subsequently recorded since the early 1800s. Antho (Plocamia) ridleyi (Hentschel, 1912) (Figs 218-220, Table 43, Plate 9D-E) Plocamia ridleyi Hentschel,1912: 387-388, pl.20, fig.44. Holoplocamia ridleyi; de Laubenfels, 1936a: 75. Antho ridleyi, Hooper & Wiedenmayer, 1994: 256. MATERIAL. HOLOTYPE: SMF1709 (fragment MNHNDCL2183): Mimien Bay, Aru L, Arafura Sea, Indonesia, 6*5, 134*50" E, 15m depth, 8.1v.1908, coll. H. Merton (dredge). OTHER MATERIAL: NT- NTMZ0299, NTMZ2108, NTMZ2110, NTMZ2112, NTMZ2131, NTMZ2142, NTMZ2201, QMG300146 (fragment NTMZ2212), NTMZ2221, QMG300507 (fragment NTMZ2230), QMG303295, NTMZ2556, NTMZ2378. WA- QMG301185. 426 TABLE 43. (continued) ALP.) Choanosomal 225-884 x nes ewe -340x 13-17 AP. ALP) AMP) AP.) delaubenfelsi "m 7 inconstans penneyi (de plena (Li we) f Top Em) Lavbentea) (Sollas)! Burton)" Mabeltate a eem ena ee 3- MEMOIRS OF THE QUEENSLAND MUSEUM AP) | AAP (P.) AAP.) AAP.) barbadensis gyrmnazus (Van Soest)" L (Schmidt)! lambei 20-2 - l 8- Renieroid — | !12-153x9 | '? mans PEDE 122x7 | 14x12 | i038 |is301xS.| agxa | 95 005 " | spicules (strongyles) (steongyles | (sirongyles (strongyles) | (strongyles | (strongyles | R (styles) | (strongyles e aci les) | Ericssimal spicules Subectosomal spicules pa ict es [Chelas — | | n2 | | 1830 | 6-20 159-733x | 230-290 x 3-IT 4-5 TE] mee ne ama ana al | 744 | Ex a es $ “Eite (1963: 2 8. Bakus (1966: 440); Simpson (1968a:43). 9, Topsent (1925: 662). 10. de Laubenfels (1936a: 76). 11. Sollas (1879: 44). 12. Bakus (1966); Duncan (1881: 478). 15. Dend 4: 351 HABITATDISTRIBUTION. Holotype collected from à sand substrate, encrusting on an Oceanapia species (Niphatidae; incorrectly identified as Phivedictyon fis- niosa (Bowerbank)); other specimens collected from subtidal laterite rock and coral reefs, associated with dead coral substrate, usually growing on the underside of coral rubble; restricted intertidal distribution to only 3m depth; known Australian distribution: Darwin Har- bour (NT); Hibernia Reef, Sahul Shelf (WA) (Fig. 2181); also Aru Is, Indonesia (Hentschel, 1912). DESCRIPTION, Shape. Thinly, thickly or bul- bous-encrusting, 2-14mm thick, forming exten- sive overgrowths on coral substrata. Colour. Live colouration consistent, even pright red or blood red (Munsell SR 5-3/10), turning grey or pinkish grey in ethanol (SR 8/2-4). Oscules. Large oscules on exterior of bulbous lobes, 0.3-1.6mm diameter, with slightly raised membraneous lip; minute pores, up to 150m diameter scattered evenly over surface; pores and oscules contract upon dessication. Texture and surface characteristics. Firm, barely compressible, easily crumbled alive; no mucous produced upon exposure to air; surface optically smooth, irregularly bulbous. mostly clear of silt in situ; surface lobes in thinly encrusting specimens collapse upon dessication or preserva- tion. but thicker specimens retain shape. Ectosome and subectosome. Surface microscopi- cally hispid, with points of smooth chnanosomal Lambe (1895: 124). 13. Van Soest (1984b: 125). 14. Ridley in Ridley & principal styles protruding and surrounded by plumose brushes of mostly smaller ectosomal auxiliary subtylostyles (with fewer larger subec- tosomal megascleres contributing to ectosomal skeleton); subectosomal region structurally vari- able: thinly encrusting specimens with peripheral Skeleton not clearly delineated from choanosomal skeleton, containing only thick tan- genual or paratangential tracts, up to 140um diameter, composed of larger subectosomal auxiliary subtylostyles; in thicker bulbous specimens silbectosomal region cavernous, con- taining numerous plumose, stellate brushes com- posed of both of choanosomal and subectosomal megascleres, clearly distinguished from the renieroid component of choanosomal skeleton; subectosomal auxiliary megascleres also in deeper choanosomal skeleton, together with smooth choanosomal principal styles, together forming vaguely ascending, multispicular, extra- fibre tracts, 25-65y.m diameter. Chounosome. Skeletal structure with 3 distinct components: hymedesmoid skeleton, with basal layer of spongin fibre lying on substrate, 60- 240um thick, with smooth choanosomal prin- cipal styles and echinating acanthostyles perpendicular to substrate; renieroid skeleton forming regularly reticulation of acanthostron- gyles, overlaying hymedesmoid basal skeleton, in pauci- or multispicular tracts (vaguely ascend- REVISION OF MICROCIONIDAE ing} and uni- or paucispicular (irregularly transverse) tracts, producing triangular (isodic- tyal) or rectangular {renieroid) meshes, 60- 150um diameter, without any obvious spongin fibre component; echinaling acanthostyles, oc- curring singly or in plumose brushes, at major nodes of renieroid skeleton. sometimes also forming irregularly plumose, discontinuous, as- cending tracts: subisodictyal extra-fibre skeleton well developed in thicker specimens but rudimentary (irregularly dispersed) in thinly encrusting specimens; subisodictyal skeleton composed of both smooth choanosomal principal styles and subectosomal auxiliary subtylostyles forming barely continuous subisodictyal tracts extending from Jeptoclathnid basal skeleton to penpheral region, becoming more plumose or dendritic towards periphery; mesohyl matrix heavy but virtually unpigmented, surrounding renieroid meshes; choanocyte chambers circular to oval, $3-95um diameter, mesohyl matrix in both basal and peripheral regions more heavily Pigmented than in choanosomal region, and microscleres also more abundant near surface. Megascleres. Choanosomal principal styles long, thick, slightly curved, with rounded or slighily subtylote bases, smooth or with lightly microspined bases, fusiform points. Length 183- (317.9 -562,0.m, width | 6.5-(14.8)-25um fholotype 212-388 x 14-204cm). Acanthostrongyles of renieroid skeleton shor, thick, straight or slightly curved, with either sym- metncal subtylote bases, or asymmetrical ends (subtylote bases, rounded or slightly subtylote points); usually evenly microspined, spines small, conical, sharply pointed. Length 98- (120.6-142ym, width | 4.5-(10.1)- I 5pm (holotype 92-104 x 6.5-10pum). Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, with prominent tylote, subtylote or polytylote bases, bases microspined or less commonly smooth. Length 231-(372.9)- 473pm, width 2.5-(5.9)-12.5m (holotype 298- 388 x 4-7pm). Ectosomal auxiliary subtylostyles identical an geometry but smaller than large auxiliary megascleres, with tylote or subtylote, smooth or mücrospined bases. Length 100-(192.7)-252,.m, width 1.1-(3.3)-6.5j.m (holotype 129-209 x 1.8- um). Echinating acanthostyles long, thick, fusiform, slightly curved, with subtylote, lightly microspined bases, entirely smooth shafts or proximal half of shafts covered with small spines (these spicules possibly small morphs of pnn- 40 cipal styles). Length 107-(194.1)-248,.m, width 5-(10, D-15j.m (holotype 170-202 x 8-13jm). Micrascleres. Palmate isochelac abundant, single size Category, unmodified, with front and lateral alae approximately same length, lateral alae com- pletely fused to shaft, front ala detached from front alae along lateral margin. Length 8-(12.5)- 19m (holotype 11-18,.m). Toxas wing-shaped, incompletely differen- trated into 2 size classes: smaller thin, extensive rounded ceniral curvature, straight or slighily reflexed points; larger thick, sharply angular or slightly rounded central curvature, straight or very slightly reflexed arms charactenstically bearing terminal bulbous swelling and a single apical spine surrounded by smaller spines. Length I: 2)-(44.3)-86,.m, width 1-(1.9)-4.] um (holotype 24-46 x 0.8-1.24 m); length [I- 62- (169.2)-355 jm, width 2-(4.7)- 10.6&.m (holotype 82-211 x 24pm). Larvae. Parenchymella larvae oval-elongate, 195-4]01.m long, 135-330üm wide, orange- brown alive, incompletely ciliated with à bare pesterior pole. Younger larvae poorly differen- tated in cellular construction, bul containing clearly visible, longitudinally disposed, whispy sinuous monactinal megascleres. Older larvae have distinct. oval. cellular differentiation. radial- ly disposed thin tvlostyles, and small toxas and isochelac. Incubated larvae were abundant in 33% of specimens, collected between January and May 1985 in the Darwin region, bit absent from samples collected during Latter part of year (Fig. 220). Probable reproductive period is wet-early dry seasons (January to June) but determination of complete reproductive period was not possible as samples could not he taken every month). Associations. All NT specimens were encrusting dead faviid coral heads, growing next to, or over other sponges (e.g., Srelleita, Mycale, Placospon- gia, Ulosa, Clathria), coralline algae and as- cidians, Variation. Ectosomal structure variable, ranging from Clathria condition (with thin oc thick fans genüal crust; 17%), Thalysias condition (with a stellate plumose ectosomal architecture; 50%), to merely paratangential (with elements of both structures; 339%), Subectosomal region cavern- ous (with plumose tracts of choanosomal and/or subectosomal megascleres: 67%), to merely tan- gential or paratangential (without subdermal cavities; 33%). Extra-axial (non-renieroid) skeleton ranges from subisodictyal, distinctly plumose, ascending, diverging towards periphery 100 um MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 218. Antho (Plocamia) ridleyi (Hentschel) (NTMZ2142). A, Choanosomal principal style. B, Echinating acanthostyles. C, Acanthostrongyles of renieroid skeleton. D, Subectosomal auxiliary subtylostyles. E, Ec- tosomal auxiliary subtylostyle. F, Accolada and wing-shaped toxas. G, Palmate isochelae. H, Section through peripheral skeleton. I, Australian distribution. J, NTMZ299. (5896), to irregularly disposed, mostly lon- gitudinal extra-fibre tracts (48%), Echinaung acanthostyles form plumose ascending structures (50%) or irregularly dispersed (50% ). Spicule geometry consistent although spicule dimensions varied slightly for all specimens (holotype had smaller toxas and acanthostrongyles than NW Australian material). REMARKS. This species is relatively common in cryptic habitats on shallow intertidal reefs in the Darwin region, and it is therefore surprising that it has not been found elsewhere in NW. Australia despite extensive sampling in similar habitats along the N. coast. Dendy (1922) merged Plocamia ridleyi with Plocamilla coriacea from the N. Atlantic and Mediterranean, but this REVISION OF MICROCIONIDAE 429 FIG. 219. Antho (Plocamia) ridleyi (Hentschel) (QMG301185). A, Choanosomal skeleton. B, Renieroid skeleton (x437). C, Acanthostrongyles of renieroid skeleton. D, Acanihostrongyle spines. E, Echinating acanthostyle. F, Acanthostyle spination. G, Base of subectosomal auxiliary subtylostyle. H, Spined toxa point. I, Palmate isochelae. J, Accolada and wing-shaped toxas. - " NO. SAMPLES SEASON TOTAL SAMPLES WITH LARVAE i | PEDRY | 1 ,| Án J| p mr [LL aes FIG, 220. Antho (Placamia) ridleyi (Hentschel), In- cidence of incubated parenchymella larvae in NT specimens. synonymy is clearly wrong. In their spicule diver- sity and growth form the two taxa are similar (Table 43), whereas comparisons between field observations on living populations of A. (P) rid- leyi (present study) and A. (P) coriacea ( Ackers, Moss & Picton, 1992: 141) show that the two species have quile different surface features, live colouration, and some differences in spicule dimensions (Table 43) indicating at most a pos- sible sibling species relationship. The separate category of echinating acanthos- tyles, a renieroid skeleton composed of diactinal or quasi-diactinal spicules, and à more-or-less plumose (non-renieroid), subisodictyal skeleton of smooth choanosomal and subectosomal spicules are typical of Antho (Plocamia). How- ever, Antho and Plocamia are barely differen- tiated on that basis and they are formally merged here. Some Plocumia have a mixture of both acanthostyles and acanthostrongyles in the renicroid skeleton (Lévi, 1960a). Megascleres echinating fibre nodes may vary from true acan- thostyles, with different geometry from other choanesomal spicules (A. (P) barbadensis (Van Soest, 1984b)), or smooth styles which are dif- ferentiated from choanosomal megascleres only by their marginally smaller size and lighl, ir- regular spinatpon (most species including A. (F) ridleyi, A. (P.) elegans (Ridley & Dendy, 1886) and A. (P) coriacea (Bowerbank, 1874)). or echinating megascleres may be entirely undif- ferentiated from choanosomal styles or absent (e.g.. A. (P) navizelanicum (Ridley, in Ridley & Duncan, 1881), A. (P) penneyi (de Laubenfels, 19362), A. (P) frondifera (Lamarck)). Similarly, the extra-axaal (non-renieroid) skeleton varies be- tween specimens. Some thinly encrusting species, such as A. (P.) ridleyi, A. (P.) delauben- felsi (Little, 1963) and A. (P.) burtoni (Lévi, 1952) have choanosomal styles embedded in the basal spongin fibre and in the renieroid fibre nodes, ascending all the way to surface in more-or-less plumose tracts. Other species (e.g., A. (P) illgi MEMOIRS OF THE QUEENSLAND MUSEUM (Bakus, 1966)) have plumose tracts of choanosomal styles mainly in the peripheral skeleton with the remainder of the skeleton being simply renieroid. Antho (Plocamia) ridleyi dif- fers substantially from the other known Australian species A. (P) frondifera in spicule geometry (particularly in having spined points on the larger toxas), spicule dimensions (Table 43), growth form and lacking algal filaments in the skeleton (*styloprothesis’). OTHER SPECIES OF ANTHO (PLOCAMIA). Antho (Plocamia) barbadensis (Van Socst, 1984) Plocamilla barbadensis Van Soest, 1984b: 125-126, lext-Fig.50 [Barbados, West Indies]. Antho barbadensis; Yan Soest & Stentoft, 1988: 123 [Barbados]. MATERIAL. HOLOTYPE: ZMAPOR3832. Province Caribbean. Antho (Plocamia) burtoni (Lévi, 1952) Plocamilla burtoni Lévi, 1952: 53-54, text-fig. 17 |Senegal, W. Africa]; Lévi, 1960b: 760 [note]. MATER HOLOTYPE: MNHN missing, Province: NW, Ca. Antho (Plocamia) circonflexa (Lévi, 1960) Plocamilla circonflexa Lévi, 19602: 81-83, text-figs 24-25 [Brest, France]; Sarà & Siribelli, 1960: 80 {Bay of Naples, Mediterranean]; Sarà & Siribelli, 1962- 51 [Gulf of Naples]: Descatoire, 1966; 242, text-liz.6B [Glenan Archipelago, Brittany]; Pulit- zer-Finali, 1983: 610 [list]. MATERIAL. HOLOTYPE: MNHN missing, NE. Atlanlic, Mediterranean. Antho (Plocamia) coriacea (Bowerbank, 1874) Isodictya coriacea Bowerbank, 1874: 136, 228, pl.76, figs 7-12 |Britain]. Dirrhopalum cariaceum; Ridley, 1881:481, pl.29, figs 3-7 [Ireland]. Plocamia coriacea; Hanitsch, 1894: 173 [Britain); Dendy, 1922: 76-77 [Amirante, Indian Ocean). Plocamilla coriacea; Topsent, 19284; 63; Burton, 1935a; 402; Burton, 1959b; 44 [Iceland]. Lévi. 1960a: 80-81, text-fig.23 [Roscoff, English Chan- nel; Atlantic]; Sarà & Siribelli, 1962: 51 [with ques- tion; Gulf of Naples]; Poggiano, 1965: 3,7. Pulitzer-Finali, 1983; 610 [list]; Solórzano et ul,. 1991; 177 [Galicia, Spain]; Ackers, Moss & Picton, 1992: 141-142 [Ireland]. i Holoplacamia coriacea; dè Laubenfels, 1936a; 75. MATERIAL. Holotype: BMNHIR877.5.21. 761/1910.- 1.1.2541, NE. Atlantic, Mediterranean, E. Africa. Antho (Plocamia) delaubenfelsi (Little, 1963) Holoplocamia delaubenfelsi Little, 1963: 45-48, lext- fig.18 [Gulf of Mexico]. REVISION OF MICROCIONIDAE MATERIAL. HOLOTYPE; USNM23596, NE. Pacific. Antho (Plocamia) elegans(Ridley & Dendy, 1886) Plocamia elegans Ridley & Dendy, 1886; 475 [var.]: Ridley & Dendy, 1887: 158-159, pl.29, fig.9, pl.31. fig.1 [var. elegans; Azores]; Topsent, 1892a: 117, pl.7, fig. 11 [var, elegans; Azores]; Topsent, 1904a: 155 [var. elegans; Azores). Plocamia elegans; Dendy, 1922: 77-78 [Cargados Carajos, Indian Ocean]; Topsent, 1928a: 64. Plocamilla elegans; Burton, 1935a: 402; Pulitzer- Finali, 1973: 35-41 [Azores]. Plocamilla coriacea var. elegans; Lévi, 1960b: 760- 761, text-fig. 13 [W. coast of Africa]. Holoplocamia elegans; de Laubenfels, 1936a: 75. MATERIAL. HOLOTYPE: BMNH1887.5.2, 109, NE Atlan- tic, E & W Africa. Antho (Plocamia) erecta(Ferrer-Hernandez, 1923) Plocamia erecta Ferrer-Hernández, 1923: 248, text- figs 1-3 [Spain]. Plocamilla erecta; Burton, 19353; 402. Holoplocamia erecta; de Laubenfels, 1936a: 75 |San- tander, Atlantic]. Antho erecta; Lévi, 19603: 80. MATERIAL. HOLOTYPE: Madrid. NE Atlantic. Antho (Plocamia) gymnazusa (Schmidt, 1870) Plocamia gyminazusa Schmidt, 1870; 62-63, pl.4, fig.17 [Florida]; Burton, 1935a: 401; de Laubenfels, 19362: 76. Dirrhopalum gymnazon; Ridley, 1881: 478-479, pl.29, figs 1-2. MATERIAL. HOLOTYPE: BMNH1870.5.370 (fragment MNHNDCLI IO0SL). Caribbean. Antho (Plocamia) illgi (Bakus, 1966) Plocamilla illgi Bakus, 1966: 440-443, pl.1A, figs 6a-j [San Juan Archipelago, Washinglon}, Simpson, 1968a: 43-47, 93, text-fig.3 [Sun Juan Is, Washington]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Bakus & Green, 1987; 73-74 [S. California]. MATERIAL. HOLOTYPE: USNM23737. NE Pacific. Antho (Plocamia) lambei (Burton, 1935) Plocamia manaarensis, in part; Lambe, 1895; 124 [California]; Lambe, 1900; 161. Not Dieryocylindrus manaarensis Carter, 1880a: 37. Heteroclathria lambei Burton, 1935a: 403. Plocamilla zimmeri Bakus, 1966: 512, MATERIAL. HOLOTYPE; USNM6331, NE Pacific. Antho (Plocamia) lithophoenix (de Laubentels, 1927) Plocamia lithephoenix de Laubenfels, 1927: 268. Isociona lithophoenix, de Laubenfels, 1932: 99-100, text-fig.59 [California]; Burton, 19352: 400 [note]; 431 Dickinson, 1945: 23, pl.35, figs 69-70, pl.36, figs 71-72 [Pacific Grove, California]. Antho lithophoenix, Van Soest, 1984b: 129 [generic synonymy for fsociona]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Sim & Bakus, 1986: 11 [California]. MATERIAL. HOLOTYPE: USNM21460, paratype BMNH1929,8,2242. NE Pacific, Antho (Plocamia) manaarensis (Carter, 1880) Dictyocylindrus manaarensis Carter, 18802: 34, pl.4. fig. | [Gulf of Manaar, Ceylon]. Dirrhopalum manaarense; Ridley, 1881: 482. Plocamia manaarensis; Dendy, 1905; 179, pl.8, fig.] [Gulf of Manaar, Ceylon]; Burton & Rao, 1932: 355 |Laccadive Sea, Mangalore and Karwar, India. Not Plocumia manaarensis, Lambe, 1895: 124, pl.2, figs | la-g [California]. Plecamilla manaarensis; Burton, 1935a: 402; Burton, 19592: 252-253 [Arabian Sea]; Bakus, 1966: 512. MATERIAL. HOLOTYPE: LFM destroyed (fragment BMNH19806.4,29.1b). India, Arabian Sea Antho (Plocamia) novizelanica (Ridley, 1881) Dirrhopalum navizelanicum Ridley, 1881: 483-485, pl.29, figs 8-16 [Bay of Islands, New Zealand]. Plocamilla novizelanicum:; Burton, 1935a: 402, Plocamilla novizelanica; Lévi & Lévi, 19839: 965- 966. Lext-fig.27 [S. of New Caledonia]; [?] Uriz, 19882: 90-9], text-fig.65 [Namibia; ? affinity]. Holoplocamia novizelanica, de Laubenfels, 19362: 75. Plocamia novizelanicum; Bergquist & Fromont, 1988; 120-122, pI.56, fig.f, pl.57, figs a-b [New Zealand : Dawson, 1993: 38 [index to fauna]. Not Plocamilla cf, novizelanica, Maldonado, 1992: 1154, fig. 11-12, table 5 [Alboran Sea; ? affinity]. MATERIAL. HOLOTYPE: BMNH1964.1.1.1. SW Pacific (N Z, New Caledonia); ISW Afnca, Mediterranean. Antho (Plocamia) ornata (Dendy, 1924) Bubaris ornata Dendy, 1924a: 351, pl.14, figs 25-27. Plocamia ornara; Burton. 1928: 129. Axoplocamia ornata; Burton, 1935a: 402. MATERIAL. HOLOTYPE: BMNH missing (fragments BMNHI923,10,.1.126, 322). W Indian Ocean, Antho (Plocamia) penneyi (de Laubenfels, 1936) Holoplocamia penneyi de Laubenfels, 1936a: 76 | Tor- tugas, Florida]. Antho penneyi; Van Soest & Stentoft, 1988: 126 {table}. MATERIAL. HOLOTYPE; USNM22460), Caribbean. Antho (Plocamia) plena (Sollas, 1879) Plocamia plena Sollas, 1879: 44, pls 6-7 [W Atnea); Topsent, 1894: 21, Holoplocamia plena; de Laubenfels, 1936a: 75 [note]. Clarhria plena; Vosmaer, 1880: 154 | Angola]. Dirrhopalum plenum; Ridley, 1881; 480-481. MEMOIRS OF THE QUEENSLAND MUSEUM 500 um 25um FIG. 221. Antho (Isopenectya) chartacea (Whitelegge) (holotype AMZA36). A, Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Acanthostyle of renieroid skeleton. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. REVISION OF MICROCIONIDAE MATERIAL. HOLOTYPE: BMNHI1909,8.15.3). W Africa. Antho (Plocamia) prima (Brondsted, 1924) Lissoplocamia prima Brondsted, 1924: 470, fig.24a-d [North Cape, New Zealand]; Topsent, 1928a: 63: Lévi, 1963: 63, fig. 73 [S Africa]. Plocamia prima, Bergquist & Fromont, 1988: 122, pl.57c-e. MATERIAL. HOLOTYPE: possibly UZC. South Africa, NZ. Bristo] (fragment Antho (Plocamia) signata (Topsent, 1904) Plecamtopsis signala Topsent, 19043: 155-157, pl.14, fig.] [Azores]; Topsent, 1928; 306-307, pl.10, fig.20 [et var. mitis; W. of Flores, Azores}; Burton, 1935a; 402 [note]. MATERIAL. HOLOTYPE: BMNH1930.7.1.36). NE Atlantic. MOM © (fragment Antho (Isopenectya) Hallmann, 1920 Isopenecrya Hallmann, 1920: 789, Clathriella Burton, 1935c: 73; Koltun, 1959: 186. TYPE SPECIES. Clathria chartacea Whitelegge, 1907: 497 (by monotypy). DEFINITION. Three skeletal components: (1) renieroid reticulation of acanthose styles, (2) overlayed by isodictyal or subisodictyal reticula- tion of smooth styles coring spongin fibres, (3) surmounted by plumose or radial extra-axial skeleton of larger smooth styles, perpendicular to axis, in peripheral region; skeleton may be slight- ly compressed at core, spongin fibres only moderately developed; echinating megascleres absent; ectosomal skeleton with single category of auxiliary subtylostyle forming tangential or paratangential tracts; microscleres absent. REMARKS. Jsopenecrya contains 4 species, 3 from the SW Pacific and | from the NW Pacific, All lack microscleres bur this is interpreted as secondary loss. Antho (Isopenectya) chartacea (Whitelegge, 1907) (Figs 221-222, Plate 9F) Clathria (?) chartacea Whitelegge, 1907: 497. Isopenecrya chartacea; Hallmann, 1920: 789. Antho chartacea; Rudman & Avern, 1989: 335; Hooper & Wiċdenmayėr, 1994; 255, Antherochalina perforata Lendenfeld, 1887b: pl.22, fig.44. Nol Antherochalina perferaia, in part; Lendenfeld, 1 887b: 788; Lendenfeld, 1888: 89-90. MATERIAL. HOLOTYPE: AMZ436: Off Coogee, NSW, 33°45’°S, 151°20'E, 98-100m depth, date of 433 collection unknown, coll. FIV ‘Thetis’ (trawl). HOLOTYPE of A. perforata: BMNH1886.8.27.459: Broughton I,, Port Stephens, NSW, 32?36'5, 152*]9'E, other details unknown.OTHER MATERIAL. NSW- NTMZ2831, AMZ3605, AMZ3604, AMZ3606, AMZ4216 (RRIMPFN 1339), AMZÁ4256 (RRIMPFENI435), AMZ4255 (RRIMPFN1434), AMZ3207, AMZ3162, AMZ4S69 (RRIMP-S9PIP), OMG30371 L, QMG303713. HABITAT DISTRIBUTION, 12-100m depth: rock platform, heads or outcrops on sand substrate, known only [rom Australia: Port Stephens, Botany Bay. Coogec, Long Reef, Dee Why, N. Sydney, Port Hack- ing, Cronulla, Manly (NSW) (Fig. 221E). DESCRIPTION. Shape. Thinly flabellate, up to 80mm long. 55mm wide, with long, thickly cylindrical stalk, very thin lamellae, up to 8mm thick, with slightly digitate or evenly rounded margins. Colour: Bright red-orange alive (Munsell 5R 5/10 - IOR 6/10], pale brown in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, barely compressible. flexible. slightly spiky: optically smooth, even surface. Evtesome and subectosome. Ectosome prominently hispid, with pauci- or multispicular plumose brushes of larger, smooth choanosomal principal styles protruding through surface, form- ing a vestigial radial extra-axial skeleton, arising from pauci- or multispicular tracts of (marginal- ly) smaller smooth principal styles in subec- tosomal region; subeetosomal auxiliary subtylostyles tangential, paratangential, or rarely plumose, at base of protruding choanosomal spicule brushes; peripheral skeleton relatively cavernous in comparison to the central choanosomal skeleton, with moderately heavy mesohyl matrix. Cheanosome. Skeletal structure with 3 distinet components: (1) slightly compressed spongin fibres forming close-meshed anastomoses at core of skeleton, more cavernous towards surface, (2) renieroid skeleton composed of acanthose styles, overlaying other structures; (3) longitudinal, as- cending tracts of smooth principal styles, mar- ginally smaller than those protruding through surface, forming subisodictyal tracts at core, more plumose in periphery, and usually (but not invariably) associated with larger spongin fibres: spongin fibres in axial skeleton heavy, 48-82 4m diameter, producing irregularly oval or elongate meshes, 32-120,.m diameter, cored by uni- or bispicular tracts of smaller, smooth chognosomal principal styles; fibres closer to surface, 19-421.m 434 diameter, regularly anastomosing, wide-meshed, 75-162,.m diameter, forming regularly renieroid (triangular) spicule meshes and oval or elongate fibre meshes, cored by uni- or bispicular tracts of smaller acanthose styles; plumose extra-fibre skeleton composed of uni-, pauci- or multi- spicular ascending tracts of smooth choanosomal styles standing perpendicular to axis, becoming increasingly plumose, larger, and typically mul- tispicular towards periphery; echinating megascleres absent; mesohyl matrix lightly pig- mented, with few auxiliary spicules scattered throughout; choanocyte chambers elongate-oval, 36-75,.m diameter. Megascleres. Smooth choanosomal principal styles long, thick, slightly curved or straight, with rounded or very slightly subtylote bases, rarely with basal microspination, fusiform points. Length 117-(232.4)-312jum, width 6-(11.8)- 15pm (holotype 168-274x13-17 um). Acanthose choanosomal styles of renieroid skeleton short, thick, fusiform, slightly curved or straight, with rounded or slightly subtylote bases, lightly microspined bases and points, with fewer spines scattered on shaft, occasionally complete- ly smooth shaft. Length 74-(86.1)-112j.m, width 4-(7.2)-8.51um (holotype 92-127x9-12.5jum). Subectosomal auxiliary subtylostyles short, thin, usually straight, with prominent subtylote, typically microspined bases, hastate points, abrupt points, or sometimes telescoped or bifid points. Length 134-(183.6)-203jum, width 2.5- (2.9)-3.8.m (holotype 163-243x2-4.5 um). Echinating megascleres absent. Microscleres. Absent. Larvae. Viviparous, parenchymella larvae oval to elongate, 340-420x180-360j.m, with central core of juvenile styles, well differentiated cellular con- struction. Associations. Obligatory (?) host for nudibranch Rostanga sp. (AMC150065) (W. Rudman, pers.comm.). REMARKS. Hallmann (1920) erected /sopenec- tya for this species based on a renieroid skeleton, with two categories of choanosomal styles, without echinating acanthostyles, and without microscleres. The type species has affinities with Antho but differs from other 'plocamid' microcionids (with myxillid-like renieroid skeletons) (viz. Antho s.s., Dirrhopalum, Plocamilla, Plocamiopsis, Labacea, Isociona, and /sociella) in having a compressed axis and more-or-less plumose extra-axial skeletons cored by smooth choanosomal (principal) styles, in one MEMOIRS OF THE QUEENSLAND MUSEUM or more size categories, together with the usual renieroid structure overlaying the remainder of the skeleton composed of acanthose (or some- times smooth) styles different from principal spicules. This species superficially resembles Oph- litaspongia tenuis (Carter) (= Echinoclathria leporina (Lamarck)) mainly due to the emphasis of the compressed central skeleton and sub- renieroid skeletal structure in both species, whereas megascleres forming these skeletons are quite different. Choanosomal megascleres in A. (L) chartacea are differentiated: small acanthose styles forming the renieroid skeleton (not echinating fibres), small smooth styles forming a secondary radial ascending skeleton, and larger smooth styles forming the peripheral perpen- dicular skeleton. By comparison, in E. leporina there is a smaller size class of smooth principal style both coring and echinating heavy spongin fibres, forming a renieroid skeletal structure, and a second, larger class of smooth principal style forming a sparse radial or plumose peripheral skeleton (embedded in peripheral fibres). This latter structure links the two groups. E. riddlei sp. nov., is also similar in skeletal structure but lacks spined spicules in renieroid skeleton and has a vestigial extra-fibre skeleton perched on the outer surface. Antho (I.) chartacea should be con- trasted with the renieroid Amphinomia (Raspailiidae), which also has acanthose struc- tural spicules (Hooper, 1991). Antho (Isopenectya) punicea sp. nov. (Figs 223-224, Plate 10A) MATERIAL. HOLOTYPE: QMG304399: Mrs Watson’s Bay, midway in bay, Lizard I., Qld, 14?39,5' S, 145?26.7 E, 18m depth, 10.iv.1994, coll. J.N.A. Hooper et al., SCUBA. HABITAT DISTRIBUTION. Sand, coral rubble, Halimeda bed substrata, in depression in sand; 18m depth; Lizard I. (FNQ) (Fig. 223E). DESCRIPTION. Shape. Bushy, subspherical, bulbous clump, 195mm long, 142mm maximum width, 138mm maximum height, composed of individual, erect, digitate projections, each up to 16mm diameter, 75mm high, forming reticulated structure, attached to coral rubble and Halimeda on base. Colour. Dull red alive (Munsell 5R 6/8), light brown in ethanol. Oscules. Small, up to 2mm diameter, mainly on lateral sides of digits, situated at junction of sur- REVISION OF MICROCIONIDAE 435 FIG. 222. Antho (Isopenectya) chartacea (Whitelegge) (QMG303711). A, Choanosomal skeleton. B, Fibre characteristics. C, Acanthostyle of renieroid skeleton. D, Acanthostyle spines. E-F, Bases of principal and auxiliary subtylostyles. G, Variability in auxiliary spicule points. 436 face aquiferous canals, surrounded hy collapsible membraneous lip Texture and surface characteristics. Firm, com- pressible. not easily torn; surface turgid in life, with distinctive ectosornal membrane. anerial- like longitudinal aquiferous canals obvious on extemal surface, branching and interconnecting, Opening into common oscules, porous surface between canals; canals, ridges and oscules col- lapse in air, producing reticulate surface upon dessication; produces abundant red mucus upon exposure. Ectosome and subectosome. Surface prominently hispid, with longer choanosomal principal styles embedded in peripheral fibres, arising from as- cending primary, plumose spicule tracts, extend- ing through surface for most of their length; subectosomal auxiliary subtylostyles tangential, occasionally paratangential, confined to exterior collagenous layer below ectosome, occasionally protruding through surface in plumose brushes; mesohyl matrix heavy in peripheral region. Choanosome. Skeleton without any compression or marked differentiation between core or subec- tosomal regions; 3 distinct skeletal components: (1) renieroid skeleton composed of both acan- those styles and smaller smooth principal styles in uni-, bi- or paucispicular tracts, coring small, light spongin fibres up to 25pm diameter, produc- ing rectangular or triangular meshes up to 902m diameter: (2) plumose, diverging Meu of smaller smooth choanosomal principal styles in multispicular ascending tracts, diverging towards periphery producing nearly radial skeletal tracts; (3) and with larger, smooth principal styles em- bedded in peripheral skeleton perpendicular to surface; echinating megascleres absent; mesohyl matnx hight, without microscleres bul few whispy (? juvenile) auxiliary subtylostyles scattered be- tween fibre meshes; choanocyte chambers small, oval. 25-45p.m diameter: Megascleres, Smooth choanosomal principal stvles lang or short, slender, slightly curved at centre, with rounded, predominantly smooth bases, occasionally microspined, telescoped points. Length 86-(155.6)-235p.m, width 2,5- (3.3)-Aj.m. Acanthose styles of renieroid skeleton slender, slightly curved towards base, rounded, sparsely microspined bases, sparsely spined shaft, spines srnall, erect, conical; points of spicules fusiform. Length 88-(114.9)-1531.m, width 2-(3.6)-61.m. Subectosomal auxiliary subtylostyles variable in length and thickness but only comprising a single category; bases subtylote, microspined, MEMOIRS OF THE QUEENSLAND MUSEUM tuberculale (granular) or occasionally smooth, points fusiform or slightly telescoped; whispy juvenile forms present scattered throughout mesoóhyl. Length 78-(169.8)-296,.m, width 0.5- (1.6)-31m. Echinating spicules absent. Microscleres. Absent. ETYMOLOGY. Latin punicews , reddish. REMARKS. The bulbous growth form, red colour and production of abundant mucus is com- mon to many other Indo-west Pacific mictocionids (such as C. (Isociella) eccentrica, C. (Thalysias) vulpina, C. (T) hirsuta, Echinoclatliria axinelloides, and Echinochalina (Protophlitaspongia) bargibunit). but this species belongs to Antho (Isopenecrya) having a renieroid skeleton composed (mainly) of a spe- cial category of acanthose styles (geometrically different from choanosomal spicules), a secon- dary, diverging, plumose skeleton of smaller, smooth choanosomal styles, and larger smooth choanosomal styles embedded in the penpheral skeleton. This latter character is reminiscent of Echinoclathria, and it could be argued for its inclusion in this genus on this basis, but the possession of 3 distinctive skeletal structures and acanthose spicules forming the renieroid skeleton support its inclusion in Antho. Antho (J.) panicea differs from A. (1.) chartacea in growth form, absence of axial skeletal coni- pression, spicule geometry and spicule sizes. The bases of auxiliary spicules in this species are also unusual, varying from swollen bases with prominent ierminal spines, granular tubercular swellings, or occasionally completely smooth. Antho (Isopenectya) saintvincenti sp. nov. (Figs 225-226) MATERIAL, HOLOTYPE: SAMS710(TS4035) (fragments QMG300486, NTMZ1671); Lead Light, Port Stanvac, St. Vincent Gulf, SA, 35°06'S, 138°27'°B, 7m depth, 16.xi.1977, coll. J. Window & H. Rapp (SCUBA). HABITAT DISTRIBUTION, Substrate unknown: 7m depth: St. Vincent Gulf (SA) (Fig. 225F). DESCRIPTION. Shape. Erect, arborescent, lamellate-digitate sponge, 235mm long, 130mm wide, with flattened or slightly cylindrical digits, up to 80mm long, 11mm diameter (cylindrical portions), or up to 20mm diameter, 8mm thick (lamellate portions of digits), repeatedly bifur- cate, rarely anastomosing. expanding towards REVISION OF MICROCIONIDAE 437 = WES SRA = SES iN FIG. 223, Antho (Isopenectya) punicea sp.nov. (holotype QMG304399). A, Choanosomal principal styles. B, Acanthostyle of renieroid skeleton. C, Subectosomal auxiliary subtylostyles. D, Section through peripheral skeleton. E, Known Australian distribution. F, Holotype. 438 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 224. Antho (Isopenectya) punicea sp.nov, (holotype QMG304399). A, Choanosomal skeleton. B, Fibre characteristics. C, Acanthostyle of renieroid skeleton. D, Acanthostyle spines. E-F, Bases of principal and auxiliary subtylostyles. G, Points of subectosomal auxiliary subtylostyles. REVISION OF MICROCIONIDAE spatula-like ends; short cylindrical basal stalk, 45mm long, 8mm diameter, and expanded basal attachment. Colour. Beige-brown in ethanol. Oscules. Small, probably contractile, 1-2mm diameter in preserved state, on edges of flattened digits. Texture and surface characteristics. Firm, com- pressible, flexible; surface smooth, even, unorna- mented, finely porous in preserved state. Ectosome and subectosome. Ectosome membraneous, microscopically hispid, with larger, smooth principal styles protruding through surface individually or in sparse, erect, plumose brushes arising from terminal subisodictyal spicule tracts; subectosomal auxiliary subtylos- tyles also protruding through surface in associa- tion with longer principal styles, in paratangential or plumose tracts; mesohyl matrix in peripheral skeleton light, poorly pigmented. Choanosome. Skeleton regularly renieroid reticulate, slightly more compressed at core than periphery, with 3 components; (1) renieroid skeleton composed of differentiated axial and extra-axial regions; axial fibres heavy, homogeneous, without clearly differentiated primary or secondary elements, 40-60j.m diameter, slightly more bulbous at fibre nodes, 70-90,.m diameter; all axial fibres cored by uni- or paucispicular tracts of acanthose principal styles forming rectangular or less often triangular meshes, 70-100jum diameter; extra-axial fibres lighter, with differentiated primary, ascending fibres, 20-40j.m diameter, cored by paucispicular tracts of both acanthose principal styles and smaller, smooth principal styles, becoming in- creasingly plumose towards surface, projecting from fibre nodes in particular as plumose brushes; primary fibres interconnected by uni- or paucispicular tracts of acanthose principal styles coring light spongin fibres, 15-30j.m diameter; (2) plumose, diverging skeleton of smaller smooth choanosomal principal styles intermixed with acanthose spicules in primary ascending tracts, diverging towards periphery, together producing nearly radial skeletal tracts; (3) larger, smooth principal styles in plumose brushes protruding through surface, embedded in ascend- ing primary fibres; echinating megascleres ab- sent; mesohyl matrix heavy but only lightly pigmented, with both fully formed and raphidiform subectosomal auxiliary subtylos- tyles scattered between fibre meshes; choanocyte chambers small, oval, 40-50j.m diameter. 439 Megascleres. Smooth choanosomal principal styles of plumose and radial surface skeleton long or short, thick or slender, slightly curved at centre, rounded or slightly subtylote, smooth bases, fusiform points, entirely smooth shaft. Length 78-(115.4)-1561um, width 4-(6.9)-10j.m. Acanthose styles of renieroid skeleton short, thick, slightly curved at centre, subtylote microspined bases, fusiform pointed, evenly microspined shaft, spines small, granular. Length 76-(84.6)-98 um, width 4.5-(5.9)-8 um. Subectosomal auxiliary subtylostyles short or long, slender or raphidiform, straight or slightly curved at centre, prominently subtylote, smooth bases, fusiform points. Length 66-(115.7)- 198um, width 1-(1.7)-3p.m. Microscleres. Absent. ETYMOLOGY. For the type locality. REMARKS. This species resembles both Antho and Echinoclathria, having a renieroid architec- ture and larger, smooth principal styles protrud- ing through the surface. Like A. (/.) punicea it is included in Antho because it has a special category of acanthose styles (geometrically dif- ferent from choanosomal spicules) forming the renieroid skeleton, and a secondary, diverging, plumose skeleton of smaller, smooth choanosomal styles. Antho (/.) saintvincenti dif- fers from A. (L) punicea in its flattened or cylindrical-digitate, arborescent growth form, reminiscent of Echinoclathria chalinoides, al- though spiculation and skeletal architecture differ substantially. OTHER SPECIES OF ANTHO (ISOPENECTYA) Antho (Isopenectya) primitiva (Burton, 1935) Clathriella primitiva Burton, 1935c: 73-4, text-fig.6 [Sea of Japan]; Koltun, 1958: 67 [Kuriles]; Koltun, 1959: 186, text-fig.148 [USSR]. MATERIAL. HOLOTYPE: BMNH1938.7.4.93 (fragment BMNH1932.11.17.69). NW Pacific. Echinoclathria Carter, 1885 Echinoclathria Carter, 1885f: 355. Ophlitaspongia; of authors; (not Bowerbank, 1866: 14). TYPE SPECIES. Echinoclathria tenuis Carter, 1885f: 355 (by subsequent designation ofBurton, 19342: 562), =Spongia leporina Lamarck, 1814: 444. DEFINITION. Two distinct skeletal components: (1) predominantly renieroid reticulate main skeleton cored by smaller, smooth principal MEMOIRS OF THE QUEENSLAND MUSEUM 440 FIG. 225. Antho (Isopenectya) saintvincenti sp.nov. (holotype SAMTS4035). A, Choanosomal principal style. B, Acanthose subtylostyle and modified style of renieroid skeleton. C, Subectosomal auxiliary subtylostyle. D, Section through peripheral skeleton. E, Holotype. F, Australian distribution. REVISION OF MICROCIONIDAE 441 FIG. 226. Antho (Isopenectya) saintvincenti sp.nov. (holotype SAMTS4035). A, Choanosomal skeleton, B, Fibre characteristics (x303). C, Acanthose subtylostyle of renieroid skeleton. D, Acanthostyle spines. E, Bases of principal subtylostyles. F, Base and point of subectosomal auxiliary subtylostyles. 442 styles, echinated by identical spicules (oc- casionally absent), typically very well developed spongin fibres sometimes slightly compressed at axis. more openly reticulate towards periphery; and (2) a vestigial radial extra-axial skeleton per- ched on the external surface, barely extending into choanosome, consisting of larger, smooth principal spicules, with identical geometry to those at core, forming radial or plumose brushes on surface; ectosomal skeleton with single size class of auxiliary subtylostyle lying paratangen- tially or embedded perpendicular to surface; microscleres include toxas and palmate isochelae. REMARKS. Sixty nine species have been in- cluded in Echinoclathria (or one of its synonyms), but only 23 are appropriately referred here. Fourteen species are known from Australia, most restricted to temperate coasts, 5 are new, Echinoclatliria 15 similar to Antho (Isopenec- rya), as noted above, differing in having only 2 skeletal components: a relatively homogeneous renieroid choanosomal skeleton composed of smaller, smooth principal spicules, amd vestigial radial extra-axial skeleton on Ihe external surface. Isopenectya has in addition à renicroid skeleton of acanthose spicules, and the smooth principal styles form longitudinal tracts extending all the way from the axis to the surface and beyond. Difficulties occur when trying to place species that have reduced structural characters: A. (4) punicea sp. nov. with spined renieroid spicules: E. riddlei sp, nov. with smooth renieroid spicules; both species with a reduced extra-fibre skeleton. Within Echinoclathria most of the variability centres around rhe development of the extra-fibre skeleton. In some species (e.g., E. leporina, E. canfragosa) there are obvious size differences between principal styles coring fibres in the choanosome and those protruding through the surface, whereas in others (e.g., E. nodosa) there is no obvious size differences between principal styles at the core and those at the periphery, although structurally these are similar to the first condition, In others (c.g.. E. egena, E. wal- daschmitt) there ts further reduction whereby the extra-fibre skeleton is virtually absent and all spicules are vestigial, poorly silicified. Groupings based on growth form (Hallmann, 1912) show little relationship to groupings made on skeletal characteristics. Thus previous clas- sifications for Echineclarhria are rejected here, MEMOIRS OF THE QUEENSLAND MUSEUM Echinoclathria axinelloides (Dendy, 18965) (Figs 227-228, Plate 10B) Ophlitaspongia axinelloides Dendy, 1896: 39: Hallmann, 1912: 268-270, pl.36, fig. 3, text-fig.58: Burton, 19342: 599, Echinochalina axinelloides; de Laubenfels, 19362: 119. Echinoclathriaaxinellnides; Carpay, 1986: 22; Hooper & Wiedenmayer, 1994; 279. MATERIAL. HOLOTYPE: NMVG2318 (fragment BMNH1902.10.18.342): Port Phillip, Vic, 380095. 144?*52'E, 36m depth, coll. J.B. Wilson (dredge), OTHER MATERIAL: VIC- AMZ802, AMZI593. TAS- QMG300269 (NCIQ66C-3655-O) (fragment NTMZ3804). HABITAT DISTRIBUTION. Rock reef: 20-36m depth; Port Phillip Bay (Vic); Furneaux Is (Tas) (Fig. 227D). DESCRIPTION, Shape. Erect, club-shaped nr arborescent, up to 115mm long, 75mm wide, with thick subeylindrical branches or slightly flattened lamellae, up to 34mm diameter, with rounded even margins, long thick basal stalk, 25-40mm long, 15mm diameter, slightly expanded basal attachment. Colour. Colour deep red alive (Munsell 2.58 4/10), pale brown in ethanol. Oscules. Numerous, moderately small, 2-4mm diameter, mainly on lateral margins of lamellae or on I side of branches, slightly raised with membraneous lip. Texture and surface characteristics. Firm, com- pressible, rubbery in life; surface optically smooth, minütely reticulated, with distinct membraneous covering. Ectosame and subectosome. Membraneous, with minutely reticulate, skin-hke membrane stretched over surface, microscopically hispid from protruding choanosomal styles forming well developed, multispicular plumose brushes just below surface producing a more-or-less con- tinuous palisade; surface spicule brushes heavier at surface than at core of skeleton; fewer subec- tosomal auxiliary styles paratangential to surface in association with oscules. Choanesome. Skeleton more-or-less dendro- reticulate, slightly sub-renieroid or irregularly reticulate in some parts, composed of heavy, well developed spongin fibre system incompletely separated into plumose primary and vestigial transverse secondary components; primary fibres (75-148,.m diameter) multispicular, cored by distinctly plumose tracts of choanosomal prin- cipal styles, whereas secondary fibres (38-72um REVISION OF MICROCIONIDAE la -— TS = p. SN N ^ = 250yum — g p ot 25um 443 s LL n ME — FIG. 227. Echinoclathria axinelloides (Dendy) (holotype NMVG2318). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary style. C, Section through peripheral skeleton. D, Australian distribution. E, Holotype. F, Larva in situ. diameter) uni-, bi- or occasionally aspicular; echinating styles sparsely dispersed on fibres mainly confined to primary fibres; fibre anas- tomoses form oval, elongate or rectangular meshes, 38-195j.m diameter, more compact in peripheral skeleton, relatively cavernous towards axis (130-275um diameter), fibre nodes slightly enlarged, bulbous; choanosomal fibres and 444 spicule tracts become more regular and more compacted towards periphery; mesohyl matrix relatively heavy, granular, with oval to eliptical choanocyte chambers (56-216j.m diameter), with both choanosomal and subectosomal megascleres scattered between fibres. Megascleres. Choanosomal principal styles and subtylostyles, coring and echinating fibres short, thick, straight, with smooth, evenly rounded or very slightly tapering, constricted bases, slightly hastate points, and approximately 5-15% modified to diactinal or quasi-diactinal forms. Length 76-(109.8)-148p m, width 4-(9.8)- 14.5j1.m. Subectosomal auxiliary styles straight, slightly curved, or rarely sinuous, relatively thick, with smooth, rounded or very slightly subtylote bases, fusiform or slightly telescoped points. Length 96-(144.3)-1874m, width 2.5-(4.1)-5j.m. Microscleres. Absent. Larvae. Viviparous, parenchymella larvae oval to elongate, 155-275x120-170j.m, without larval spicules. REMARKS. This species differs from other Australasian Echinoclathria in its growth form and rubbery texture, having a well developed ectosomal membrane covering a thick ectosomal palisade of principal styles, a dendro-reticulate skeletal structure verging on subrenieroid, its fibre characteristics and spiculation. It is most similar to E. nodosa in spicule geometry and gross skeletal architecture although differs in most other respects. Hallmann (1912) suggested his specimen (AMZ802) differed from Dendy's (1896) description having greater fibre diameter, less extensive spicule tract development, and a denser ectosomal skeleton, but comparison be- tween both specimens showed them to be clearly conspecific (i.e., supposed discrepancies were a consequence of Dendy's incomplete description). The collector of AMZ1593 is unknown; the AM register indicates Port Phillip, Vic. That specimen contained numerous small parenchymella larvae. According to Burton (19342) the Saville Kent collection contains this species, but this record is questionable as the specimens have not been dis- covered in the BMNH collections. Echinoclathria bergquistae sp. nov. (Figs 229-230, Plate 10C) MATERIAL. HOLOTYPE: QMG303872: S. of Tri- angle Reef, Hook Reef, Whitsunday Is region, 19?49.2'S, 149?07.1'E, 28m depth, 09.xii.1993, coll. J.N.A. Hooper & L.J. Hobbs (SCUBA). PARATYPE: MEMOIRS OF THE QUEENSLAND MUSEUM QMGL952 (fragment NTMZ1534): E. of Murdock 1., Howick Group, Great Barrier Reef, 14?36'S, 145°03’E, 14m depth, 18.ix.1979, coll. A. Kay (trawl). HABITAT DISTRIBUTION. Coral reef, coral rubble; 14-28m depth; Howick Reefs (FNQ); Hook Reef (MEQ) (Fig. 229F). DESCRIPTION. Shape. Erect or clumped, clathrous digitate mass, 90-110mm high, 65- 150mm wide, attached directly to substrate without basal stalk, composed of fused lobate or vaguely cylindrical digits, up to 55mm long, 30mm wide. Colour. Bright red alive (Munsell 2.5R 5/10), pale brown in ethanol. Oscules. Small, upto 3mm diameter, with slightly raised membraneous lip alive, scattered on ex- terior surface of lobate digits. Texture and surface characteristics. Soft, com- pressible, fibrous, difficult to tear, produces slight, clear mucus alive (on deck), stains ethanol orange; surface highly clathrous with large, flat- tened lobate or pointed conules covering exterior surface of digits, 5- 15mm long, up to 5mm wide; surface porous in preserved state, membraneous alive. Ectosome and subectosome. Surface prominently hispid with longer, smooth choanosomal prin- cipal styles embedded in peripheral fibres, ex- tending nearly 70% of their full length through surface; near bases of protruding principal styles are relatively heavy multispicular tracts of subec- tosomal auxiliary subtylostyles, usually tangen- tial to surface; mesohyl matrix in peripheral skeleton heavy but only lightly pigmented; choanosomal fibres extend directly to surface. Choanosome. Skeleton irregularly renieroid reticulate, slightly compressed at axis, with renieroid structure partially obscured by both larger principal styles echinating and subec- tosomal auxiliary subtylostyles scattered throughout mesohyl; spongin fibres large, 40- 60j.m diameter, well developed but only lightly invested with spongin, without any marked dif- ferences between thickness of fibres at core or surface; ascending fibres approximately same thickness as transverse fibres but generally longer and containing more coring spicules; ascending fibres pauci- or multispicular, cored by smaller choanosomal principal styles, with 2-5 spicules per tract; transverse connecting fibres generally shorter, containing the same spicules, 1-3 spicules per tract, and both fibres sparsely echinated by same spicules; fibre anastomoses produce elongate-oval meshes; axial fibre REVISION OF MICROCIONIDAE 445 FIG. 228. Echinoclathria axinelloides (Dendy) (holotype NMVG2318). A, Choanosomal skeleton. B, Fibre charac- teristics. C, Principal style. D, Ends of principal style. E, Subectosomal auxiliary style. F, Ends of auxiliary style. reticulation slightly more compressed than peripheral skeletal network, with meshes up to 90um diameter in axis, 160m diameter near surface; in addition to renieroid skeleton of smaller principal styles, larger principal styles also core ascending fibres and echinate fibre nodes especially closer to surface, forming sparse plumose bundles; mesohyl matrix heavy but only lightly pigmented, containing numerous toxas; choanocyte chambers oval, 35-55jum diameter. 446 Megascleres. Choanosomal principal styles (coring and echinating fibres) variable in length, straight or slightly curved at centre, with rounded bases, predominantly smooth but occasionally microspined, fusiform points. Length 71-(149.8)- 309m, width 2.5-(5.6)-1 2pm. Subectosomal auxiliary subtylostyles long, slender, straight, subtylote, smooth or less com- monly microspined bases, fusiform points; numerous smaller and raphidiform styles also scattered through mesohyl presumably being younger forms, Length 203-(356.7)-480pm, width 2-(3.7)-6yum. Microscleres. Palmate isochelae not common, relatively Jarge. with short thin alae, lateral alae completely fused to shaft, front ala nearly com- pletely detached from lateral alae, shaft straight. Length 18-(24.2)-325 m. Toxas wing-shaped, relatively thick, with slightly rounded central curvature, slightly reflexed points. Length 32-(49.7)-68m, width 0.5-(1.9)-3.0n m. ETYMOLOGY. For Dame Professor Patricia Bergquist for her work on Indo-Pacific sponges: REMARKS. Generic placement is not straight forward, with affinities to Echinoclathria and Antho (Isopenectya). The smaller, smooth choanosomal principal styles coring and echinat- ing all spongin fibres, producing an irregularly renieroid reticulation, a vestigial radial skeleton of larger, smooth principal styles protruding through the surface, and a slightly compressed axial region are typical of Echinoclathria, and in this respect the species is similar to E. leporina. However, the larger principal styles coring the ascending spongin fibres, occasionally echinat- ing fibre nodes, is reminiscent of Antho Usopenectya). It is included in Echinoclathria because the ascending tracts of larger principal styles do not form a subisodictyal skeleton; rather, these spicules end abruptly at fibre nodes in sparse plumose brushes and usually do not form continuous tracts. This evidence it weak and illustrates the difficulty in separating some species in both genera. This species is distinct from other Echinoclathria in toxa morphology and plumose brushes‘tracts of larger principal styles within the choanasome. MEMOIRS OF THE QUEENSLAND MUSEUM Echinoclathria chalinoides (Carter, 1885) (Figs 231-232) Axinella chalinoides Carter, 1885F: 358: Carter, 18862: 377 [et var glutinosa, cribrosa]. Axinella cladoflagellata Carter, (836g: 377. Echinochalina chalinoides, de Laubenfels, 19362: 119. Ophlitaspongia chalinoides, Dendy, 1896: 36, Echinoclathria chalinoides, Hooper & Wiedenmayer, 1994: 279, Not Ophlitaspongia chalinoides; Hallmann, 1912: 270-272, text-fig.59. MATERIAL. LECTOTYPE: BMNH1886.12. [5.401 (dry): Port Phillip, Vic, 38°09"S, 144°52°E, coll. J.B. Wilson (dredge), PARALECTOTYPES: BMNH1886.12.15.402 (dry) (fragment AMG2900a): same locality. BMNH1886.12.15.403 (dry) (fragment AMG2900b): same locality. HOLOTYPE of A. cladoflagellata: BMNHI886.12.15.407: same locality às lectotype. HABITAT DISTRIBUTION. Ecology unknown; Port Phillip (Vic) (Fig. 231D). DESCRIPTION. Shape. Arborescent branching, up to 250mm long, 170mm maximum width, with small basal stalk up to 45mm long, 22mm diameter, long cylindrical branches up to 105mm long, 9mm diameter, slightly flattened, bifurcat- ing repeatedly, rarely anastamosing. Calour. ‘Dull brick-red" alive (Carter, 1885e), pale brown in ethanol and dry. Oscules. Large. up to 4mm diameter, scattered mainly on lateral sides of branches, with series of radial subectosomal drainage canals radiating towards each oscule, and slightly raised membraneous lip. Texture and surface characteristics. Soft, com- pressible but difficult to tear. flexible branches, more rigid stalk; surface slightly microconulose, with fine surface network of radiating spicules associated with aquiferous system. Ectosome and subectosame. Membraneous, with points of larger principal styles protruding through surface, singly or in paucispicular brushes, for up to 30% of their length, and also with subectosomal auxiliary styles lying paratan- gential to surface; subectosomal region slightly cavernous, meshes up to 450m diameter, sub- stantially more wider-meshed than in axial region skeleton, Choanosome. Skeletal architecture dendro- reticulate, vaguely subrenieroid and more regular towards periphery than axis, composed of heavy, well developed spongin fibres, 25-70um diameter, thicker and slightly bulbous at fibre REVISION OF MICROCIONIDAE 447 = | FIG. 229. Echinoclathria bergquistae sp.nov. (holotype QMG303872). A, Principal style/ subtylostyles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyles. C, Wing-shaped toxas. D, Palmate isochela. E, Section through peripheral skeleton. F, Known Australian distribution. G, Paratype QMGL952. H, Holotype. 448 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 230. Echinoclathria bergquistae sp.nov. (holotype QMG303872). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Bases of principal and auxiliary styles. E, Wing-shaped toxa. F, Palmate isochela. nodes, without any marked differentiation be- tween primary and secondary components but substantially compressed in axis and diverging near surface; ascending fibres slightly sinuous, larger than transverse fibres, containing pauci- or multispicular core of more-or-less plumose tracts of choanosomal styles; in periphery these tracts form radial bundles and composed predominant- ly of longer spicules whereas towards core of skeleton coring spicules generally smaller and contained mainly within fibres; transverse con- necting fibres uni-, bi- or aspicular; echinating REVISION OF MICROCIONIDAE 100 um 500 um 449 FIG. 231. Echinoclathria chalinoides (Carter) (lectotype BMNH1886.12. 15.401). A, Principal styles and anisos- tyles (coring and echinating fibres). B, Subectosomal auxiliary styles and anisostyles. C, Section through peripheral skeleton. D, Australian distribution. E, Lectotype. megascleres not definitely present, although choanosomal principal styles protrude through fibres at oblique angles ('quasi-echinating"); fibre anastomoses form circular, polygonal or triangular meshes, 90-320j.m diameter in axis; mesohyl matrix heavy but only lightly pig- mented, with ovoid choanocyte chambers (90- 120um diameter), and numerous subectosomal auxiliary styles dispersed throughout. Megascleres. Choanosomal principal styles, anisostyles or anisoxeas (asymmetrical), thin, slightly curved at centre, occasionally straight, entirely smooth, bases rounded or slightly taper- ing, sometimes subtylote or telescoped, with has- 450 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 232. Echinoclathria chalinoides (Carter) (lectotype BMNH1886.12.15.401). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Bases of principal and auxiliary anisostyles. tate or telescoped points, rarely completely Subectosomal auxiliary styles, anisostyles or modified to quasidiactinal forms (symmetrical anisoxeas with similar geometry to principal spicules but generally longer, more slender, ends). Length 176-(264.7)-325ym, width 8- straight or occasionally slightly curved or (10.8)-13j.m. sinuous, with smooth rounded bases, sometimes REVISION OF MICROCIONIDAE telescoped, fusiform or telescoped points. Length 243-(315.5)-365 2m, width 4-(5.3)-8.m. Microscleres. Absent. REMARKS. This species is restricted to the type material following re-examination of a number of other specimens assigned here (Hallmann, 1912) which were misidentifications. Hallmann's (1912: 270) concept and illustrations are rejected here. It is presumed that he based his descriptions on several older AM specimens from Port Phillip, allegedly donated to the Museum by Carter but their spiculation and skeletal structures are quite different from the types (see E. subhispida). Echinoclathria chalinoides has a markedly compressed axial skeleton composed of heavy fibres and close-meshed spicule tracts together forming a dendro-reticulate skeleton; the skeleton becomes very wide-meshed near the surface, composed of poorly developed fibres and spicule tracts become more plumose; and coring/echinating spicules are predominantly anisostyles. Dendy (1896) suggested that it was a synonym of E. subhispida given their similarities in having a Haliclona-like branching growth form, distribution of oscules on lateral margins, soft compressible texture, and very heavy spon- gin fibres producing a compressed axial skeleton. However, there are major differences between these species in spicule geometries and skeletal architectures showing that they are not closely related. Echinoclathria confragosa (Hallmann, 1912) (Figs 233-234) Ophlitaspongia confragosa Hallmann, 1912: 255-257, pl.35, fig.2, text-fig.53. Axociella confragosa; de Laubenfels, 1936a: 113. Echinoclathria confragosa; Hooper & Wiedenmayer, 1994: 279. MATERIAL. HOLOTYPE: AMZ992 (dry): Shoal- haven Bight, NSW, 34?49'S, 151°04’E, 30-90m depth, 1.vii.1911, coll. FIV ‘Endeavour’ (trawl). HABITAT DISTRIBUTION. Epizootic on Bryozoa; 30-90m depth; S. coast (NSW) (Fig. 233F). DESCRIPTION. Shape. Irregularly digitate, lamellate branches, up to 50mm high, 5mm thick, arising from semi-encrusting base; branches vary from cylindrical to flattened lamellate, bifurcat- ing and anastomosing, forming loose reticulate mass, with shaggy lobate surface projections on points of branches. Colour. Grey-brown in dry state. Oscules. Not seen. 451 Texture and surface characteristics. Firm, com- pressible, brittle, fibrous; surface porous, pitted, slightly arenaceous. Ectosome and subectosome. Membraneous ec- tosome, microscopically hispid, with principal subtylostyles erect on peripheral fibres, singly or in bundles of up to 3 spicules, protruding through surface for most of their length and forming a sparse, vestigial, radial extra-axial skeleton; sub- ectosomal auxiliary styles form tangential and paratangential tracts on surface; thickness of ec- tosomal skeleton ranges from tangential (three spicules abreast) to paratangential (tracts of up to 20 spicules at obtuse angles to surface, forming low microconules). Choanosome. lrregularly renieroid reticulate skeleton, more-or-less homogenous throughout; spongin fibres thin, relatively light, 18-48j.m diameter, without any obvious differentiation be- tween primary or secondary components; fibres cored by uni- or paucispicular tracts of choanosomal principal styles, occasionally aspicular, echinated by choanosomal styles spar- sely and irregularly dispersed over fibres; fibre meshes predominantly rectangular (=renieroid), less often oval or triangular (=isodictyal), 112- 345jum diameter, slightly more cavernous at core and more compacted in peripheral regions of skeleton; mesohyl matrix heavy but only lightly pigmented, oval choanocyte chambers 52- 110j.m diameter; numerous microscleres, subec- tosomal auxiliary styles, and also few choanosomal styles scattered between fibres. Megascleres. Choanosomal principal subtylos- tyles (coring and echinating fibres) short, thick, straight, with smooth, slightly constricted sub- tylote bases, almost hastate points, slightly rounded, telescoped or pointed. Length 142- (164.8)-197j.m, width 5-(9.6)-121.m. Subectosomal auxiliary styles long, thin or thick, straight, slightly curved or sinuous, with smooth, slightly subtylote or rounded bases, fusiform points, sometimes slightly telescoped. Length 136-(214.4)-291.m, width 3.5-(5.1)- 6pm. Microscleres. Palmate isochelae abundant, small, poorly silicified, lateral alae completely fused to shaft, front ala detached from lateral alae for most of length, both alae sculptured with marginal ridges; shaft without any curvature. Length 8- (10.6)-14um. Toxas oxhorn, small, thick, with slight to moderate, evenly rounded central curvature, straight arms, slightly reflexed points. Length 32-(58.2)-95 1m, width 1-(2.2)-3.5jum. 452 25um MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 233. Echinoclathria confragosa (Hallmann) (holotype AMZ992). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary style and subtylostyle. C, Palmate isochelae, D, Oxhorn toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype. REMARKS. Echinoclathria confragosa is only known only from a single specimen which differs notably from all other species in several impor- tant respects: encrusting lamellate-digitate growth form; more-or-less homogeneous renieroid (Haliclona-like) skeletal architecture (see also E. notialis sp. nov.); retaining only the vestiges of a radial extra-axial skeleton on the extreme outer edge of the ectosomal region; having a relatively thick tangential and paratan- gential ectosomal skeleton covering most of the surface; and distinctive oxhorn toxa morphology. REVISION OF MICROCIONIDAE Echinoclathria digitata (Lendenfeld, 1888) (Figs 235-236, Plate 10D) Thalassodendron digitara Lendenfeld, 1888: 223. Echinochalina digitata; Thiele, 1903a: 962; Hooper & Wiedenmayer, 1994: 277. MATERIAL. HOLOTYPE: Missing from AM and BMNH collections. NEOTYPE-QMG304763: NW. of Snake Reef, Howick Gp., Great Barrier Reef, Qld, 14?28.6'S. 145?04.6' E, 21m depth, 03.ix.1994, coll. J.A. Kennedy (trawl), HABITAT DISTRIBUTION. Soft substrata inter-reef region; 21m depth; Torres Strait and Howick Reefs (FNQ) (Fig. 235E). DESCRIPTION. Shape. Bushy, clathrous mass of erect, bifurcate digits 54mm high. 76mm wide: digits short, stout, claviform, subcylindrical, bul- bous, expanding and slightly flattened towards apical extremities, up to 32mm long, 6mm wide, bifurcating several times, occasionally anas- tomosing, with ! or more blind branches; mass growing from a semi-encrusting common base. Colour, Bright red alive (Munsell SR 4/10), grey- brown in ethanol. Oscules. Small, up to 3mm diameter, on apex of each digit. Texture and surface characteristics. Firm, com- pressible, flexible, fibrous; surface bulhous, prominently microconulose, hispid. Ectosome and subectosome. Membraneous, granular collagenous heavier than in choanosomal mesohyl. with protruding primary fibres from ascending choanosomal skeleton and plumose bundles of principal subtylostyles erect on surface producing hispid ectosome; sparse tracts of thinner auxihary styles tangential to sur- face. Choanosome. Skeletal architecture more-or-less renieroid reticulate, with heavy spongin fibres producing wide-meshed rectangular reticulation; primary ascending fibres long, multispicular 60- 904m diameter, interconnected by numerous, shorter, secondary fibres, 30-55m diameter cored by 1 or few principal spicules; fibres spar- sely echinated by principal subtylostyles, identi- cal to but marginally thicker than those coring fibres, confined mostly to distal margins (periphery) af fibres; fibre anastomoses produce cavernous meshes, 80-3204um diameter; mesohyl matrix sparse in choanosome, lightly pigmented, granular, containing numerous whispy auxiliary styles and fewer microscleres; chounocyte cham- bers elongate 20-30j.m diameter. Megascleres. Principal subtylostyles coring and sparsely echinating fibres short, moderately thick, straight or slightly curved at centre, entirely smooth, with slightly subtylote bases, blumened or slightly telescoped points, Length 186- (214.4)-238j.m, width 4-(4.6)-7 jum. Auxiliary styles very similar in geometry to principal spicules except for being much thinner, whispy, and lacking subtylote bases); auxiliary styles straight or sinuous, rounded bases, pointed or telescoped points. Length 182-(204.2)- 246m, width 1.5-(1.9)-2.5j m. Microscleres. Palmate isachelae small, un- modified, long lateral alae approximately same length as front ala, entirely fused to shaft, front alae nearly completely detached. Length 10- (11.7-13um. REMARKS, Lendenfeld's (1888) original material is not extant in the collections of either the AM or BMNH, but we know from his bref description that the species has a growth Form reminiscent of. Ciocalypta (Halichondrida) and spiculation of Echinoclathria or Echinochalina. The specimen described here from the Howick Islands group, close to the type locality of Torres Strait, agrees completely with Lendenfeld’s (1888) brief description and is nominated neotype of this species. Echineclatiria digitata is similar to E. berguis- ie in its digitate growth form and having a cav- ernous, predominantly renieroid, reticulate skeletal architecture. It differs from E. bergquis- tae in geometry and dimensions of all its spicules, lacking toxa microscleres, and having relatively homogeneous megascleres throughout the skeleton, conng and echinating tracts and scat- tered interstitially. In possessing relatively homogeneous megascleres E, digitata ulso resembles E. levii, although the latter has corn- pletely different skeletal architecture, growth form and toxa microscleres. Echinoclathria egena Wicdenmayer, 1989 (Figs 237-238, Plate 10E) Echinoclathria egena Wiedenmayer, 198%: 64-66 pl.6, fig.8, pl.24, fig.6, pl.25, figs 1-2, text-fig.44; Hooper & Wiedenmayer, 1994: 279. MATERIAL. HOLOTYPE: NMVF51978: Winter Cove, E. side of Deal 1., Kent Group, Bass Strait, Tas, 39°29'S, 147?20' E, 26.11.1981, 3-6m depth. coll. F. Wiedenmayer et al, (SCUBA).OTHER MATERIAL: TAS- QOMG300669 (NCIQ66C-3721-L) (fragment NTMZ23817). 454 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 234. Echinoclathria confragosa (Hallmann) (holotype AMZ992). A, Choanosomal skeleton. B, Fibre charac- teristics (x288). C, Choanosomal principal subtylostyle (coring and echinating fibres). D, Ends of principal subtylostyles. E, Subectosomal auxiliary style. F, Ends of auxiliary spicules. G, Palmate isochela, H, Oxhorn toxa. HABITAT DISTRIBUTION. On granite boulders and | DESCRIPTION. Shape. Erect, digitate, ranging xd = E ire TD) depth; Kent Is, E. St ^ (rom young forms thickly encrusting basal mat, Pm up to 5mm thick, with irregularly bifurcate and REVISION OF MICROCIONIDAE 455 25um FIG. 235. Echinoclathria digitata (Lendenfeld) (neotype QMG304763). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary styles. C, Palmate isochela. D, Section through peripheral skeleton, E, Australian distribution. F, Neotype. occasionally anastomosing lobo-digitate branches, up to 38mm long, 5mm diameter, to arborescent branching, up to 220mm long, 115mm breadth, cylindrical branches up to 17mm diameter, with bulbous terminal and subterminal processes along branch length, and with short stalk, up to 50mm long, 21mm diameter, and enlarged basal attachment. 456 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 236. Echinoclathria digitata (Lendenfeld) (neotype QMG304763). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Bases and points of auxiliary styles. E, Palmate isochela. Colour. Dull orange-brown alive (Munsell 5 YR7-8/12), beige-brown in ethanol. Oscules. Large oscules, up to 2mm diameter, mainly on lateral sides of branches, particularly on edges of lobate bulbs along branches, less common on basal mat; oscules usually raised on small conical projections with slight membraneous lip. Texture and surface characteristics. Compres- sible rubbery texture, difficult to tear; surface contorted, macroscopically even but microscopi- cally reticulate. REVISION OF MICROCIONIDAE 200 um 25um 457 FIG. 237. Echinoclathria egena Wiedenmayer (holotype NMV51978). A, Principal styles/ subtylostyles (coring and quasi-echinating fibres). B, Subectosomal auxiliary anisostyles/ quasi-strongyles. C, Section through peripheral skeleton. D, Australian distribution. E, Holotype. Ectosome and subectosome. Fibrous, micropapil- lose ectosome, with well developed surface fibres lying paratangential to surface, and with sparse subectosomal auxiliary megascleres lying on or just below surface, orientated tangential or paratangential to it; peripheral fibres swollen, bud-like, containing sparse core of plumose brushes of choanosomal principal styles, which rarely protrude beyond surface; subectosomal auxiliary megascleres not found directly in peripheral skeleton but some way below it. Choanosome. Skeletal architecture regular, dendro-reticulate, slightly renieroid, with heavy spongin fibres divided into primary ascending 458 FIG. 238. Echinoclathria egena Wiedenmayer (QMG300669). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal spicules. E, Base of subectosomal auxiliary stongylote style. elements, cored by uni- or multispicular tracts of choanosomal principal subtylostyles, fibres be- coming swollen and compressed towards periphery (27-52um diameter), and secondary transverse, usually aspiculose fibres (16-401.m diameter); true echinating spicules absent, but choanosomal principal styles occasionally protrude through spongin fibres at oblique angles; fibre anastomoses form more rectangular cavernous meshes deeper within choanosome (270-420j.m diameter), whereas meshes towards MEMOIRS OF THE QUEENSLAND MUSEUM peripheral skeleton become more. compacted and oval (70-904.m diameter); fibres commonly stratified near periphery, with granular spongin; subectosomal auxiliary megascleres only sparsely dispersed within mesohyl; peripheral fibres also contain abundant microalgae. Megascleres. Choanosomal principal subtylostyles and styles (coring and quasi-echinat- ing fibres) small, thin, straight, with smooth, tapering, slightly subtylote or rounded bases, fusiform or slightly telescoped points. Length 38-(47.7)- 53pm; width 2.3-(2,5)- 3.1 pm. Subectosomal auxiliary megascleres thin, straight, or rarely sinuous, stron- gylote styles (asymmetri- cal, anisostyles), with evenly rounded points and slightly thicker bases. Length 88-(158.2)-1784.m, width 1-(1.5)-2.4,.m. Microscleres. Absent. Larvae. Viviparous parenchymella larvae, 150- 195j.m diameter, in various stages of development througout mesohyl. REMARKS. This species is placed with Burton's (19592) group of flabel- late, massive, ramose Echinoclathria sponges lacking microscleres (axinelloides, chalinoides, inornata, leporina, nodosa, none of which are synonyms contrary to Burton’s (1959a: 247) opinion). It differs from these species in having much smaller, almost vestigial choanosomal styles and lacking true echinating spicules. It is close to E. confragosa in growth form, and E, nodosa in skeletal architecture and fibre charac- teristics, but differs from these species in spicule geometry and compression of peripheral spongin fibres. It should also be compared with E. REVISION OF MICROCIONIDAE leporina which has heavily spiculose fibres and ectosomal skeleton, whereas spiculation of E. egena is very much reduced, virtually vestigial. Echinoclathria inornata (Hallmann, 1912) (Figs 239-240) Ophliaspongia inernata Hallmann,1912: 265-268, pl.36, fig.2, text-fig.57; de Laubenfels. 1954: 163; Coombe etal.. 1987: 38]; Chong et al.. 1987: 85. MATERIAL. HOLOTYPE: AM "cf. E826" (in same specimen jar as O. tenuis): 24km S.of St. Francis Is, Nuyts Archipelago, SA, 32°31'S, 133^18'E, 60m depth. coll, FIV ‘Endeavour’ (dredge), OTHER MATERIAL: WA- WAM622-81(1) (fragment NTMZ1714). S AUST- SAMTS4055 (fragment NTMZ1656), AME938, AME768. HABITAT DISTRIBUTION, Rock reef; 3i-60m depth; Rottnest I. (WA); Nuyts Archipelago and Port Noarlunga (SA) (Fig. 239E). DESCRIPTION. Shape. Erect arborescent digitate or club-shaped sponges. 55-270mm high, 25-65mm maximum width, with irregular cylindrical or lohate branches, bifurcate, ex- panded and bulbous at their ends, 7-25 mm diameter; long cylindrical basal stalk, 12-35mrm long, up ta 15mm diameter, enlarged basal attach- ment. Colour. Live colouration unknown, pale brown or yellowish grey in ethanol. Oscules, Small, 1-21nm diameter, only seen on upper portions of digits, Texture and surface characteristics. Firm, com- pressible, difficult to tear; surface membraneous in places, porous in poorly preserved material, uneven, lumpy towards extremities of branches. Ectosome and subectosome. Surface membraneous, microscopically hispid, with longer choanosomal principal styles protruding through surface, individually or in multispicular brushes, arising from ends of ascending primary Spicule tracts within choanosome; subectosomal auxiliary subtylostyles form tangential or paratangential bundles lying just below ec- tosome, surrounding bases of protruding prin- cipal spicules; choanosomal fibres immediately subectosomal: mesohyl matrix in peripheral skeleton heavy, granular. Choanosome. Skeleton irregularly renieroid reticulate, with slightly compressed axis and plumo-reticulate extra-axial regions; spongin fibres in axial region relatively homogeneous, thick, 60-90j.m diameter, bulbous, not clearly divided into primary or secondary elements; axial fibres contain only 1-2 smaller principal styles 459 per tract, producing nearly regular renieroid skeleton, whereas fibres running longitudinally through branches {seen in cross-section in skele- tal preparations) are multispicular, partially obscuring renieroid appearance of axial skeleton; axial fibre anastomoses form tight oval meshes, 40-90,.m diameter, and echinating acanthostyles sparsely dispersed; extra-axial skeleton with more poorly developed spongin fibres, clearly divided into primary and secondary elements; primary fibres ascending, 30-50jum diameter, bifurcating and anastomosing, cored hy 1-3 smaller principal styles and very heavily echinated by the same spicules, particularly near surface: secondarv connecting, transverse fibres 20-40m diameter, 1-2 spicules per tract, oc- casionally aspicular, also heavily echinated; extra-axial fibre meshes not bulbous as in axial region, forming large oval cavernous meshes up to 250j.m diameter; ascending primary fibres also contain tracts of long, sinuous subectosomal auxiliary subtylostyles secondarily incorporated into fibres; spongin fibres closer to surface much thinner than axial fibres bui very heavily cchinated, eventually producing plumose hundles af larger principal styles protruding through surface; mesohyl matrix heavy, granular, with few extra-fibre spicules; choanocytes large, oval, 55-90um diameter. Meeaseleres. Choanosomal principal styles (coring and echinating fibres) long or short, rela- tively thick, slightly curved towards basal end, rounded or faintly subtylote bases, predominant- ly smooth, rarely slightly micróspined bases, fusiform points. Length 88-(124.2)-199,.m, width 3-16.8)-10um. Subectosomal auxiliary subtylostyles long, slender. straight, slightly curved or sinuous, prominently subtyolote, smooth or microspined bases, hastate points. Length 170-(205.6)- 2354m, width 0.5-(1.9)-33.m. Microscleres. Raphidiform toxas uncommon, found in only 2 of 5 specimens (presumably as- sociated with larvae; impossible 10 retain intact on SEM stubs); very long. hair-like, with slight angular central curvature, siraight arms, straight points. Length 205-(225.4)-238.m, width up to (. 5i. m. Associations. Three of the five known specimens of this species are covered with a zoanthid, al- legedly a cornulanid (Hallmann, 1912). REMARKS. Hallmann (1912) erected this species muinly by comparison with E. leporina (as O. tenuis), remarking on their close 460 MEMOIRS OF THE QUEENSLAND MUSEUM Sd FIG. 239. Echinoclathria inornata (Hallmann) (holotype AM"cf.E826"). A, Principal styles/ subtylostyles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Raphidiform toxa. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. G, SAMTS4055. H, WAM622-81(1). similarities. Hooper & Wiedenmayer (1994) used Hallmann's observations to suggest their synonymy, whereas the types and other material indicate that the Sw and SE populations are con- sistently different and are distinct species. The skeleton of E. inornata is dominated by echinating principal spicules, particularly on peripheral fibres, moreso than any other species. These echinating spicules tend to obscure the characteristic unispicular ascending and REVISION OF MICROCIONIDAE 461 Ac FIG. 240. Echinoclathria inornata (Hallmann) (holotype AM"cf.E826"). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal styles. D, Ends of subectosomal auxiliary subtylostyles, transverse tracts, whereas in E. leporina tracts are consistently pauci- or multispicular and echinat- ing spicules are sparse. Echinoclathria inornata also has a thick palmate-digitate or lobate- digitate growth form (whereas E. leporina is thin- ly flabellate), principal megascleres are generally larger, and some specimens of E. inornata have thin raphidiform toxas (although these are not consistently present and might be associated wilh larvac}. Echinoclathria leporina (Lamarck, 1814) (Figs 241-242) agia leporina Lamarck, 1814; 444, 373. chinocknhela leporina, Topsent, 1932: 101, pLå, figl; de Laubenfels, 1936a: 119; de Laubenlels, 1954; 163; Wiedenmayer, 1989: 61-63, pi.6, fig.1, pl.24, figs 4-5, text-fig.42; Carpay. 1986: 24; Hooper & Wiedenmayer, 1994- 2 X PORES leporing, Burton, [9343: $58, 562, bL die tenuis Carter, 1885f: 355. Ophlitaspongia tenuis, Dendy, 1896; 37; Hallmann, 1912: 261-265, pl.35, fgl, text-fig.56; Topsent, 1932: 101, pl.6, fig. 1. Burton, 19342; 558, 562, 599, Not Clathria tenuis Hentschel, 1911; 377-379, text- fig.49; Parish, Jakobsen, Coombe & Bacic, 1991: 56-64. Phakellia papyracea Carter. 1886g: 379. Antherochalina tenuispina Lendenfeld, 1887b: 789; Hallmann, 1912: 265; Burton, 1934a: 558. MATERIAL. HOLOTYPE: MNHNDTS67: ‘Australian Seas’, Peron & Lesucur collection. HOLOTYPE of E tenuis: BMNHI886,12,15,147: Port Phillip Heads, Vic, 40m depth, 38?17'S, 144?39' E, coll. J.B. Wilson (dredge), LECTOTYPE of P. papyracea: BMNH1886.12.15.231 (dry) (fragment AMG2907: same locality. PARALECTOTYPE of F. porrecen: BMNH1886,12.15.232 (dry): same ocality. HOLOTYPE of A. Lenuispina: BMNH1886.8.27,448 (dry) (fragment AMG2467]: Westernport Bay, Vic, 38°26'S, 145*08'E. OTHER MATERIAL: S AUST- AMESs26. VIC- NMVRN1075, AMZ318, AMZII67. NSW- AMZ1642, AME820. TAS- AMZ2136, AMZ22]D. HABITAT DISTRIBUTION. 5-42m depth in shallow coastal waters on rock recf substrate; known only from Australia: Coogee (NSW); Port Phillip, Westernport Bey vie) N. coast (Tas); Cape Martin (SA) (Fig. 41D). DESCRIPTION. Shape. Persistently very thin, flabellate digits, up to 330mm high, 190mm wide, 2-8mm thick, ranging from single elongate planar fans with evenly rounded margins, to hifurcate palmate digits growing in more than 1 plane, with uneven margins; usually with long or short cylindrical basal stalk, up to 65mm long, 18mm diameter. Colour. Pale red or red-orange alive (Munsell LSR 5/8-5/10), light brown in ethanol, Oscules. Relatively small, up to 2.5mm diameter, dispersed over margins of digits, without. as- sociated subectosomal drainage canals. MEMOIRS OF THE QUEENSLAND MLISEUM Texture and surface characterisites. Firm, flexible. moderately difficult to tear: surface even, without pronounced sculpturimg; some with distinct radial growth lines on lamellae. Ectosome and subectosome. Microscapically hispid, with larger sizes of principal stvle/suh- tylostyle protruding through surface for up to 100um, singly or in brushes, forming a vestigial plumose or radial extra-axtal skeleton in peripheral region, and with a distinct tangential layer of subectosomal auxiliary subtylostyles, in pauci- or multispicular tracts, underlying erect principal spicule brushes; mesohyl matrix moderately heavy, particularly near outer margin of peripheral skeleton. Cheoanosome, Skeletal architecture with 3 com- ponents differentiated: (1) irregularly isodictyal, slightly compressed axis; (2) more open-reticu- late renieroid or subrenieroid extra-axial region; and (vestigial) plumose or radial skeleton in the peripheral region; axial skeleton with single, thickened central core of heavy fibres, vaguely separated into primary ascending pauci- or mul- Hispicular fibres, 28-65 1m diameter, and secon- dary, mosily transverse. uni- or paucispicular fibres, 28-36um diameter; primary fibres plumose, arborescent, producing radial tracts; secondary fibres regularly renieroid; fibres cored and sparsely echinated by smaller choanosomal principal styles/subtylostyles in choanosomal skeleton (larger in peripheral skeleton); echinat- ing principal spicules located predominantly on pnmary fibres; fibre anastomoses in axis form predominantly triangular meshes (=isodictyal), less often rectangular or oval meshes, 75-1801.m diameter, obviously more compressed in axis than at periphery; peripheral fibres form more regular, rectangular meshes (=renieroid); primary spicule tracts mostly confined to within fibres in axial skeleton, becoming increasingly plumose. protruding through fibres, in peripheral skeleton; mesohyl matrix heavy but only lightly pig- mented; choanocyte chambers ovoid, 43- | 121m diameter, with sparsely dispersed subectosomal auxiliary megascleres particularly in peripheral skeleton. Megascleres. Choanosomal principal styles and subtylostyles long or short (larger in peripheral region than in axis), thick, straight or slightly curved, with smooth, evenly rounded, or slightly tapering subtylote bases, sometimes quasi- oxeote, rarely microspined bases, usually with fusiform points. Length 62-(185.3)-305um, width 4-(9.6)- 141m. REVISION OF MICROCIONIDAE 463 FIG. 241. Echinoclathria leporina (Lamarck) (holotype MNHNDT567). A, Principal styles/ subtylostyles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyles. C, Section through peripheral skeleton. D, Known Australian distribution. E, Holotype. F, Holotype of E. tenuis. G, Holotype of A. tenuispina. Subectosomal auxiliary subtylostyles long, thin, straight, slightly curved, or frequently sinuous, with predominantly smooth, occasional- ly microspined, subtylote bases, hastate points. Length 148-(265.8)-321j.m, width 1.5-(3.1)- 4.2m. Microscleres. Absent. REMARKS. This species is better known under the junior names Echinoclathria or Ophlitaspon- gia tenuis. In growth form it is remarkably close to A. (Isopenectya) chartacea. This similarity, also noted by Hallmann (1912), is emphasised by their respective renieroid and isodictyal skeletal architecture, compression of the axial skeleton and fibre characteristics, whereas other charac- ters indicate that similarities may be convergent and functionally related to flabellate growth form. Major features which differentiate the two species are the possession of 3 distinct skeletal structures (renieroid, plumose-isodictyal and peripheral skeletons) in A. (Z) chartacea, the 464 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 242. Echinoclathria leporina (Lamarck) (holotype MNHNDTS567). A, Choanosomal skeleton. B, Fibre characteristics. C, Larger principal style (protruding through surface). D, Smaller principal styles (in renieroid skeleton). E, Ends of subectosomal auxiliary subtylostyle. presence of acanthose choanosomal styles form- ing the renieroid skeleton, and much larger smooth choanosomal styles protruding through the ectosome. Choanosomal styles in E. leporina are homogeneous, entirely smooth shafts, and the renieroid/ isodictyal skeleton is cored and echinated by the same smooth principal spicules, with differentiated primary and secondary fibre REVISION OF MICROCIONIDAE structures, there is less pronounced axial com- pression, larger, entirely smooth principal spicules are embedded only in the peripheral skeleton, barely extending into the choanosome, and there is no clear differentiation between renieroid/ isodictyal and plumose skeletons as in Anrho. The two species may be confused and hence care has been taken in checking all avail- able voucher specimens 10 ascertain con- specificity. Echinoclathria leporina is probably wide- spread throughout temperate SE Australia al- though some published records (in the Zoological Record) are nol corroborated by voucher samples and are not included in the present synonymy. Echinoclathria leyii sp. nov. (Figs 243-244, Plate 10F) MATERIAL. HOLOTY PE: QMG300675(NCIQ66C- 3764-1) (fragment NTMZ3832): Trap Reef, Bicheno, Tas., 41*51.7' S, 148°18.6'E, 30m depth, 26,11, 1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION, Large houlder reef; 30m depth; E. coast (Tas) (Fig. 243F). DESCRIPTION. Shape. Erect, thickly flabellate fan in several planes. 155mm long. 135mm wide, each lamella 7-12mm thick, up to 80mm wide, margins irregularly digitate or palmate-digitate, with digits up to 30mm long, Smm diameter; irregular digits and small lamellae (*buds') also ansing from lateral sides of fanz stalk long, thick- ly cylindrical, 45mm long. 9mm diameter, wath expanded hasal attachment. Colour. Dark red-orange alive (Munsell SR 5/10), brown 1n ethanol. Oscules. Large, up to 2 5mm diameter, scattered over L side of fan, without any obvious membranecous Tip, collapsing in air. Texture and surface characteristics, Sialk stiff, lamella firm, flexible. slightly compressible; sur- face membraneous. optically hispid, relatively even, slightly lumpy but without any conules or other ornamentation. Ectosome and subectosome, Surface microscopi- cally hispid, with longer principal styles protrud- ing through surface in thick, bushy, erect brushes arising from ascending choanosomal fibres in peripheral skeleton; subectosomal auxiliary sub- tylostyles in sparse tangential brushes on surface; mesohyl matrix heavy but only lightly pigmented in peripheral skeleton. Choanosome, Skeleton with 2 components: ir- regularly renieroid renieroid reticulate and plumo-reticulate, without any marked axial com- 465 pression but clearly differentiated axial and extra- axial regions; (1) axial skeleton more-or-less renieroid, with heavy spongin fibres divided inu» primary and secondary elements; primary fibres ascending, 70-901um diameter, cored by pauci- or multispicular tracts of generally smaller choanosomal principal styles; secondary fibres transverse, short, thinner 30-40p.m diameter, in- terconnecting primary fibres, cored by unispicular tracts of smaller principal styles; fibre anastomoses in axial region produce cavernous oval or elongate meshes, 150-250,.m diameter, fibre nodes heavy, slightly bulbous, containing multispicular tract ascending through lon- gitudinal plane of lamellae; axial fibres sparsely echinated by smaller principal styles, mainly at fibre nodes; (2) extra-axial skeleton distinctly plume-reticulate, with disctinctly different primary and secondary spongin fibre systems; primary ascending fibres very thick, upto 130m diameter. cored by multispicular tracts of smaller and larger choanosomal principal subtylostyles ascending to surface: spicule tracts become heavier, more plumose towards periphery; size of principal subtylostyles coring fibres generally in- crease towards surface; secondary, connecting fibres in extra-axial skeleton small, less than Qum diameter, uni- or aspicular; echinating spicules in extra-axial region mostly obscured hy ascending plumose spicule tracts; mesohyl matrix heavy, with numerous auxiliary megaseleres (sometimes also incorporated inta fibres) and numerous toxas scattered throught; choanocyte chambers small, oval, 40-60jum diameter. Megaseleres. Choanosomal principal styles (conng and echinating fibres) long or short, moderately slender, slightly curved at centre, with smooth, rounded or slightly subtylote bases, fusiform points, Length 172-(244.4)- 3d l jum, width 5-(6.1)-7 pm. Subectosomal auxiliary styles relatively short, slender, slightly curved at centre, with slightly subrylote, smooth bases, fusiform or hasiate- rounded points. Length 154-(187.8)-2054.m, width 2-(3.1)-4.5.m, Microscleres. Palmate isochelae small, with lateral alae longer than front ala, completely fused to shaft, front ala wide, curved, nearly completely detached from lateral alae; shaft straight. Length 13-(14.5)-164um. Toxas oxhorn (although smaller wing-shaped forms also present), thick, with prominent, wide, even central curvature, shghtly reflexed arms, 466 MEMOIRS OF THE QUEENSLAND MUSEUM .FIG. 243. Echinoclathria levii sp.nov. (holotype QMG300675). A, Principal subtylostyles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Oxhorn toxas. D, Palmate isochela. E, Section through peripheral skeleton. F, Known Australian distribution. G, Holotype. slightly reflexed points. Length 23-(91)-135jm, ETYMOLOGY. For Professor Claude Lévi, MNHN A Paris, for his contributions to Indo-Pacific sponges. width 1-(1.4)-3j.m. REVISION OF MICROCIONIDAE 467 FIG. 244. Echinoclathria levii sp.nov. (holotype QMG300675). A, Choanosomal skeleton. B, Fibre charac- teristics. C-D, Ends of principal and auxiliary spicules. E, Oxhorn toxas. F, Palmate isochela. REMARKS. This species is an unusual Echinoclathria having well developed extra- axial plumo-reticulate skeleton in which the thick plumose spicule tracts completely dominate the thin unispicular connecting fibres, partially obscuring the renieroid skeleton (seen only clear- ly at the core of the skeleton). Although the thick- ness of the plumo-reticulate and renieroid portions of the skeleton varies from place to place within the sponge (the former dominant towards the edge of Jamellae or surface digits (growing edges) and the latier predominant towards the basal stalk region), it is à prominent feature of skeletal structure. This species also differs from related flabellate species such as E. ridddlei sp. nov. and £, leporina in having distinctive oxhorn toxas (cf. no toxas), megasclere dimensions (cf. substantially smaller, thinner megascleres), and a thickly flabellate, palmate-digitate growth form (cf. thinner flabellate lamellac). Echinoclathria nodosa Carter, 1335 (Figs 245-246) Echinoclatheia nodosa Carter, 1885f: 356; Ridley & Dendy, 1887: 160; Carpay, 1986; 25; Hooper & Wiedenmayer, 1994: 280. Ophlitaspongia hodesa; Dendy, 1896: 37. Litaspongia nodosa, de Laubenfels, 1954: 162. MATERIAL. HOLOTYPE: BMNH 1886.12.15.96 (fragment AMG2770} Port Phillip, Vic, 38°09"S, 144*52'E, 8m depth, coll, unknown (dredze). OTHER MATERIAL: VIC- NMVRN264, NM VRN628. HABITAT DISTRIBUTION. Sand and shell gril, 8- 38m depth; Pon Phillip (Vic) (Fig. 245E). DESCRIPTION. Shape. Bulbous-digitate, up to 55mm long, 30mm maximum width, 35mm breadth, small cylindrical basal stalk 5-15mm long, 10mm diameter; with bifurcate and oc- casionally anastomesing, irregularly cylindrical, slightly swollen branches, upto 22mm Jong, 8mm diameter. Colpur. Bright red to crimson alive, dark brown in ethanol. Oscules. Numerous small oscules, up to 2mm diameter, scattered between surface conules, Texture and surface characteristics. Soft, com- pressible, difficult t» tear; surface nodulose, granular. Ectosome and subectosome. Membrancous, with protruding spongin fibres from ascending peripheral skeleton, and points of principal styles protruding beyond surface in sparse plumose brushes or individually, not forming continuous MEMOIRS OF THE QUEENSLAND MUSEUM palisade: subectosomal auxiliary styles form sparse paratangential, tangential or occasionally erect layers below protruding skeleton of prin- cipal spicules. Choanosome. Skeleton dendro-reticulate, more- or-less renieroid and homogeneous throughout, without any axial compression and only slight differentiaüon between peripheral skeleton and core (the former with plumose primary mars becoming increasingly dense towards periphery, the latter evenly renieroid with relatively sparse spicule skeleton); heavy spongin fibres dominate skeleton, clearly differentiated into primary and secondary components; primary fibres, 35-55 um diameter, ascending, multispicular with 3-8 spicules per fibre, spicules confined to within fibres at core but become plumose in subec- tosomal and peripheral regions; secondary fibres uni- or paucispicular. 15-40um diameter, predominantly transverse; fibre nodes usually bulbous, up to 804m diameter; fibres cored by choanosomal principal styles and sparsely echinated by same spicules; echinating spicules confined mainly to core of skeleton, less common in periphery; fibre meshes triangular, rectangular or rhomboidal, 150-270um diameter, slightly more cavernous in subectosomal region than al core; mesohyl matnx light, unpigmented, with few interstitial spicules; choanocyte chambers small, oval, 15-25,cm diameter. Megascleres. Choanosomal principal styles short, thick, straight or very slightly curved at centre, smooth, evenly rounded bases. oc- casionally slightly subtylote, fusiform points; sometimes modified to quasi-oxeote spicules. Length 109-(128.6)-164p,m, width 6-(8.4)- i3pm. Subectosomal auxiliary styles long, slender, straight or very slightly curved at centre, smooth rounded bases or very slightly subtylote, long tapering fusiform points. Length 116-(161.1)- 242m, width 2-(3.6}-6um. Microscleres, Absent. REMARKS. Similar to E. thielei and E. notlalis sp. nov, this species lacks noticeable size dif- ferences between principal styles coring choano- somal fibres and those protruding through ectosomce. It has bulbous branching growth form; bulbous fibre nodes in the skeleton are similar to E. levii sp. nov. It is also reminiscent of E. axinel- loides in its spicule skeleton, which becomes increasingly dense and plumose towards the sur- Tace, but this resemblance is superficial, whereas characters such as growth form, bulbous fibres, REVISION OF MICROCIONIDAE 469 FIG. 245. Echinoclathria nodosa Carter (holotype BMNH1886.12.15.96). A, Principal styles (coring and echinating fibres). B, Subectosomal auxiliary styles. C, Abherrant principal spicule. D, Section through peripheral skeleton. E, Known Australian distribution. F, Holotype. G, Specimen NMVRN264. absence of microscleres, and the geometry and Kangaroo I., SA, 35?46.9'S, 137?46.5'E, 6m depth, size of both categories of megascleres together — 31.1.1989, coll. NCI (SCUBA). differentiate it within the genus. HABITAT DISTRIBUTION. Rock reef, sand, mussell beds; 6m depth; Kangaroo I. (SA) (Fig. 247D). Echinoclathria notialis sp. nov. . (Figs 247-248, Plate 11A) DESCRIPTION. Shape. Erect, massive, club- shaped, bulbous-digitate, 75mm high, 85mm MATERIAL. HOLOTYPE: QMG300614(NCIQ66C- diameter; digits subcylindrical, bulbous, up to 2243-F) (fragment NTMZ3541): American River, 45mm long, 38mm diameter, rounded margins, 470 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 246. Echinoclathria nodosa Carter (NM VRN264). A, Choanosomal skeleton. B, Fibre characteristics. C- D, Ends of principal and auxiliary spicules. REVISION OF MICROCIONIDAE FIG. 247. Echinoclathria notialis sp.nov. (holotype QMG300614). A, Principal styles (coring and echinating fibres). B, Subectosomal auxiliary style. C, Section through peripheral skeleton. D, Australian distribution. E, Holotype. 471 fused to adjacent digits; attached directly to substrate, no stalk or basal ‘holdfast’. Colour. Pale red- brown alive (Mun- sell 5R 5/6), pale brown in ethanol. Oscules. Large, 3- 6mm diameter, on apex or subapex of digits, raised above surface with prom- inent membraneous lip. Texture and surface characteristics. Firm, compressible, relatively easily torn; surface with prominent surface conules when alive (mostly white and offset from darker ectosome), each conule bearing fine trace of radiating subectosomal auxiliary spicules which collapse upon exposure to air and preservation; in air surface membrane collapses, surface becomes highly porous (each pore up to 1mm diameter, scattered evenly over surface), super- ficially resembling a red-brown Acropora coral. Ectosome and sub- ectosome. Fibrous, with tips of ascend- ing primary fibres protruding a long way through surface, with bundles of cho- anosomal spicules also protruding slightly from ends of 472 MEMOIRS OF THE QUEENSLAND MUSEUM l : / iR jd - A] 5 d "c D Fs IE pts AM ct p^ é m m) A Tse FIG. 248. Echinoclathria notialis sp.nov. (holotype QMG300614). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal styles. D, Ends of principal styles. E, Subectosomal auxiliary styles. F, Ends of auxiliary styles. primary fibres; moderately heavy tracts of subec- surrounding surface pores and collapsing upon tosomal auxiliary styles lying tangential, paratan- exposure to air and preservation. gential or erect on surface; auxiliary spicules Choanosome. Skeleton regularly renieroid heaviest between protruding fibres, presumably reticulate, relatively homogenous skeletal tracts, REVISION OF MICROCIONIDAE without any differentiation between axial and extra-axial regions; spongin fibres moderately well developed, 15-25 jum diameter: fibre meshes cavernous throughout, rectangular, square or oc- casionally triangular, 125-250j.m diameter, few obvious differences between ascending and transverse fibres; ascending fibres cored by 1 or few choanosomal principal styles, occasionally more near surface of sponge, with tracts becom- ing multispicular, plumose in ectosomal] and protruding fibres; transverse fibres with similar spicule content; all fibres sparsely echinated by smooth principal styles, identical to those inside fibres; echinating spicules predominant at fibre nodes; mesohyl with abundant, mesohyl matrix only lighüy pigmented, subectosomal auxiliary styles scattered throughout; choanacyte cham- bers small, round or oval, 30-50jum diameter. Megascleres. Choanosomal principal styles {coring and echinating fibres) short, slender, straight, smooth rounded bases, fusiform or slightly telescoped points. Length 107-[123.6)- 151 p.m, width 1.5-(2.7)-44.m. Subectosomal auxiliary styles long, slender, straight, rounded smooth bases, fusiform or prominently telescoped points. Length 123- (163.2)-191 um, width 1-(1.9)-2.5pum. Microscleres. Absent ETYMOLOGY. Greek noros, south. REMARKS, In growth form, texture, coloura- tion, surface sculpturing and conules E. notialis is quite different from all other Echinoclathria, yet tts skeletal structure, spicule geometry and distribution of spicules are more difficult to dif- ferentiate from other species, Skeletal differences are a matter of degree rather than absolute (which seems to be a feature of the genus), including regularity of the renieroid skeletal reticulation (no axial compression; cf. E, confragosa), homogeneity of fibre meshes (without any marked differences between ascending and transverse tracts). and virtually. unispicular spicule tracts throughout the skeleton (except at the surface where they become multispicular and slightly plumose; cf. E. inornata}. The new species differs from all these others in most other respects. In fibre development, density of spicule tracts coring fibres, and possession of both fibres and terminal principal spicule brushes protruding through the surface this species is most similar to E. subhispida, differing substantially in spicule geometry, spicule size, skeletal architecture and field characteristics (see E subhispida below). 473 It is not known if the live surface ornament (small white conules covered by a fine trace of radiating auxiliary spicules, overlaying surface pores) Is unique to the genus as many named species are only known from preserved material. Echinoclathria parkeri sp. nov. (Figs 249-250) MATERIAL. HOLOTYPE: SAMTS4091 (fragments NTMZ16¢H1, QMG300473): 20km due W. of Outer Harbour, Adelaide, St. Vincent Gulf, SA, 34°45°S, 138°20°E, 23-25m depth, 23.1v.1975, coll. J. Me- Phalain. PARATYPE; SAMTS4097 (fragments NTMZ1649, QMG300128): same locality. HABITAT DISTRIBUTION, Gravel, rock reef; 23- 25m depth; St. Vincent Gulf (SA) (Fig. 249E). DESCRIPTION. Shape. Massive, subspherical- bushy or erect digitate growth forms, 75-1 imm high. 45-70mm wide, with short, irregularly cylindrical, bulbous lamellae, up to 50mm long, 20mm diameter, expanded at apex, partially fused to adjacent branches, with small, irregularly cylindrical or subconical digits arising from apex of lamellac and basal mass, up to. 15mm long, 9mm wide, tapering al apex. Colour Colour alive dark or bright red, dark brown in ethanol. Oscules. Numerous small oscules. up to 3mm diameter, apical or subapical on surface digits Texture and surface characteristics. Firm, com pressible, not easily torn, harsh due to incorporá- tion of detritus and bivalve shells into lamellae, surface rugose, porous on preserved state, with irregularly dispersed conules. Ectosome and subectosome. Membraneous, fibrous, with terminal fibres protruding slightly and larger principal styles protruding a long way through surface, in plumose brushes or singly; subectosomal auxiliary subtylostyles relatively dense, usually erect, forming plumose or paratan- gential tracts on surface: mesohyl in ectosomal region heavy, granular, darkly pigmented, Choanosome. Skeleton. irregularly renieroid reticulate, slightly compressed in axis, slightly plumose at surface: spongin fibres very heavy, regularly anastomosing, incompletely divided into primary and secondary fibre systems; axial skeleton with very heavy primary fibres, 90- 120m diameter, running longitudinally (through longitudinal sections of surface digits), ascending to apex of digits, interconnected by heavy but shorter, thinner secondary fibres, 42- 954m diameter, with slightly bulbous nodes at their junction, 155-195j2m diameter; fibre anas- 474 tomoses in axial skeleton close-meshed. 60- 155,um diameter, primary axial fibres cored by multispicular tracts of shorter choanosomal prin- cipal si Be aes secondary fibres unispicular, oc- ly paucispicular; all fibres echinated hy naer principal styles; extra-axial skeleton more cavernous, fibre meshes 110-360um diameter, with more-or-less evenly reticulate fibres and irregularly renieroid reticulate spicule tracts; extra-axtal primary fibres heavy, thick, 65-945 um diameter, clearly ascending and arising perpendicularly from primary fibres in axis, mul- tispicular, with spicule tracts composed of smaller and larger principal styles diverging in- creasingly towards periphery; secondary con- necting fibres heavy, short, mainly transverse, 18-à3p.m diameter. unispicular; echinating spicules moderately common in extra-axis; mesohyl matrix heavy, darkly pigmented, con- tàining many scattered. subectosomal auxiliary styles mostly near surface; choanoc yte chambers oval, 30-50j.m diameter. Megascleres. Choanosomal principal styles (coring and cchinating fibres) long or short, rela- tively thick, slightly curved at centre, with smooth, rounded or slightly subtylote bases, fusiform points. Length 124-(196.6)-3124«m, width 5-(7.1)-10,.m. Subectosomal auxiliary subtylostyles thin (oc- casionally raphidiform), straight, slightly curved or sinuous, smooth subtylote bases, fusiform points. Length 123-4(238.4)-355j.m, width 1.5- (2.9}4 Spm. Microscleres. Chelae absent. Toxas accolada, moderately short, thin, with slightly rounded central curvature, straight arms, straight points. Length 63-+129.4)-L 75m, width 0.5-(1.6)-2.5]um. ETYMOLOGY, For the late Dr Shane Parker of the SA Museum, bryozoologist, sponge enthusiast, or- nithologist and gentleman. REMARKS. This species differs from other Echinoclathria in toxa geometry, growth form, skeletal structure and spicule dimensions. The massive, more-or-less bulbous-digitate growth form is similar to E. notialis sp. nov., although E. parkeri 1s irregularly digitate, lacks the specialised surface sculpturing and porous reticulation peculiar to the prominently bulbous E. notialis. Ks skeletal architecture is also slightly similar to E. subhispida, both having heavy fibres, multispicular primary (ascending) tracts and greatly reduced secondary (transverse) tracts, but those of E. parkeri are consistently MEMOIRS OF THE QUEENSLAND MUSEUM unispicular and renieroid in construction whereas in E. subhispida secondary fibres are aspicular. The two species differ in growth form. fibre thick- ness, spicule geometry and size. Only one other species (E. inornata) has toxas that approach the accolada geometry of E. parkeri. In E. inornata toxas are much longer, raphidiform, with slight angular central curvature whereas in E. parkeri they are short and have rounded central curvature (these two species also differ in most other respects). Echinoclathria parkeri has the heaviest spongin fibres of all species, and its skeletal ar- chitecture is also dominated by the longitudinal, multispicular fibres running through digits, with the renieroid component of the skeleton not as obvious as in most species of Echinoclathria. Echinoclathria riddlei sp. nov. (Figs 251-252, Plate 11 B) MATERIAL. HOLOTYPE: QMG305005 (NCIQ66C- 3637-1) (fragment NTMZ3801): Channel between Chappell I. and Badger 1., Furneaux Is, Bass Strait, 40°16.8'S, 147^54.4' E, 15m depth, 22.11.1990, coll. M. Riddle, NCI (SCUBA). PARATYPE: QOMG2300271 (NCIQ66C-3752-T) (fragment NTMZ3827): Tra Reef, Bicheno, E. coast Tas., 41°51.7'S, 148°) 8.6'E 30m depth, 27.11.1990, ‘coll. NCI. OTHER MATERIAL: TAS- QMG30C664 (NCIOQG66C-3556- H) (fragment NTMZ378 1). HABITAT DISTRIBUTION. Rock reel, Ecklonia kelp and Sargassum beds; 15-30m depth; Furneaux Is, Kent Is, Bass Strait, Bicheno (Tas) (Fig. 251D). DESCRIPTION. Shape. Thinly flabellate, up to 190mm long, 150mm maximum width, with long or short, cylindrical basal stalk, 30-75mm long. up to 18mm diameter, and expanded basal attach- ment; fan very thin walled, 1-4mm thick, flat, growing face-on to current, or convoluted grow- ing in several planes; margins of fan digitate. palmate-digitate or heavily convoluted, never even. Colour. Red or orange-hrown alive (Munsell 2.5R S/10 - 5YR 7/10), pale brown in ethanol. Oscules. Minute, 0.5mm diameter, scattered over interior face of convoluted fan (holotype), or on osculiferous face of flat fans. Texture and surface characteristics. Soft, slightly compressible, flexible; surface smooth, perfectly even, without any sculptunng, or with only faint ridges on surface near margins of convoluted fan; surface porous in preserved state, collagenous alive. Ectosome and subectosome. Ectosome micros- copically hispid with protruding large REVISION OF MICROCIONIDAE 475 25um ne, LA SS Le 92, FIG. 249. Echinoclathria parkeri sp.nov. (holotype SAMTS4091). A, Principal styles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Accolada toxa. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. G, Paratype SAMTS4097. choanosomal principal subtylostyles embedded on outer edge of peripheral skeleton, extending up to 300ju.m from surface, surrounded by ir- regular bundles of smaller thin subtylostyles lying tangential, paratangential or rarely forming brushes on ectosome; mesohyl matrix light in subectosomal region, ectosome membraneous. Choanosome. Skeletal architecture renieroid reticulate, without any division between axial or peripheral regions except for larger principal 476 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 250. Echinoclathria parkeri sp.nov. (holotype SAMTS4091). A, Choanosomal skeleton. B, Fibre charac- teristics. C-D, Ends of principal and auxiliary styles. E, Accolada toxas. REVISION OF MICROCIONIDAE spicules protruding through surface; spongin fibres light, imperfectly divided into primary and secondary elements; primary fibres ascending, 30-40pm diameter, cored by pauci- or multi- spicular tracts of smaller subectosomal auxiliary subtylostyles; secondary fibres connecting primary elements, mainly transverse, 10-20j.m diameter, cored by uni- or paucispicular tracts of smaller principal spicules; primary and secon- dary fibre anastomoses form rectangular, ir- regular, or occasionally triangular meshes, 40-80j.m diameter; fibres echinated by smaller principal styles, mainly at core of skeleton, whereas in peripheral skeleton primary spicule tracts become plumose with principal spicules protruding through fibres at acute angles; mesohyl matrix light, unpigmented; choanocyte chambers small, oval, 15-45j.m diameter. Megascleres. Choanosomal principal subtylos- tyles long or short (the longer ones confined to the ectosomal skeleton), thick, slightly curved at centre, rarely straight, with smooth or microspined subtylote bases, entirely smooth shafts, fusiform points. Length 84-(146.4)- 348jum, width 4-(6.8)10-j.m. Subectosomal auxiliary styles long, straight, slender, with prominently subtylote bases, smooth or slightly microspined, points that are fusiform, hastate or slightly telescoped. Length 127-(191.9)-282j.m, width 1-(1.9)-3p.m. Microscleres. Absent. ETYMOLOGY. Named for Dr Martin Riddle for col- lecting this and many other undescribed sponge species throughout Australasia during the AIMS NCI contract 1984-1991, and providing the author with many oppor- tunities to collect from remote sites throughout the Indo-west Pacific. REMARKS. This species has a very thinly flabel- late growth form and renieroid skeletal structure similar to Antho (1.) chartacea, with which it was initially confused. However, A. chartacea has 3 distinct skeletal components: the smaller, smooth choanosomal principal styles forming the renieroid skeleton in the compressed axis and ascending to the surface in longitudinal tracts, the larger, smooth principal styles embedded in ter- minal ascending tracts, and the entirely spined styles forming extra-axial renieroid skeletal tract; principal styles are longer and thicker, with even- ly rounded smooth bases, and tracts extend all the way into the choanosomal skeleton; auxiliary spicules are shorter, the skeletal meshes are more cavernous and the skeletal meshes comprising the renieroid skeleton are relatively homogenous. By 477 comparison, E. riddlei sp. nov. has its principal styles with an entirely smooth shaft, bases are smooth or microspined (and spines are mor- phologically quite different from those in A. char- tacea), bases of principal spicules are subtylote, the longer principal styles are embedded only in external surface (i.e., there is no secondary, lon- gitudinal subisodictyal skeleton), skeletal meshes are much closer, fibres are poorly developed and heterogeneous (differentiated into primary and secondary elements), and points of auxiliary spicules do not vary in their terminations as do those of A. chartacea. These two species may be confused in the field, and their convergent field characteristics further illustrates the difficulty in objectively defining a generic boundary between Antho and Echinoclathria. Echinoclathria subhispida Carter, 1885 (Figs 253-254) Echinoclathria subhispida Carter, 1885f: 356 [et var. subhispida]; Ridley & Dendy, 1887: 160; Hooper & Wiedenmayer, 1994: 280. Ophlitaspongia ^ subhispida; Dendy,1896: 36; Hallmann, 1912: 257-260, pl.36, fig.1, text-figs 54- 55. Echinoclathria gracilis Carter, 1885f: 356. Thalassodendron viminalis Lendenfeld, 1888; 225. ? Echinoclathria viminalis; in part, Whitelegge, 1901: 87-88. Not Thalassodendron viminalis, Whitelegge, 1902a: 214-215. Not Echinochalina glabra; Thomas, 1977: 115-116. Ceraochalina multiformis; Whitelegge, 1902b: 287. Ophlitaspongia chalinoides, in part, Hallmann, 1912: 270-272, text-fig.59. Not Axinella chalinoides Carter, 1885f: 358. MATERIAL. LECTOTYPE: BMNH1886.12. 15.70: Port Phillip, Vic, 38?09'S, 144°52’E, other details unknown. PARALECTOTYPE: BMNH1886.12.15.- 73: same locality. LECTOTYPE of E. gracilis: BMNH1886.12.15.45 (fragment AMG2768): Port Phillip Heads, Vic, 38?17'S, 144?39'E, 40m depth, coll. J.B. Wilson (dredge). PARALECTOTYPE of E. gracilis: BMNH1886.12.15.90 (fragment AMG2773): same locality. LECTOTYPE of T. viminalis: AMZ948 (dry): Illawarra region, NSW, 34?32'S, 150?50'E, other details unknown (label 'Distylinia viminalis (Thalassodendron); type’). PARALECTOTYPE of T. viminalis: BMNH1887.4.27.116 (dry) (fragments AMG3589, ZMB7135): same locality. OTHER MATERIAL: VIC- AMZ922, AMZ797, AMZ1141, AMZ1142, AMZ1144, AMZ1763. S AUST- QMG300240 (NCIQ66C-2194-Z) (fragment NTMZ3534). 478 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 251. Echinoclathria riddlei sp.nov. (holotype QMG305005). A, Principal subtylostyles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyles. C, Section through peripheral skeleton. D, Australian distribution. F, Holotype (at right) and unregistered specimen from same locality. HABITAT DISTRIBUTION. Rock reef, sand, seagrass substrata; 38-440m depth; Port Phillip (Vic); Kangaroo I. (SA); Illawarra (NSW) (Fig. 253). DESCRIPTION. Shape. Erect, digitate, up to 220mm high, 90mm maximum width; digits long, whip-like, thinly cylindrical or slightly flat- tened, up to 120mm long, 12mm maximum thick- ness, repeatedly bifurcate, rarely anastomosing, with rounded, flattened, or bifid apical margins, occasionally palmate; short basal stalk, 15-65mm long, 18mm maximum diameter and expanded basal attachment. Colour. Red to dark red alive; brown, dark red- brown to maroon in ethanol. Oscules. Large oscules, up to 4mm diameter, scattered on lateral margins of branches, slightly raised on small conules, slightly stellate or ir- regular in shape. Texture and surface characteristics. Digits com- pressible, rubbery, flexible; surface optically smooth, even except for microscopic conules REVISION OF MICROCIONIDAE 479 FIG. 252. Echinoclathria riddlei sp.nov. (paratype QMG300271). A, Choanosomal skeleton. B, Fibre charac- teristics, C, Principal subtylostyles. D, Subectosomal auxiliary subtylostyles. bearing apical oscules on lateral margins of digits. Ectosome and subectosome. Surface membraneous, microscopically hispid, with points of longer choanosomal principal styles protruding slightly through surface, singly or in paucispicular plumose brushes, arising from ter- minal fibres, and fibres themselves also protrude through surface; few subectosomal auxiliary styles scattered tangential or paratangential to ectosome. Choanosome. Skeleton regularly dendro-reticu- late, composed of heavy spongin fibres more compressed in axial than extra-axial regions; axis irregularly reticulate, heavy fibres incompletely differentiated into primary and secondary ele- ments, 46-182j.m diameter, with markedly bul- bous axial fibre nodes; extra-axial skeleton nearly 480 perfectly regular, dendro-reticulate, with clearly differentiated primary ascending, radial, paucispicular fibres branching at outer surface (30-62j.m diameter), interconnected at regular intervals by thinner transverse, aspicular or rarely unispicular secondary fibres (18-53jum diameter); primary fibres both cored and echinated by choanosomal principal styles, slightly smaller than those at the surface; echinat- ing spicules marginally more abundant on peripheral fibres; fibre anastomoses form oval or elongate meshes in axis (160-295j.m diameter), square, rectangular or oval meshes towards periphery (125-290jm diameter); mesohyl matrix heavy but only lightly pigmented, large ovoid choanocyte chambers 75-110j.m diameter, numerous loose choanosomal styles and fewer subectosomal auxiliary megascleres scattered be- tween fibres. Megascleres. Choanosomal principal styles (coring and echinating fibres) moderately short, thick, slightly curved or straight, with smooth, tapering, constricted and slightly subtylote bases, fusiform points. Length 75-(131.3)-168m, width 4-(5.4)-7jum. Subectosomal auxiliary subtylostyles long, thin, straight or slightly curved at centre, with smooth subtylote bases, fusiform points. Length 89-(183.1)-235jum, width 1-(2.2)-3j.m. Microscleres. lsochelae absent. Toxas wing-shaped to oxeote, thin with slightly rounded or slightly angular central curvature, straight arms, straight or slightly reflexed points. Length 30-(72.5)-95 jum, width 0.8-(1.2)-1.5j.m. Larvae. Viviparous, spherical or elongate parenchymella larvae, 180-280x150-210,.m, heavily pigmented, and with toxas and juvenile styles dispersed. REMARKS. AMZ948 of Whitelegge (1902b) and Hallmann (1912), was labelled ‘Cerao- chalina multiformis var. dura Lendenfeld (no. 332)’, but is now labelled (in Hallmann's hand- writing) ‘Distylinia viminalis Lendenfeld, ?type of Thalassodendron viminalis’. However, the holotype of var. dura (with a surviving fragment AMG3456) is different (cf. Lendenfeld, 1887b). This variety was also described from Port Chal- mers, New Zealand (the only locality cited in the original description), containing oxea megascleres (cf. Whitelegge, 1902b). Thus, AMZ948 is not a syntype contrary to Lendenfeld's (1887b) descriptions. The specimen Thalassodendron viminalis Lendenfeld, no.365 missing from AM (pers. obs.) was rejected as type MEMOIRS OF THE QUEENSLAND MUSEUM by Whitelegge (1902b), who made it the holotype of his E. intermedia. Consequently, T. viminalis is a junior synonym of Echinoclathria subhispida var. viminalis. Further discussion on T. viminalis and E. intermedia are given below under E. inter- media. Some material (AMZ1141, 1142, 1144, 1763) described by Hallmann (1912) as E. chalinoides also belong to E. subhispida. based on different spiculation and skeletal architecture. This species differs from most other Echinoclathria in its renieroid fibre skeleton (i.e., rectangular meshes) rather than dendro-reticulate spicule skeleton, but this is because the transverse, connecting spicule tracts in secondary fibres are either absent or vestigial, leaving only the primary ascending spicule tracts (more-or- less plumose, branching) dominating the mineral skeleton. Size differences between principal styles protruding through the surface and those coring and echinating choanosomal fibres is only marginal (the former slightly larger), whereas in most Echinoclathria this feature is more pronounced. Reductions of these characters in E. subhispida are not good reasons to exclude it from this genus. This species is similar to F. notialis sp. nov. in microscopic surface features (having spongin fibres and sparse spicule brushes protruding through the surface), and in skeleton dominated by spongin fibres (as opposed to well- developed spicule tracts). However, E. notialis has a bulbous-digitate growth form (cf. a digitate, Haliclona-like habit in E. subhispida), lacks toxas (cf. wing-shaped toxas), has substantially smaller spicules, lacks any (or has very little) compression of the axial skeleton (cf. well-dif- ferentiated axial and extra-axial fibres), has vir- tually no difference in size between principal spicules coring fibres and those protruding through the surface (cf. clear localisation of larger spicules in the peripheral skeleton), and both ascending and transverse fibres are cored by greatly reduced, predominantly unispicular tracts (cf. ascending fibres have well developed, multi- spicular spicule tracts whereas transverse fibres are often aspicular in E. subhispida). — OTHER SPECIES OF ECHINOCLATHRIA Echinoclathria arborea (Tanita, 1968) Litaspongia arborea Tanita, 1968: 49-50, pl.1, fig.7, text-fig.9 [Ariake Sea]. Not Echinoclathria arborea; Hallmann, 1912: 202. MATERIAL. HOLOTYPE: MMBS. Japan. Echinoclathria arcifera (Schmidt, 1880) REVISION OF MICROCIONIDAE 481 FIG. 253. Echinoclathria subhispida Carter (B-C,E-F, holotype BMNH1886.12.15.70; A,D,G, holotype of T. viminalis AMZ948). A-B, Principal styles/ subtylostyles (coring and echinating fibres). C, Subectosomal auxiliary subtylostyles. D-E, Wing-shaped/ oxeote toxas. F-G, Sections through peripheral skeleton. H, Australian distribution. I, Holotype. J, Holotype E. gracilis. K, Holotype T. viminalis. 482 Tenacia arcifera Schmidt, 1880: 81, p10 [Gulf of Mexico]; Hallmann, 1920: 771. Ophlitaspongia arcifera, Burton, 1959a: 247 [key to species]. MATERIAL. HOLOTYPE: LMJG, Caribbean. Echinoclathria atlantica Sara, 1978 Echinoclathria atlantica Sarà, 1978: 76-79, text-figs 49-51 [Cape Domingo, Tierra del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 50 [Tierra del Fuego, Argentina]. MATERIAL. HOLOTYPE: MNHNDCL2739. SW. Atlantic. Echinoclathria beringensis (Hentschel, 1929) Phakellia papyracea, in part: Lambe, 1892: 76 [Alas- ka] IZUG34, fragment Phakellia beringensis Hentschel, 1929: 975, Echinoclathria beringensis; de Laubenfels, 1954: 164 [note]. Echinoclathria schmitti de Laubenfels, 1942. 264 [Baffin Bay, Arctic]. MATERIAL. HOLOTYPE: USNM. Arctic. Echinoclathria contexta Sarà, 1978 Echinoclathria contexta Sarà, 1978: 79-82, text-figs 52-53 | Tierra del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 49 [Tierra del Fuego. MATERIAL, HOLOTYPE: IZUG 170, fragment MNHNDCL2749. SW. Atlantic. Echinoclathria dichotoma (Lévi. 1963) Ophlitaspongia dichotoma Lévi, 1963:59, pl.9G, text- fig.68 [South Africa]. MATERIAL. HOLOTYPE: MNHNDCL6IS. South Africa. Echinoclathria digitiformis (Row, 1911) Ophlitaspongia digitiformis Row, 1911: 351-353, pl.37, figs 14-15 [Shab-ul-Shubak, Red Sea]. Echinochalina digitiformis, de Laubenfels, 1936a- 119 [note]. MATERIAL. HOLOTYPE: BMNH1912.2.164. Red Sea. Echinoclathria hjorti Arnesen, 1920 Echinoclathria hjorti Arnesen, 1920: 21-22, pl.2, fig.5, pl.5, fig.3 [off Cape Bojador, Sahara, N. Atlantic]. Axociella hjorti; dc Laubenfels, 1936a: 119 [note]. Ophlitaspongia hjorti; Burton, 1939b: 43-44 [Iceland]. MATERIAL. HOLOTYPE: ZMUU. NW. Africa, NE. Atlan- "je. Echinoclathria minor (Burton, 1959) Ophlitaspongia minor Burton, 19592: 246, text-fig.27 [Arabian Sea; key to species]; Sim & Kim. 1988: 24, pl.1, figs 1-2 [S. Korea, doubtful conspecificity . aah alia HOLOTYPE: BMNH1936.3.4. 609. Arabian wif, Echinoclathria mutans (Sarà, 1978} Ophlitaspongia mutans Sarà, 1978: 73-76, text-ligs 47-48 [Cape San Sebastiano, Cape Domingo, Tierra MEMOIRS OF THE QUEENSLAND MUSEUM del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 50 [Tierra del Fuego, Argentina]. MATERIAL. HOLOTYPE: IZUGI24, fragment- MNHNDCL2750. SW. Atlantic. Echinoclathria noto (Tanita, 1963) Ophlitaspongia noto Tanita, 1963: 124-125, pL4. fig.3, text-fig.3 [Sea of Japan]; Tanita, 1964: 17-18, 21, pl.1, fig.4 [Japan]; Tanita, 1965: 48 [Sea of Japan]. Sim & Kim, 1988: 24-25 [Korea]; Sim & Byeon, 1989: 37 [Korea]. MATERIAL. HOLOTYPE: MMBS. Japan, Korea, Echinoclathria oxeata(Bergquist & Fromont, 1988) Onhlitaspongia oxeata Bergquist & Fromont, 1988: 113, pl.52, figs c-e [New Zealand]; Dawson, 1993: 38 [index to fauna]. MATERIAL, HOLOTYPE: NMNZPOR117. New Zealand. Echinoclathria reticulata(Bergquist & Fromont, 1988) Ophlitaspengia reticulata Bergquist & Fromont, 1988: 113-114, pl.52, fig. f, pl.53, figs a-c [New Zealand]; Dawson, 1993: 38 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPOR118. New Zealand. Echinoclathria translata(Pulitzer-Finali, 1977) Ophlitaspongia translata Pulitzer-Finali, 1977: 63, lext-figs 20-22 [Tyrrhenian Sea]. MATERIAL. HOLOTYPE: BMNH1977,7.69, Mediter- ranean. Echinoclathria waldoschmitti de Laubenfels, 1954 Echinoclathria waldoschmitti de Laubenfels, 1954; 163-164, text-fig. 106 [Kuop Atoll, W Pacific]. MATERIAL. HOLOTYPE: USNM23092, NW Pacific TRANSFERS Other species described in Echinoclathria but now transferred to other genera. Ophlitaspongia fucoides Bowerbank, 1876: 771. pl.80 [Shark Bay, WA]. Terpiosella fucoides; Burton, 1930b: 675. MATERIAL. HOLOTYPE: BMNH1877.5.21,8. Note: referred to Suberitidae. Holopsamma Carter, 1885 Holopsamma Carter, 1885c: 211 Pleciispa Lendenfeld, 1888: 225. Aulena; in part, Lendenfeld, 1888: 228; (not Lenden- feld, 1885c: 309). [Halme] Lendenfeld 1885c: 285 (preocc.). TYPE SPECIES. Holopsamma crassa Carter, 1885c: 211 (by subsequent designation of de Luubenfcls, 1936a: 98). REVISION OF MICROCIONIDAE 483 | ] [ | FIG. 254. Echinoclathria subhispida Carter (holotype BMNH 1886.12.15.70). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal styles/ subtylostyles. D-E, Ends of principal and auxiliary subtylostyles. F, Wing-shaped - oxeote toxas. DEFINITION. Honeycombed reticulate growth skeletons; principal spicules core and echinate form of tightly anastomosing flattened fibre- branches (lacunae); choanosomal skeleton simp- . ly reticulate, without any axial compression or COmpletely replaced by detritus although those differentiation between axial and extra-axial — echinating fibres usually always present. fibres, those inside fibres sometimes partially or 484 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 255. Holopsamma arborea (Lendenfeld) (holotype AMZ946/G9160). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary style and points. C, Section through peripheral skeleton. D, Known Australian distribution. E, Holotype. F, NTMZ2907. REMARKS. Holopsamma and Echinoclathria Halme have been used interchangeably for all have been confused throughout the literature and species. Holopsamma is separated from Echino- Echinoclathria, ^ Ophlitaspongia, — Plectispa, | clathria by its honeycombed reticulate growth REVISION OF MICROCIONIDAE 485 1 S Na k à " i ; ' z : à w | : AK i aS zc 4 AN 1 Lae MASS ky n Lec j m dq ^ v Te. 9. = A as Na d = ie & fe : t T W * & eS T I —c— — R 5 =) >) M" FIG. 256. Holopsamma arborea (Lendenfeld) (QMG303756). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal style. D-F, Ends of principal style. G, Ends of auxiliary style. form, homogeneous fibre reticulation, single size class of choanosomal spicule found both inside fibres and echinating fibres (ie., lacking the larger, extra-axial styles found in the peripheral skeleton of most Echinoclathria): when type species are compared it is surprising why these genera have been confused. Twenty four species have been referred to this genus, of which 10 are well established and live in Australian waters; they are probably part of the Gondwanan fauna. 486 Holopsamma arborea (Lendenfeld, | 888) (Figs 255-256, Plate 11C) Pleciispa arborea Lendenfeld, 1888: 226, Echinoclathria arborea; Hallmann, 1912: 202, 230- 282, pl.29, fig.2, text-fig 63. Hotopsamma arborea; Hooper & Wiedenmayes, 1994; 281, Not Clathria (Plectispa) arborea, Whitelegge, 1901: 65, 88, 89, pl.11, figs 15-15a: Hallmann, 1912: 211 [nomen nudum]. Nat Litaspongia arbaren Tanita. 1968: 49, Nol Thalassodendron reticulaia Lendenfeld, 1888: 227, Whitelegge, 1902a: 212. Echinoclathria elegans, Whitelegge, L901: 90, Not Plectispa elegans Lendenfeld, 1888: 226. Not Clathria mullipes Whilelegge, 1901: 88, ? Echinoclathria laxa, de Laubenfels, 1954: 164, MATERIAL. HOLOTYPE: AMZ946/G9160 (dry): [- lawarra, NSW, 34732'S, 150°50'E, other details un- known (old specimen label “Halme alborea, type or paypas new label ‘Echinoelathria arborea Lendi. ? "LOTHER MATERIAL: NSW- QMG303756, OMG303760, QMG303797, QMG303808, VIC- GMa 1287 (NCIQ66C-3499-T) (fragment NTMZ3772). TAS- QMG311450 (NCIQ66C-3759- A) (fragment NTMZ3826). S AUST- QMG300239 {NCIQ66C-2148-Z) (fragment NTMZ3526). WA- QMG300192 (fragment NTMZ2907), N'TMZ2368. HABITAT DISTRIBUTION. Widely distributed in temperate coastal waters usvally on rocky reefs; shal- low subtidal-50m depth; Mawarra, N. & S. Sydney (NSW); Westernport Bay (Vic), Bicheno (Tas), Kan- garoo L (SA), Houtman Abrolhos, Barrow I. (WA) (Fig. 255D). DESCRIPTION. Shape. Arborescent, cavemous- reticulate, tubulo-digitate sponge, up to 450mm long, with short basal stalk, up to 35mm long, 18mm diameter, and single or multiple, regularly or irregularly cylindrical branches which repeatedly bifurcate and occasionally anas- tumose; branches 120-330mm long, 15-38mm diameter, often tapering towards ends. Colour. White or off-white alive, grey or prey- brown in ethanol, Oscules, Large, up to 7mm diameter, recessed hetween honeycomb surface meshes, surrounded by membraneous lip. Texture and surface characteristics. Firm, com- soap flexible branches, slightly harsh sur- ace; surface regularly or irregularly honeycomb reticulate, with fibre-branches (lacunae) forming close hexagonal meshes, 8-22mm diameter; lacunae ndge-like, projecting up to 10mm above surface, with thin membrane stretched between adjacent lacunae. MEMOIRS OF THE QUEENSLAND MUSEUM Ectosome and subectosome, Ectosomce membraneous, collagenous, with few subec- tosomal auxiliary megascleres lying tangential or paratangential to surface, and single or paucispicular plumose brushes ‘of principal styles protruding through surface; subectosomal skeleton undifferentiated from choanosomul skeleton, immediately below surface Choanosome. Skeletal architecture regularly reticulate, with relatively light, small spongin fibres differentiated into ascending primary, pauci- or multispicular (30-521.m diameter), and transverse hi-, uni- or rarely aspicular connecting elements (12-221.m diameter); fibre anastomoses form oval to elongate meshes, 60-i140j.m diameter, fibres cored and echinated by choanosomal principal styles, and echinating spicules most common on peripheral fibres; mesohy] matrix light, virtually unpigmented, with few scattered subectosomal auxiliary spicules between fibres; choanocyte chambers small, oval, 22-35jum diameter, Megascleres. Choanosomal principal styles or subtylostyles short, thick, with tapering or slight- ly subtylote bases, typically smooth, sometimes vesligially microspined bases, points tapering fusiform. Length 78-(104,8)-lI8um, width 4- (5.6)-7.5. m. Subectosomal auxiliary megascleres thin, straight or slightly curved at centre, rownded smooth bases and telescoped hastate-pninted or rounded strongylote points somelimes resem- bling anisostrongyles. Length 109-(133.2}- 143pm, width 1.5-(2.2)-2. 5j. m. Micrascleres. Absent. REMARKS. There tsonly a single type specimen in the AM collections, despite Hallmann's (1912: 282) remarks to the contrary, and J must assume that the double-registered AMZ946/G9160 is a holotype and not part of a type series (syntype). No other specimens with true affinities to Hf. arborea were found in the AM collections al- though there are several specimens labelled Clathria or Plectispa arborea that Hallmann may have been referring to, but these do not belong here (see remarks for C. mudtipes). This species is distinctive in the field in its nearly pure white colour, thinly cylindrical digitate branches (often bifurcate or arborescent in older specimens), and regular, close-meshed honeycomb reticulation (whereas jn most other species the surface reticulation is generally wider and the lacunae thicker). This species ts closes! to H. ramosa and H. macropora, with comparuble REVISION OF MICROCIONIDAE spicule geometry and skeletal architecture, but differs in fibre diameter, spicule size and field characteristics. Holopsamma crassa Carter, 1885 (Figs 257-258, Plate 11D) Holopsamma crassa Carter, 1885c: 211; Hooper & Wiedenmayer, 1994: 281. Aulena crassa Lendenfeld, 1889a: 101-102, pl.8, figs 1,2, 5, 6, 22, 23, pl.9, figs 5,9. Echinoclathria crassa; Hallmann, 1912: 287. Not Antherochalina crassa Lendenfeld, 1887b: 787. Halme micropora Lendenfeld, 1885c: 303; Lenden- feld, 1889a: 461, pl.9, fig.12, pl.11, fig.4. Halme globosa Lendenfeld, 1885c: 303; Lendenfeld, 18892: 456-457, pl.9, fig.11. Echinoclathria globosa; Carpay, 1986: 24; Wieden- mayer, 1989: 63-64, pl.6, figs 2-5, 7, text-fig.43. MATERIAL. LECTOTYPE: BMNH1886.12.15. 313: Port Phillip, Vic, 38*09'S, 144°52’E, 38m depth, coll. J.B. Wilson (dredge). PARALECTOTYPES: BMNH1886.12.15.314, 316, 410, 485: same locality. LECTOTYPE of H. globosa: BMNH1886.8.27.71 (fragment AMG3753): Port Phillip, Vic, 38?09'S, 144°52’E, 38m depth, coll. J.B. Wilson (dredge) (only remaining syntype of two. LECTOTYPE of H. micropora: BMNH1886.8.27.89 (fragments BMNH1886. 12.15.90, AMG3779): Illawarra, NSW, 34?33'S, 150°55°E, no other details known. PARALECTOTYPE of H. micropora: AMG8828 (dry): same locality. OTHER MATERIAL: QLD- QMG303235. S AUST- QMG301379, SAMTS4082 (fragment NTMZ1678), SAMTS4085 (fragments QMG300472, NTMZ1599), SAMTS4057 (fragment NTMZ1664), QMG310777 (NCIQ66C-2143-U) (fragment NTMZ3524), QMG310804 (NCIQ66C- 2192-X) (fragment NTMZ3532). HABITAT DISTRIBUTION. Rock reef, sand and kelp bed substrata; 2-38m depth; Stradbroke 1., Moreton Bay (SEQ); Byron Bay, Illawarra, (NSW); Port Phillip Bay, Westernport Bay (Vic); Bass Strait (Tas); Inves- tigator Strait, St. Vincent Gulf, Kangaroo I., Port Noar- lunga, Great Australian Bight (SA) (Fig. 257E). DESCRIPTION. Shape. Massive, subcylindri- cal, or lobate-digitate honeycomb reticulate sponge, 75-150mm diameter, with enlarged, flat- tened, slightly elongated basal attachment. Colour. Yellow-brown to cream coloured alive (Munsell 2.5Y 8/6 or paler), brown in ethanol. Oscules. Large, up to 6mm diameter, slightly recessed within reticulate meshes, surrounded by thin membraneous lip in life. Texture and surface characteristics. Texture varies according to amount of detritus in skeleton, ranging from flexible, soft, compressible, to dis- tinctly brittle, arenaceous; surface reticulate, 487 lacunose, consisting of irregularly meandering ridges forming large meshes producing charac- teristic honeycomb growth form, in life covered by moderately thin, translucent dermal membrane stretched between adjacent ridges. Ectosome and subectosome. Ectosome heavily arenaceous, with a thick sandy external cortex covered by a fine membraneous ectosomal skeleton usually containing a tangential layer of auxiliary spicules, in tracts or scattered singly across surface, but easily detached when surface membrane collapses upon preservation (hence and not seen in some specimens); subectosomal region undifferentiated from choanosome, fibres immediately subectosomal. Choanosome. Skeletal architecture irregularly reticulate, with heavy, relatively homogeneous spongin fibres, up to 200pm diameter, lamellated and virtually fully cored by sand grains, with or without a core of choanosomal principal styles; fibres usually always echinated by principal styles, sometimes echinating spicules reduced in heavily arenaceous specimens; mesohyl matrix heavy, darkly pigmented, granular, also contain- ing foreign particles and some auxiliary megascleres scattered between fibres; mesohyl cavernous in places, with cavities up to 260m diameter, and elongate choanocyte chambers 80- 120j.m diameter usually obscured by detritus. Megascleres. Choanosomal principal styles coring and echinating fibres short, thin, straight, with rounded or tapering smooth bases, fusiform, sharply pointed. Length 53-(67.4)-81j.m, width 1.2-(4.1)-61.m. Subectosomal auxiliary spicules long, slender, strongylote styles, straight or slightly curved or sinuous, with slightly subtylote bases, evenly rounded or bifid points. Length 188-(234.1)- 261m, width 1.5-(3.9)-6].m. Microscleres. Palmate isochelae small, straight shaft, long, thick lateral alae fully fused to shaft, nearly completely detached from front ala, front ala shorter than lateral alae. Length 8-(11.6)- 15pm. Toxas absent. Larvae. Viviparous parenchymella larvae spheri- cal or subspherical, 320-410j.m diameter, with vestigial quasidiactinal megascleres scattered throughout mesohyl. REMARKS. Under Echinoclathria | globosa Wiedenmayer (1989) provided a detailed dicus- sion of additional type specimens of each nominal species he examined (whereas only the specimens that have a direct bearing upon the 488 MEMOIRS OF THE QUEENSLAND MUSEUM 500 um Wwe z- = FIG. 257. Holopsamma crassa Carter (lectotype BMNH1886.12.15.313). A, Subectosomal auxiliary subtylos- tyles/ strongylotes. B, Principal style (coring and echinating fibres). C, Palmate isochela. D, Section through peripheral skeleton. E, Known Australian distribution. F, Lectotype. G, SAMTS4085. identity of this species were examined in this study). The polymorphic concept of this species was slightly overstated by Wiedenmayer (1989) and it does have a distinct identity. He also described only auxiliary spicules present in the species whereas all specimens examined have at least some echinating principal styles in the skeleton, even if most of the other spicules have been displaced by sand. In fact, most of the described variability in H. crassa is a conse- REVISION OF MICROCIONIDAE 489 FIG. 258. Holopsamma crassa Carter (0MG303235). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal style. D-E, Ends of principal and auxiliary spicules. F, Base and bifid point of auxiliary spicule. G, Palmate isochelae. quence of the amount of sand incorporated into reflected in the external ‘honeycomb’ reticulate the skeleton, which appears to be directly related appearance of the sponge. With the nomination to the loss of spicules from within fibres and also of a lectotype of H. crassa by Hooper & Wieden- 490 mayer (1994) the senior name of Carter (1885c) now takes precedence over Lendenfeld's (1885c) name H. globosa (which allegedly appeared several months later; Wiedenmayer, 1989). Wiedenmayer (1989) did not take this step due to the ambiguity of Carter's original description. and the lack of a lectotype designation for H. crassa from a composite series of 31 syntypes (consisting of at least five different species belonging to three families). Holopsamma crassa is well characterised by its extemal morphology (superficially resembling u faviid coral), texture (close to Xestospongia ex- iga, Petrosiidae), and skeletal characteristics (superficially resembling the arenaceous ‘keratose” sponges). but variability in these fea- tures can be illustrated by comparing 3 specimens from the St Vincent Gulf region. SAMTS4057 was almost entirely permeated by sand grains, obscuring most of the fibres and mesohyl matrix {with only echinating principal styles as abvious components of the sponge's spiculation). SAMTS4085 had very heavy spongin fibres, heavily arenaceous fibre core, and a heavy, darkly pigmented mesohyl matnx superficially resem- bling a verongid sponge. Both these specimens had fine, sharp ridges forming the external honeycomb reticulation, with a bryozoan lace- like external reticulation in the preserved state. Both these specimens also had very few in- digenous megascleres, and were initially con- fused with a dictyoceratid or verongid species (Fig. 258B). SAMTS4082 contained numerous subectosomal auxiliary megascleres scattered throughout the mesohyl, choanosomal styles echinating lightly arenaceous fibres in moderate quantities, and rounded edges on ridges forming the external reticulation producing a more robust. honeycomb reticulate structure. Holopsamma crassa is most similar to H, laminagfavosa, and it is conceivable that they belong to a single species. They are retained here as distinct species in order to highlight their dif- ferences and not submerge them, although addi- tional non-skeletal characters are needed to confirm or refute the validity of these taxa. Holopsamma crassa has a brownish live coloura- tion; the lacunae forming its extemal honeycomb reticulate ridges are more robust, recessed more deeply, and meshes have only a light membraneous ectosomal covering between them; there is a more prominent sandy cortex; principal spicules are fusiform, sharply pointed; auxiliary strongylote styles have rounded or bifid points; and chelae are present, H. laminaefavosa MEMOIRS OF THE QUEENSLAND MUSEUM js white alive; shallow meshes formed by the surface honeycomb reticulation are connected by a well formed tympanized membrane stretched between adjacent ridges; principal styles have telescoped points; auxiliary spicules have rounded or slightly hastate points; and chelae are absent. Wiedenmayer (1989) also suggested that the growth form of H, crassa was reminiscent of Phoriospongia reticulum Marshall, although the spiculation of the latter is quite different (stron- gyles, subtylotes and sigmas). Holopsamma elegans (Lendenfeld, 1888) (Figs 259-260) Plectispa elegans Lendenfeld,) 888: 226, Echinoclathria elegans; Hallmann, 1912: 203. — Holopsamina elegans; Hooper & Wiedenmayer, 1994: 2 282. Not Echinoclathria elegans, Whitelegge, 1901: 90. Nat Antherochalina elegans Lendenfeld, 1887b: 787. MATERIAL. HOLOTYPE; BMNH1887.4.27.98: Port Jackson, NSW, 33°51°S, 151°16'E, other details unknown. HABITAT DISTRIBUTION. Ecology unknown; Port Jackson (NSW) (Fig. 259D}. DESCRIPTION, Shape. Honeycomb reticulate mass of irregularly cylindrical, thin digitate branches, up to 145mm long, 25mm diameter, with branches composed of tight-meshed fibre bundles up to Smm wide. Colour. Unknown live colouration, grey-brown dry. Oscules. Large, up to 7mm diameter in dry state, within surface lacunae. Texture and surface characteristics. Harsh, brittle in dry state; honeycomb reticulate surfuce, lacunae smooth Ectosome and subectosome. Membraneous, minutely hispid, with abundant subectosomal auxiliary subtylostyles erect on surface in plumose brushes; choanosomal fibres immedi- ately subectosomal. Choanosome. Itregularly reticulate skeletal structure, with differentiated multispicular primary, ascending fibres, up to 200jum diameter, and numerous transverse, pauci- or aspicular secondary fibres, up to 702m diameter; echinat- ing principal styles sparse. scattered only over ascending fibres; spongin fibres heavy; fibre meshes rounded, 30-ISO0pm diameter; choanocyle chambers not seen (poor dry material). REVISION OF MICROCIONIDAE Megascleres. Choanosomal principal styles long, robust, straight or very slightly curved at centre, evenly rounded, entirely smooth bases, fusiform or slightly telescoped points. Length 78-(90.3)- 105pm, width 4-(4.6)-61.m. Subectosomal auxiliary subtylostyles long, slender, slightly curved near basal end, subtylote smooth bases, tapering rounded or quasi-stron- gylote points. Length 90-(141.2)-255jum, width 1.0-(1.6)-2.5p.m. Microscleres. Absent. REMARKS. This species is known only from the poor, dry holotype from Port Jackson. It is pos- sibly a synonym of H. arborea, based on similiarities in their spicule geometries, although there are some differences in their respective skeletal structures. The species also shows similarities to H. crassa, H. macropora, H. ramosa and H. rotunda in having strongylote points on auxiliary spicules, although all species differ in most other respects. Holopsamma favus (Carter, 1885) (Figs 261-262) Echinoclathria favus Carter, 1885e: 292; Ridley & Dendy, 1887: 160-161, pl.31, figs 4-5; Dendy, 1896: 40; Hallmann, 1912: 276-277, text-fig.61; Wieden- mayer, 1989: 59-60, pl.5, fig.10, pl.24, fig.2, text- fig.40; Carpay, 1986: 23. Axociella favus; de Laubenfels, 1954: 164. Holopsamma favus; Hooper & Wiedenmayer, 1994: 282. Not Echinoclathria favus var. arenifera; Carter, 1885f: 350. Spongia cellulosa; Ellis & Solander, in part; Lamarck, 1814: 373; Topsent, 1932: 20, pl.1, fig.3. Not Spongia cellulosa Esper,1797. Echinoclathria carteri Ridley & Dendy, 1886: 476; Ridley & Dendy, 1887: 162, pl.29, fig.12, pl.31, fig.3; Hallmann, 1912: 284-285, text-fig.65. Holopsamma carteri; Hooper & Wiedenmayer, 1994: 281. Axociella carteri; de Laubenfels, 1936a: 119. Echinoclathria macropora; Whitelegge, 1907: 504. Not Echinoclathria macropora Whitelegge, 1901: 89, 117. Not Plectispa macropora Lendenfeld, 1888: 226. MATERIAL. HOLOTYPE: BMNH1887.5. 13.35: Port Phillip, Vic, 38?09'S, 144°52'E, other details unknown. HOLOTYPE of E. carteri BMNH1887.5.2.751: Off Port Jackson, NSW, 33°51’S, 151?16'E, HMS ‘Challenger’ (dredge). PARATYPE of E. carteri: BMNH1887.5.2.44: same details. OTHER MATERIAL: TASMANIA - MNHNDT556 [specimen of S. cellulosa sensu Lamarck]. WESTERN 49] AUSTRALIA - NTMZ3498. HABITAT DISTRIBUTION. Growing on bivalves, seagrass, rock reefs, vertical rock walls of cave, in sand and shell-grit substrata; 3-333m depth; known only from Australia: Port Jackson, Shoalhaven Bight (NSW); Port Phillip Bay, Westernport Bay (Vic); King L, Bass Strait (Tas); Houtman Abrolhos, Monte Bello Is (WA) (Fig. 261E). DESCRIPTION. Shape. Elongate digitate, con- tort cylindrical or lobate-cylindrical honeycomb reticulate sponge, forming masses up to 380mm high, 450mm wide, composed of bifurcate, usual- ly non-anastomosing branches, with individual branches up to 160mm long, 20-45mm diameter; branches formed by close-meshed fibre bundles (lacunae) producing a characteristic honeycomb surface; multiple points of attachment to sub- strate, without basal stalk or expanded point of attachment. Colour. External surface usually red-orange or orange-brown in life (Munsell 7.5 YR 7/10-12), palerorange below surface, deeper water samples honey-brown, in ethanol specimens turn yellow- brown or dark brown. Oscules. Small, no more than 2mm diameter, within meshes of surface reticulation. Texture and surface characteristics. Soft, com- pressible, rubbery, difficult to tear; regular, close- meshed honeycomb reticulate surface; meshes (alveoles) small, rounded or elongate formed by surface lacunae, up to 4mm diameter, more-or- less regular, usually smaller than meshes within deeper regions of sponge. Ectosome and subectosome. Membraneous, with thin transparent ectosomal membrane stretched across adjacent lacunae at surface of sponge; points of principal styles from ascending choanosomal skeleton protrude slightly produc- ing hispid surface, and paucispicular bundles or single auxiliary subtylostyles lying paratangen- tial to surface; peripheral lacunae rounded or bulbous, not flattened. Choanosome. Skeleton regularly or irregularly reticulate, cavernous; spongin fibres well developed, more-or-less differentiated into primary ascending and secondary transverse ele- ments; primary fibres 35-60j.m diameter, cored by pauci- or multispicular, plumose or oblique tracts of principal styles and small quantities of foreign detritus; secondary fibres up to 20pm diameter, unispicular or aspiculose; fibres sparse- ly echinated by principal styles evenly dispersed throughout skeleton; fibre anastomoses form round to polygonal meshes, 20-100j.m diameter; NTMZ3211, NTMZ3226, 492 MEMOIRS OF THE QUEENSLAND MUSEUM 250um 25um FIG. 259. Holopsamma elegans (Lendenfeld) (holotype BMNH1887,4,27.98). A, Principal style (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Section through peripheral skeleton. D, Known Australian distribution. E, Holotype. mesohyl matrx light, with abundant principal and auxiliary megascleres strewn between fibre meshes; auxiliary spicules predominantly in deeper regions of choanosomal mesohyl, less abundant in peripheral skeleton; choanocyte chambers large, elongate-oval, 35- 55m diameter. Megascleres. Principal styles coring and echinat- ing fibres straight or faintly curved at centre, relatively thick, entirely smooth, with rounded, tapering or oc- casionally anisoxeote bases, fusiform points. Length 69-(96.4)-132jum, width 2-(4.8)-7 um. Subectosomal auxiliary subtylostyles long, thin, mostly straight, with slight- ly constricted bases, usual- ]y hastate, slightly rounded or slighty telescoped points, sometimes sub- tylostrongylote with barely differentiated ends. Length 132-(120.7)-174jum, width 1.4-(1.9)-2.1 jum. Microscleres. Palmate isochelae sparse, small, with long lateral alae en- tirely fused to straight shaft, front ala shorter, often with expanded, spoon-shaped alae. Length 8-(11.9)- 15jum. Toxas absent. REMARKS, Holopsamma favus has a red-orange ex- ternal live colouration (most species of Holop- samma are pale or white), a cartilaginous, algae-like, contort lobate-digitate habit (most are globular or evenly cylindical-digitate), palmate isochelae (most lacking chelae), skeletal ar- chitecture consisting of slightly differentiated as- REVISION OF MICROCIONIDAE 493 FIG. 260. Holopsamma elegans (Lendenfeld) (holotype BMNH1887.4.27.98). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal style. D, Ends of principal style. E, Subectosomal auxiliary strongylote styles. F, Ends of auxiliary spicule. 494 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 261. Holopsamma favus (Carter) (holotype BMNH1887.5.13.35). A, Principal style (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle, C, Palmate isochela. D, Section through peripheral skeleton. E, Known Australian distribution. F, Holotype. G, Paratype of E. carteri. cending and transverse fibres in reticulation (most have relatively homogeneous spongin fibres), and dominance of spicules over detritus within fibres (most are much more heavily arenaceous than H. favus). Comparing their respective type specimens it is obvious that H. carteri is à synonym of H. favus, given their identical growth forms, megasclere spicule geometry and spicule size, and dominance of spicule over detritus within fibres (although these similarities were overlooked by several recent authors; Carpay, 1986; Wiedenmayer, 1989; Hooper & Wiedenmayer, 1994). Wiedenmayer (1989) indicated that the pale red-orange live colouration was typical and consistent for this species, but this is only confirmed for shallow water specimens, whereas deeper water material is reportedly paler, honey-brown (e.g., Ridley & REVISION OF MICROCIONIDAE 495 FIG. 262. Holopsamma favus (Carter) (NTMZ3211). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal style. D, Ends of principal style. E, Auxiliary style. F, Ends of auxiliary spicule. G, Palmate isochela. Dendy, 1887; Hallmann, 1912). Similarly, some theless, although none of the material described specimens were reported to shed their palmate isochelae microscleres (Wiedenmayer, 1989), making confident identification difficult. Never- others has yet been examined their descriptions by Dendy, Hallmann, Whitelegge, Carpay and 496 match fairly closely those of the type specimens 1o be confident of conspecificity, This species is a common component of the benthic fauna in Bass Strait (Wiedenmayer, 1989), and the pecimens from the Houtman Abrolhos and Monte Bello Islands indicate a more widespread distribution. Holopsamma laminaefavosa Carter, 1835 (Figs 263-264, Plate 1 1E) Holopsamma laminaefavesa Carter, 1885c; 212; Hooper & Wicóenmayer, 1994; 282. Echinoclathria laminaefavosa; Wiedenmayer, 1989- 60-61, pl.5, fig. 11, vexr-fig.41, Carpay, 1986: 24. Halme nidus-vesparum Lendenfeld, 1885c: 288, pl. 26, figs 1-2, pl.27, fig.4-5,7, pl.28, figs 8-9,11, pl.29, figs 12-13; Lendenfeld, 1888: 157; Lendenfeld, 1889a: 457, pl.1l, figs 1, 7, 9-10, 15-18; Whitelegge, 1889: 184, Halme giganea Lendenfeld, 886a: 847-9 [et varr intermedia, macropora, micropora |. Aulena gigantea, Lendenfeld, 1888; 230-232 [et varr Irermedia, macrcopora]; Lendenfeld, 18899; 97, pl. 8, figs 3,4, 7, 8, 18, 29, pl.9. figs 2-4; Whitelegee, 1889: 187; Whitelegge, 1901: 93, 1 18; Whitelegge. 1907: 504; Hallmann, 1912: 268, 285-287; Burton, 19382; 20; Guiler, 1950: 10, Not Halme micropera Lendenfeld, 1885c: 303; Len- denfeld, 18894: 461. pl.9. fig.12. pl.11, fig.4. Halme laxa Lendenfeld, 18863: 845-7 [et varr digito, minima]. Aulena laxo; Lendenfeld, 1888: 228-250; Lendenfeld, 18892: 95-97, pl.8, figs 10-12. 15, 20-21, pl.9, fie 1; Whitelegge, 1889; 187; Hallmann, 1913: 287; Hallmann, 1914a: 268; Burton. 19383: 20. 7 Aulena laxa; Dragnewitsch, 1906: 442, Echinoclathria laxa; Hallmann, 1912; 287. Halme irregularis Lendenfeld, 1389a: 49, 453-5, pl.9, fig.R, pl.11, fig.3 [et varr micropora, lamellosa) Echinoclathria favus var. arenifera Carter, | &85f- 350. Echineclathria wrenifera; Dendy, 1896: 40:41; Hallmann, 1912: 287; Wiedenmayes, 1989; 60-61. cf. Plectispa urborea Lendenfeld, 1888: 226; de Laubenfels. 1954; 164. MATERIAL. LECTOTYPE: BMNHI1886,12.15.312 (dry) (fragments BMNH1886.12.15.311, AMG2879)- Port Phillip, Vic, 38*09'S, 144752" E, colL J.B. Wilson (dredge). PARALECTOTY PES: BMNH1886.12.15.- 420 (dry): same locality. BMNH1886,12.15.490 (dry }- same locality. BMNHIRA6.12.15.49 (dry) (fragment AMG2875): same locality, BMNH [886.12.15.419 (dry) same locality. LECTOTYPE of H. nidus- vesparum! — BMNH!R8S85.8.27.73 (fragment BMNH1886.8.27.74): Port Jackson, NSW, 33751'5, 151*16'E. PARALECTOTYPE of H. nius- vesparum; AMGSB899; same locality, HOLOTYPE of H. laxa: BMNH1886.12.13,24: Port Jackson. NSW, 33*51'S, 151^16' E. LECTOTYPE of H. laxa digitata: BMNHIB86.8.27.266. (fragment AMG3652): Porn MEMOIRS OF THE QUEENSLAND MUSEUM Jackson, NSW, 33°51°S, 151*16'E (label 'Psamella digitata’), PARALECTOTYPES of H. laxa digitata: AMZI21-124 (dry) (fragments AMG3759, ZMB6396): same locality. AMZ120: same locality. AMG9125 (dry); same locality. LECTOTYPE of H, lax: minima: BMNHI1886,8.27.75 (fragments BMNH1886.8.27.76-77): Port Jackson, NSW, 33*5I'S, 151^ 16 E. PARALECTOTYPES of H. laxa minima: AMZ]25: same locality, AMG9157 (dry) {fragment AMG3780): same locality. HOLOTYPE of H. gigantea: BMNH1886,12. 13.19: 'E. coast of Australia’. LECTOTYPE of H, gigantea intermedie BMNHLERSG.8.27.91 (fragment BMNH1886.8.27.92)- Broughton L, NSW, 32°36'S, 152?19' E. PARALEC- TOTYPES of H. gigantea intermedia: BMNH- 1886.8.27.79 (dry) (fragments BMNH1886.8.27.80, AMG3762, ZMB1177): Port Jackson, NSW, 33°51°S, 151"16'E. AMAS544 (dry): same locality. AMZIL7: same locality, LECTOTYPE of H. gigantea macro- pora: BMNH1886.8.27.85 (fragment BMNH- 1886.8 27,86); Port Jackson, NSW, 3375['5, 151^I6'E. PARALECTOTYPES of H. gigante mine- ropora: BMNHISS86.8.27.83 (dry) (fragments BMNHISRE6.8,27.84, AMZ714, AMG3760, AMG2878): same locality. AMG9158 (dry): same locality. BMNH1886.12.15.312 (dry): same locality. HOLOTYPE of H. gigantea micropora; BMNH- | 886.8.27.81 (dry) (fragments BMNH 1886.8.27.82, ÀMG3761)- Hllawarra, NSW, 34?33'8, 150°55'E. LECTOTYPE of H. irregularis micropora; BMNH- 1886.2.15.49] (fragment AMG2875): Port Fhilip, Vic, 3809'S, 144^52'E. PARALECTOTYPE of H. irregularis micropera; BMNH1886,12.15,504 (frag- ment AMG2874); same locality. LECTOTYPE of H. irregularis. lamellosa: BMNH1886.12.15.490 (dry) (Fragment AMG2876): Port Phillip, Vic, 38°09°S, 144°52°E, PARALECTOTYPES of H, irrepularis lamellosa; BMNHi886.12.15.419, 420 (dry): same locality. OTHER MATERIAL: QLD- QMG303957. NSW- QMG301359, QMG301441, AMZ2225, VIC- NCIQRGC-3402-O (fragment NTMZ3852). HABITAT DISTRIBUTION. Rocky reefs, sand and shell grit substrata, 3-80m depth; Bass Strait, Maria L, Blackman's Bay (Tas); Port Phillip Bay, Westernport Bay (Vic); Byron Bay, Jervis Bay. Pon Jackson. Port Stephens, Maroubra, Barranjoey, Wlawarra, Broughton l., Sandon Bluffs (NSW); Noosa Heads, Tweed River (SEQ) (Fig. 263D). The reported localities of Torres Strait (FNQ) (Lendenfeld, 1888), and Palaa Brant, Singapore (Dragnewitsch, 1906) are suspect. DESCRIPTION. Shape. Massive, Jobate, cylindricul-digitate, honeycomb reticulate growth forms, up to 385mm long, 240mm wide at basal attachment, with individual digits free at apical end. up to 210mm long, 75mm diameter, usually fused at base. forming globular lobate REVISION OF MICROCIONIDAE 497 FIG. 263. Holopsammui laminaefavosa Carter (holotype BMNH 1886.12.15.312). A, Principal style (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle/ style. C, Section through peripheral skeleton. D, distribution, E, Holotype. F, QMG30144]. mass; attached directly to substrate, no expanded point of attachment. Colour. White alive, brown, sandy brown or dark orange-brown in ethanol. Oscules. Large, up to 7mm diameter, mostly on apex of digits but also in between some meshes of lacunae. Texture and surface characteristics. Flexible digits, harsh, arenaceous, lacunae brittle; surface composed of regular, slightly raised ridges formed by honeycomb reticulation of lacunae; meshes produced by reticulation of lacunae typi- cally widely spaced, 4-13mm apart, cavities formed by meshes relatively shallow, covered by thick tympanised membrane stretched across ad- jacent ridges. Ectosome and subectosome. Membraneous, heavily arenaceous, without a well defined tan- gential spicule skeleton, but often with points of ascending principal styles protruding through surface in sparse bundles, below which are scat- tered individual auxiliary spicules; subectosomal 498 FIG. 264. Holopsamma laminaefavosa Carter (QMG303957), A, Choanosomal skeleton, B, Fibre characteristics. C, Principal styles. D, Ends of principal style. E, Auxiliary styles. F, Ends of auxiliary spicule. region undifferentiated from choanosome, fibres immediately subectosomal. Choanosome. Skeleton irregularly reticulate, with very heavy spongin fibres forming radiating lamellae, fibres radiating and diverging slightly towards periphery, and more-or-less differen- tiated into primary ascending and secondary transverse components; fibre anastomoses form close or open meshed reticulations, 40-7301.m MEMOIRS OF THE QUEENSLAND MUSEUM diameter, without any con- sistent size differences be- tween meshes at core or periphery; primary fibres usually cored by detritus and sparse tracis of choanosomal principal styles, and echinated by pauci- or multispicular tracts of same principal styles; coring and echinat- ing spicules together produce plumose or obli- que, vaguely ascending tracts, usually heaviest on peripheral fibres; principal styles may be absent entire- ly from fibre core in heavily arenaceous specimens; secondary tracts arenaceous or entirely clear of detritus, in the latter case tracts of principal spicules clearly seen; mesohyl matrix usual- ly darkly pigmented, usual- ly with subectosomal auxiliary megascleres dis- persed throughout; choanocyte chambers oval to elongate, 50-90um diameter, often obscured by detritus. Megascleres. Choanosomal principal styles coring and echinating fibres short, straight, rela- tively thick, with tapering (hastate) or rounded smooth bases, telescoped or oc- casionally fusiform points. Length 52-(86.4)-115jum, width 2-(4.1)-7m. Subectosomal auxiliary megascleres strongylte styles, long, thin, straight or flexuous, with rounded or slightly subtylote bases, rounded, telescoped or sometimes hastate points. Length 128-(174.3)- 195,.m, width 1-(2.8)-44um. Microscleres. Absent. REMARKS. This species is similar to H. crassa in growth form, skeletal structure and more-or- less in spiculation, although both species may show considerable variation between regional REVISION OF MICROCIONIDAE populations in these features; they are alsa sym- patric although they have not yet been found on the same reefs (present study), This polymor- phism has been described in detail by Hallmann (1912; as Echinoclathria gigantea) and Wieden- mayer (1989; as E. laminaefavosa), and the "ümerous synonyms erected for this species by Lendenfeld (1885-1889) testify to the confusion that this variability can produce. Similarly, the range of variability within each species may over- lap such that observable differences may be not be sufficient to consistently differentiate all populations, and it is conceivable that the two species may be synonyms, Nevertheless, H, laminaefavosa differs from H, crassa mainly in having a white colouration underwater; a shallow meshed surface honeycomb reticulation with a well formed tympanized membrane stretched be- tween adjacent surface ridges: principal styles have telescoped points; and chelae are absent (see remarks For H. crassa). The species has been well illustrated by Lendenfeld (1885c: 18893) under its numerous synonyms, and more recently by Wiedenmayer (1989), Wiedenmayer (1989) and others indicated that some specimens lack a spicule skeleton, with spicules being replaced by foreign detritus. but this cannot be corroborated from materal examined in this study. Holopsamma macropora (Lendenfeld, 1888) (Figs 265-266) Pleciispa macropora Lendenfeld, 1888: 226, Wilsonella macroporu; Hallmann, 1912: 240. Holopsamme macropora; Hooper & Wiedenmayer, 1994; 283. Not Echinoclathria macropora Whitelegge, 1901: 65, 89, 117; Whitelegge, 1902a: 212; Hallmann, 1912: 271. Not Echinoclathria niacropera Whitlegge, 1907; 304. Not Clathria macropora; Whitelegge, 1901: 91. MATERIAL. HOLOTYPE; AMG9159 (dry); Pon Jackson, NSW, 33*5I'S, ]51* 16" E (published locality of Torres Strait incorrect). OTHER MATERIAL: S AUST- SAMTS4094 (fragments QMG300476, NTM1622). HABITAT DISTRIBUTION. Rock reef; 15-25m depth; Port Jackson (NSW); St. Vincent Gulf (SA) (Fig. 265E). DESCRIPTION. Shape. Small, lobate-digitate, honeycomb reticulate sponge, 80-100mm high, 85-140mm wide, with irregular, bulbous surface lobes up to 30mm high, 35mm diameter. Colour. Live colouration unknown, light brown dry, dark brown in ethanol. 499 Oscules. Small, up to 3mm in preserved state, scattered within meshes of surface lacunae. Texiure and surface characteristics. Firm, com- pressible, flexible; surface honeycomb reticula- tion close-meshed, meshes 3-6mm diameter, surface fibre bundles. (lacunae) with scattered microconules. on exterior edges, relatively deep meshes. Ectosome and subectosome. Surface microscopi- cally hispid with points of principal styles protruding through surface, usually in plumose multispicular brushes; stibectosomal auxiliary spicules tangential to surface in sparse bundles or individually. associated with protruding spicule brushes; choanosomal fibres immediately suhec- tosomal Choanosome. Skeletal architecture reticulate, in- creasingly plumose or plumo-reticulate near periphery, with a clearly differentiated primary and secondary fibre skeletons; primary ascending fibres very heavy, up to 120j2m diameter, vaguely stratified, containing plumose uni-, pauci- or multispicular tracts of choanosomal principal styles. generally increasing in density towards periphery but overall sparsely cored, secondary transverse fibres relatively heavy, up 16 551m diameter, very shor, interconnecting clase-set ascending fibres, usually aspicular, rarely unispicular; some fibres also cored by auxiliary spicules; all fibres abundantly echinated by prin- cipal styles, particularly at core of skeleton; spicule tracts within Hbres become increasingly plumose towards periphery; fibre anastomoses form smal! oval or elongate meshes, 25-1261m diameter; mesohy] matrix very lightly pigmented and auxiliary spicules scattered between fibres, particularly towards periphery; some detritus also Incorporated into mesohyl but usually noi into fibre skeleton; choanocyte chambers oval, 20- 65pm diameter. Megascleres. Choanesomal principal styles. oF subtylostyles coring and echinating fibres straight, moderately thin, with shghtly con- stricted, subtylote or rounded. smooth bases. tapering to fusiform points. Length 49-(68.4j- 96m, width 3-(3.9)-6um. Subectosomal auxiliary megascleres range [rom strongvlote subtylostyles to tornotes; usually long, thin, straight or slightly curved with blackened axial canals, sometimes setuceous, sinuous. with rounded or hastate points. Length 123-(166.1)-184.m, width 1.5- (3.2)-Ap.m. Microscleres, Palmaie isochelae uncommon, many smaller forms twisted, with very short alac. MEMOIRS OF THE QUEENSLAND MUSEUM xL M T Z NNV DAN N 25um FIG. 265. Holopsamma macropora (Lendenfeld) (holotype AMG9159). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary strongylote style. C, Palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, SAMTS4094. G, Holotype. REVISION OF MICROCIONIDAE 501 FIG. 266. Holopsamma macropora (Lendenfeld) (holotype AMG9159). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal styles. D, Ends of principal subtylostyle. E, Auxiliary strongylote style. F, Ends of auxiliary spicule. G, Palmate isochelae. straight shaft, lateral alae completely fused to REMARKS. The two specimens are the only shaft, front ala usually partially, or sometimes validated records for the species. Other records wholly fused to lateral alae. Length 4-(9.3)- quoted as ‘macropora’ by Lendenfeld and 12pm. Whitelegge are spurious (see remarks for Plec- Toxas absent. tispa). From Lendenfeld (1888) and Whitelegge 502 (1901, 1902a. 1907) the species has been misinterpreted such that the holotype was referred (sight unseen) to Wilsonella by Hallmann (1912). This error is a direct result of Lendenfeld's (1888) inaccurate description and probably also the confusion resulting from his continual usage of mecropora for several sup- posedly unrelated species (see remarks for Clathria (Axociella) macropora). The holotype belongs to Holopsamma, with the characteristic honeycombed reticulate growth form and smooth echinating spicules (not acan- those as supposed by Lendenfeld (1888) and Halimann (1912)), whereas all other alleged syn- Lypes of Plectispa macropora belong to different species. In growth form A. macropora shows similarities to A. laminaefavosa, and to a lesser extent H. faves. In spiculation and skeletal con- struction it is reminiscent of specimens referred lu H. laminaefavosa by Wiedenmayer (1989) and to a lesser extent H. ramasa (Hallmann, 1912) (i.e, those specimens with a diverging choanosomal skeleton which becomes increas- ingly dense towards the periphery), Tt is possible that H. macropora is simply a non-arenaceous morph of H. laminaefavosa alihough there are differences in spicule geometry and spicule dimensions to suggest that they are different species, In particular, the possession of long sctaceous quasi-diactinal auxiliary megascleres (which also core fibres, are scattered throughout the mesohyl and associated with spicule brushes on the surface) do not appear in these other species; isochelae are also definitely present in H. macropora (alhiet uncommon in both the holotype and SA specimen), allhough they were not recorded in the original descnption of the species by Lendenfeld (1888), whereas microscleres are definitely not present in H. laminaefuvosa. The type locality was given as Torres Strait by Lendenfeld (1888) but it is probably actually from Port Jackson, NSW (as noted on the AM register). Holopsamma pluritoxa (Pulitzer-Finali, 1982) (Figs 267-268) Echinoclathria pluritoxa Pulitzer-Finali, 1982: 108- 109, text-figs 18-19. Holopsamma pluritoxa, Hooper & Wiedenmaycer, : 283. MATERIAL. HOLOTYPE: MSNG 46938 (not seen): Heron I, Great Barrier Reef, Qld, 23726 S, 151°55'E, 8.x11,1979, coll. A.J. Bruce (by hand).OTHER MEMOIRS OF THE QUEENSLAND MUSEUM MATERIAL: QLD- QMG6786 (fragments QMG300471, NTMZ1588). HABITAT DISTRIBUTION. 3-8m depth, on sand, shell erst and coral rubble substrates; Moreton Bay. Heron L (Fig. 267E). DIAGNOSIS OF HOLOTYPE. Honeycombed reticulate growth form; irregularly reticulate skeleton with pale spongin fibres 20-1004um diameter, fibre meshes 300-500j.m diameter, fibres sparsely cored by thin choanosomal prin- cipal subtylostyles (270-320x4-91,m); echinating spicules absent; sirongylote subectosomal auxiliary subtylostyles scattered throughout mesohyl (230-250x2-3um); abundant toxas wing-shaped or slightly oxhorn (10-I188xI- Aam), palmate isochelae unmodified (13.5jm) (Pulitzer-Finali, 1982). DESCRIPTION. Shape. Subspherical-lobate, shaggy and irregularly honeycombed reticulate construction, 55mm wide, 40mm long. Colour. Live colouration unknown. light grey- brown in ethanol. Oscules. Not seen. Texture and surface characteristics. Sott, com- pressible; surface reticulate fibre bundles in peripheral regions slightly flattened, with micropapillose, shaggy points. Ectosome and subectosome, Surface microscopi- cally hispid, with plumose brushes of choanosomal principal styles protruding, inter- dispersed with subectosomal auxiliary megascleres lying tangential or paratangential to surface; subectosomal skeleton plumoreticulate. Choanosome. Skeleton irregularly pluma-reticu- late, with poorly developed spongin fibres, 20- I53ym diameter, forming irregular ovoid to elongate meshes, 145-500j.m diameter; fibres contain pauci- to mullispicular tracts of chosnosomal principal styles, without any division of primary or secondary fibre com- ponents, forming plumose ascending structures which coalesce aad diverge to produce fibre anas- lomoses; fibres not echinated although choanosomal principal styles may protrude obli- quely; mesohy! matrix relatively heavy, darkly pigmented, granular, with numerous toxas and scattered. subectosomal auxiliary megascleres: choanocyte chambers large, oval, 40-90j.m. Megascleres. Choanosomal principal styles straight or slightly curved, with smooth, rounded or slightly subtylote bases, fusiform points. Length 167-(217.2)-289y1.m, width 4-(5.1)}-7 um. Subectosomal auxiliary megascleres straight, asymmetrical, varing from quasistrongyles with REVISION OF MICROCIONIDAE 503 100 um oring FIG. 267. Holopsamma pluritoxa (Pulitzer-Finali) (QMG300471). A, Principal subtylostyle (c and echinat- ing fibres). B, Subectosomal auxiliary subtylostyle. C, Accolada toxas. D, Section through peripheral skeleton. E, Australian distribution. F, Lateral view. MEMOIRS OF THE QUEENSLAND MUSEUM ML i ia €— € — o oaetie SE FIG. 268. Holopsamma pluritoxa (Pulitzer-Finali) (QMG300471). A, Choanosomal skeleton. B, Fibre charac- teristics. C, Principal subtylostyles. D, Ends of principal subtylostyle. E, Auxiliary strongylote subtylostyle. F, Ends of auxiliary spicules. G, Accolada toxas. REVISION OF MICROCIONIDAE smooth rounded extremities, to quasitylotes with subtylote microspined points. Length 195- (218.8)-262,um, width 2-(2.8)-4j.m. Microscleres. Isochelae absent. Toxas wing-shaped, variable in size, thin or thick, with rounded slight, or large central curva- ture, straight or slightly reflexed points, arbitrari- ly divisible into two size classes. Length I: 8-(26.7)-30j.m, width 0.5-(0.8)- 14cm; length II: 65-(177.0)-2501.m, width 1.5-(2.3)-4.m. REMARKS. This species is unusual in its fibre characteristics and skeletal architecture, lacking true echinating spicules, and having quasidiac- tinal auxiliary megascleres. The specimen from Moreton Bay also differs in some respects from the holotype from Heron I. (known from the original diagnosis; type not seen). The skeletal architecture of the specimen is more obviously plumoreticulate than reticulate (reminiscent of Stylinos; Halichondriidae), ectosomal structure is distinctly plumose (reminiscent of Lissoden- doryx; Myxillidae; ectosomal structure was not mentioned by Pulitzer-Finali, 1982), and isochelae are absent (abundant in the holotype). In other respects the two specimens are obviously conspecific having virtually identical spicule geometries, growth forms and surface features although Pulitzer-Finali's (1982) description is incomplete. Holopsamma ramosa (Hallmann, 1912) (Figs 269-270) Echinoclathria ramosa Hallmann, 1912: 277-279, pl.30, fig.3, text-fig.62. Axociella ramosa; de Laubenfels, 1936a: 119. Holopsamma ramosa; Hooper & Wiedenmayer, 1994: 283. Echinoclathria macropora; Whitelegge, 1901: 89, 117. Not Plectispa macropora Lendenfeld, 1888: 226. Not Echinoclathria macropora; Whitelegge, 1907: 504. MATERIAL. LECTOTYPE: AMZ949 (dry): Swan- sea, off Lake Macquarie, NSW, 33°11°S, 151°59’E, depth unknown, coll. FIV ‘Thetis’ (trawl). PARALEC- TOTYPE: AMZ40: 64km W. of Kingston, SA, 36°50’S, 139°0S’E, 60m depth, coll. FIV ‘Endeavour’ (trawl), OTHER MATERIAL: S AUST- SAMTS4105 (fragments QMG300487, NTMZ1695). HABITAT DISTRIBUTION. Rock reef and sandy substrata; 8-60m depth; Lake Macquarie (NSW), Kingston SE., Nuyts Archipelago (SA) (Fig. 269E). DESCRIPTION. Shape. Profusely bushy, ar- borescent branching, honeycomb reticulate 505 sponge, up to 165mm high, 220mm wide, with small basal stalk and expanded point of attach- ment; branches numerous, bifurcate, occasional- ly anastomosing, relatively thin, cylindrical or slightly laterally compressed, up to 75mm long, 32mm diameter. Colour. Grey-brown to dark brown in ethanol. Oscules. Moderately large, up to 5mm diameter, scattered between honeycombed lacunae. Texture and surface characteristics. Harsh, slightly rubbery; fibre bundles form more-or-less regular hexagonal or elongated meshes, 1-3mm diameter. Ectosome and subectosome. Membraneous, with points of sparse principal styles protruding through surface, but generally smooth, non- hispid; sparse subectosomal auxiliary megascleres tangential to ectosome, never protruding through surface. Choanosome. Skeletal architecture irregularly reticulate, with small, heavy, evenly compressed spongin fibres, 23-95j.m diameter, divided into primary longitudinal and vaguely ascending, and secondary connecting components; primary fibres contain uni- or paucispicular tracts of choanosomal principal subtylostyles; secondary fibres mostly aspicular, rarely unispicular; fibre meshes close-set near core, 35-85jum diameter, more cavernous towards periphery, 112-220j.m diameter; tendency for peripheral fibres to con- tain spicule tracts of styles in more-or-less- plumose brushes; fibres sparsely echinated by principal subtylostyles particularly on distal mar- gins of peripheral fibres (cf. Hallmann, 1912); mesohyl matrix heavy, darkly pigmented, with scattered subectosomal auxiliary megascleres; choanocyte chambers small, oval, up to 50pm diameter. Megascleres. Choanosomal principal styles straight, short, relatively thick, with tapering, constricted, smooth, slightly subtylote bases, slightly fusiform sharp points. Length 75- (100.7)-144qj.m, width 4.5-(6.8)-8.5 jum. Subectosomal auxiliary subtylostyles or quasi- strongyles (asymmetrical) straight or curved at centre, thin, with smooth, slightly subtylote bases and rounded points. Length 122-(139.8)-157p.m, width 1.5-(2.7)-41um. Microscleres. Palmate isochelae small, poorly silicified, with approximately 30% contort forms, small alae, lateral alae fused entirely to shaft, front ala completely detached, straight shaft. Length 7-(10.2)-124.m. MEMOIRS OF THE QUEENSLAND MUSEUM 506 25um FIG. 269. Holopsamma ramosa (Hallmann) (holotype AMZ949). A, Principal subtylostyle (coring and echinat- ing fibres). B, Subectosomal auxiliary subtylostyle. C, Palmate isochelae. D, Section through peripheral skeleton. E, Known Australian distribution. F, Holotype. G, SAMTS4105. REVISION OF MICROCIONIDAE 188um G^ z | | ! | FIG. 270. Holopsamma ramosa (Hallmann) SAMTS4105). A, Choanosomal fibre characteristics. B, Echinating principal style on fibres. C, Choanosomal skeleton. D, Principal subtylostyle. E, Ends of principal subtylostyle. F, Auxiliary subtylostyle. G, Ends of auxiliary spicule. H, Palmate isochela. 508 REMARKS. Hallmann (1912) suggested that spicule dimensions were highly variable in this species, particularly choanosomal styles, but in comparison with other Holopsamima species these are in fact relatively homogencous, Similar- ly, palmate isochelae are probably native to this species because they were found in all material examined, although Hallmann did not describe them in the dry lectotype. Spicule geometry and certain aspects of skeletal architecture indicate that H. ramosa is allied to H. macropora and H. arborea, diffenng mainly in growth form, spicule dimensions and the presence of microscleres, In- deed Hallmann (1912) erected H. ramosa for Whitelegge's (1901) specimen of Echinoclathria macropora (as distinct from Plectispa macropora Lendenfeld. 1888), but there has obviously been some confusion in the interpretation of the latter species. Any proposed merger of A. macropora, H. arborea and H. ramosa ts not presently sup- ported on the basis of existing material. Holopsamma rotunda (Hallmann, 1912) (Figs 271-272) Echinaclathna rotunda Hallmann, 1012: 282-284, pl.30, Fig... texi-fig.64. Axociella rotunda; de Laubenfets, 193Ga: 119 [note]. Holopsamma rotunda; Hooper & Wiedenmayer, 1994: 284, MATERIAL. LECTOTYPE: AMZI54: Off Swansea, Lake Macquarie, NSW, 33°) lS, 151°59"E, coll. FIV ‘Endeavour’ (dredge). PARALECTOTY PE: AME 1272 (dry): same locality. OTHER MATERIAL. QLD- QMGL2165. VIC- QMG300275 (NCIQ66C- 3288-P) (fragment NTMZ3865). HABITAT DISTRIBUTION. Substrate unknown; up 1a 330m depth; Lake Macquarie (NSW): off Fraser I. (SEQ); Portsea (Vic) (Fig. 271E). DESCRIPTION. Shape. Massive, subspherical, plobulat, honeycomb reticulate sponge, up to 155mm high, 90mm maximum diameter, with short basal stalk and expanded point of attach- ment; sponge insubstantial, hollow intemally, Colour, Dark bluc-grey alive (Munsell 2.5B 6/2), dark brown in ethanol, Oscules. Large, up to 6mm diameter, scattered between lacunae. Texture and surface characteristics. Firm, com- pressible alive, harsh in dry state; open, porous, tuhular external construction; honeycombed sur- face fibre bundles produce very thin, flattened, erect lamellae with regular meshes, up to 4mm diameter, MEMOIRS OF THE QUEENSLAND MUSEUM Ectosome and subectosome. Membraneous, with points of principal subtylostyles protruding through surface in sparse plumose brushes, few subectosomal auxiliary subtylostyles tangential to and below surface. Choanasome. Skeletal architecture distinctly plumo-reticulate, with ascending multi- ni paucispicular primary fibres, interconnected by pauci- or aspicular secondary fibres; fibres rela- tively light, 43-92,um primary fibre diameter, 18-56,.m secondary fibre diameter, prominently flattened; fibre anastomoses form almost regular, circular or oval meshes, 38-96).m diameter, more compacted towards periphery than at core; fibres cored and echinated by single category of choanosomal principal subtylostyle; echinating megascleres predominant on (although not con- fined to; cf. Hallmann, 1912) distal edges of fibres; primary spicule tracts increasingly plumose towards periphery; mesohyl heavy in ectosomal region, relatively light in deeper choanosomal region, containing sparsely dis- persed subectosomal auxiliary subtylostyles; choanocyte chambers small, oval, up to 55m diameter, Megascleres. Choanosomal principal subtylos- tyles straight or slightly curved at centre, relative- ly Jong, thick, with lapering, constricted, smooth, subtylote bases. fusiform pointed or slightly tele- scoped points. Length 85-(107.5)-L38um, width 4-(7.2)-9.5j m. Subectosomal auxiliary subtylostyles thin, straight, slightly curved or sometimes sinuous, slightly subtylote bases, varying from asymmetri- cal strongylote with rounded points or stylosd with tapering points. Length 123-(152.3)- 196m, width 2,5-(2.9)-41.m. Microscleres. Palmate isochelae abundant, un- modified, with long lateral alae completely at- tached to shaft, shorter front ala partially fused to latera! alae, straight shaft. Length 8-(10.7)- }2yum. REMARKS. Holopsanima rotunda has an insub- stantial, hallow morphology, paper thin, kamel- late fibre bundles (‘lacunae’), and regularly lamellate fibre reticulation with prominent flat- tened fibres, whereas in spicule geometry it could match most other species, particularly H. crassa, H. elegans, H. macropora and H. ramosa (1... with strongylote points on auxiliary spicules). So far the species is only known only from two type specimens, à poorly preserved deep water specimen from the E. continental shelf (repre- sented only by a fragment), and a more recent REVISION OF MICROCIONIDAE 509 FIG. 271. Holopsamma rotunda (Hallmann) (lectotype AMZ154). A, Principal subtylostyle (coring and echinat- ing fibres). B, Subectosomal auxiliary subtylostyle. C, Palmate isochelae. D, Section through peripheral skeleton. E, Known Australian distribution. F, Lectotype. 510 MEMOIRS OF THE QUEENSLAND MUSEUM FIG, 272. Holopsamma rotunda (Hallmann) (QMG300275). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal subtylostyle. E, Auxiliary subtylostyle. F, Ends of auxiliary spicule. G, Palmate isochela. collection from Portsea, Victoria (unfortunately also now only represented by a fragment). The deeper water specimen differs from the others in having some detritus scattered throughout the mesohyl, and also lacking any clear division be- tween ascending and transverse fibre elements, but otherwise it is clearly conspecific. Hallmann (1912) did not record a locality for the type but specimen labels and registers indicate Lake Mac- quarie, NSW. Holopsamma simplex (Lendenfeld, 1885) (Figs 273-274) Halme simplex Lendenfeld, 1885c: 301-303, pl.26, fig.3, pl.27, fig.6. Holopsamma simplex; Hooper & Wiedenmayer, 1994: 284. MATERIAL. LECTOTYPE: AMG8822 (dry): Torres Strait, Qld, 9?41'S, 142*17'E, no other details known. PARALECTOTYPE: BMNH1886.8.27.68 (dry) (fragment AMG3778): off Port Jackson, NSW, HABITAT DISTRIBUTION. Substrate unknown, 10-20m depth; Torres Strait (FNQ); Port Jackson (NSW) (Fig. 273D). DESCRIPTION. Shape. Thickly encrusting spon- ges, reportedly up to 20mm thick, with lobate margins anda maximum lateral spread of 100mm; surviving por- tion of lectotype is 30mm diameter and 10mm thick. Colour. Live colouration apparently brown, beige- brown in dry state. Oscules. Undifferentiated from honeycomb reticulate trabeculae. Texture and surface char- acteristics, Surviving por- tion of lectotype brittle, partially crumbled when dry; surface minutely porous, partially arenaceous; close reticulation of round- ed branches or 'lacunae' form interweaved ridges on encrusting base produced by irregularly honeycombed reticulate surface fibres. Ectosome and subec- tosome. Membraneous, with thick sand cortex; no specialised spicules or spongin fibres visible on surface; subectosome arenaceous. REVISION OF MICROCIONIDAE 51i 50 um FIG. 273. Holopsamma simplex (Lendenfeld) (lectotype AMG8822). A, Principal styles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyles. C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. Im -— t3 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 274. Holopsamma simplex (Lendenfeld) (lectotype AMG8822). A, Principal styles. B, Ends of principal styles. C, Auxiliary styles/ subtylostyles. D, Ends of auxiliary styles. Choanosome. Skeleton completely arenaceous, consisting of densely packed sand grains side-by- side, bonded together with abundant brown col- lagen; fibres absent; collagen covers surface of individual sand grains with smooth choanosomal principal styles embedded in and protruding from (? echinating) this basal layer, or sand grains may be free within mesohyl; subectosomal auxiliary subtylostyles scattered throughout mesohyl; choanocyte chambers not observed. Megascleres. Choanosomal principal styles small, smooth, straight, with smooth, rounded or very slightly swollen bases, usually with tele- scoped points, sometimes slightly fusiform. Length 43-(50.8)-62j.m, width 3-(4.1)-5j.m. Subectosomal auxiliary spicules thin subtylos- tyles, occasionally quasi-strongyles, straight, REVISION OF MICROCIONIDAE slightly curved or sinuous, with asymmetrical ends. Length 68-(84.8)-994m, width 2-(2.7)- 3.5pm. Microscleres. Absent. REMARKS. This species is known only from two relatively poor type specimens. Lendenfeld’s (1885c: 303) published geographical distribution for this species, supposedly including also Port Phillip Bay, Vic, NT waters, and Mauritius, is unsubstantiated by voucher material and not ac- cepted here. This species is similar to E. (Protophlitaspongia) bispiculata in geometry of principal megascleres, and with several other arenaceous Holopsamma species (i.e., H. favus, H. crassa, H. laminaefavosa, H. felixi), although in H. simplex fibres cannot be differentiated from aggregated sand grains. Holopsamma sp. indet. (Fig. 275, Plate 11F) MATERIAL. QMG300620 (NCIQ66C-2389-N) (fragment NTMZ3563): Marion Reef, off Edithburgh, S. Yorke Peninsula, SA, 38?08.5'S, 137?48.0' E, 6m depth, 10.ii.1989, coll. NCI. HABITAT DISTRIBUTION. 6m depth; on algae covered rock reef; SAust (Fig. 275B). DESCRIPTION. Shape. Groups of subspherical, bulbous lobes fused together, each composed of honeycombed reticulate lamellae producing a delicate Bryozoan-like lace. Colour. Pale red-brown alive (Munsell 10R 7/8), brown in ethanol. Oscules. Large, up to 6mm diameter, on apex of lobes; small ostia 1-2mm diameter scattered amongst surface lacunae. Texture and surface characteristics. Harsh; sur- face membraneous, with distinct concentric ridges, slightly arenaceous layer on outer edge of ridges, fleshy between ridges. Ectosome and subectosome. No ectosomal spicule skeleton; ectosome prominently arenaceous, fleshy below surface; ectosomal sand particles dispersed in association with primary ascending fibres. Choanosome. Irregularly reticulate skeletal structure, with broad fibres up to 400um diameter, partially or fully cored with detritus; mesohyl matrix very heavy; no native spicules; choanocyte chambers oval to elongate, 20-55p1.m diameter. Megascleres. Absent. Microscleres. Absent. 513 REMARKS. This single specimen is similar to several Holopsamma species (e.g., H. crassa, H. laminaefavosa) in being able to shed its spicules completely and replacing these with foreign detritus, particularly sand particles. Wieden- mayer (1989) discusses these arenaceous species in detail and the difficulty in assigning them to any particular taxon. On the basis of its growth form, fibre characteristics and skeletal architec- ture the specimen could be assigned to one of several species, and for the time being its specific identity is left indeterminate. Echinochalina Thiele, 1903 Refer to subgenera for synonymy. TYPESPECIES. Ophlitaspongia australiensis Ridley, 1884a: 442 (by subsequent designation of Hallmann, 1912: 288)). DEFINITION. Monactinal, quasi-monactinal or thin diactinal auxiliary megascleres tangential or erect on ectosome; choanosomal skeleton ir- regularly reticulate; fibres cored by tracts of auxiliary megascleres, identical to those in ec- tosomal skeleton, and echinated by principal megascleres varying from true monactinal, quasi- diactinal to true diactinal forms, smooth or acan- those; microscleres may include palmate isochelae and toxas. REMARKS. Twenty six species have been in- cluded in Echinochalina, of which 20 are valid, of which 16 live in Australian waters including 5 new species. All species are known from the Indo-west Pacific region (Hooper & Lévi, 19932). The genus contains 2 groups: one (Echino- chalina (Echinochalina)) with true monoactinal spicules, showing superficial affinities with Holopsamma (having honeycomb reticulate growth forms) and the Raspailiidae (fibre charac- teristics), and the other with quasi-diactinal or secondarily modified diactinal megascleres (Echinochalina (Protophlitaspongia)) superfi- cially resembling Niphatidae (Haplosclerida). Both groups are linked by the common posses- sion of auxiliary megascleres coring fibres and principal megascleres echinating fibres. Echinochalina (Echinochalina) Thiele, 1903 Echinochalina Thiele, 1903a: 961; sensu Hallmann, 1912: 288. 514 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 275. Holopsamma sp. indet. (QMG300620). A, Choanosomal skeletal structure. B, Australian distribution. Tablis de Laubenfels, 1936a: 76. [Echinoclathria]; Uriz, 1988: 89. TYPESPECIES. Ophlitaspongia australiensis Ridley, 1884a: 442. DEFINITION. Monactinal, quasi-monactinal or secondarily derived true diactinal auxiliary megascleres core fibres, and monactinal principal spicules echinate fibres. Echinochalina (Echinochalina) anomala Hallmann, 1912 (Figs 276-277) Echinochalina glabra, in part; Whitelegge, 1907: 507. Not Echinoclathria glabra Ridley & Dendy, 1887: 163. Echinochalina anomala Hallmann, 1912: 292-294, text-fig.68; Hooper & Wiedenmayer, 1994: 276. Tablis anomala; de Laubenfels, 1936a: 76. cf. Echinochalina anomala Burton, 1934a: 563. MATERIAL. HOLOTYPE: AMG10548 (dry): Off Woolongong, NSW, 34°25’S, 151?10'E, 110-112m depth, coll. FIV ‘Thetis’ (dredge). PARALEC- TOTYPES - AMG10549 (dry): same locality. AMG10550 (dry): same locality. AMG10551 (dry): same locality. HABITAT DISTRIBUTION. 110-112m depth; sub- strate unknown; S. coast (NSW) (Fig. 276D). DESCRIPTION. Shape. Massive, clathrous- honeycomb reticulate sponge, 125mm long, 75mm maximum width, composed of numerous, small, flattened fibre-branches, up to 12mm long, 4mm diameter, anastomosing and bifurcating REVISION OF MICROCIONIDAE 100 um 515 FIG, 276. Echinochalina (Echinochalina) anomala Hallmann (holotype AMG10548). A, Principal subtylostyle (echinating fibres). B, Auxiliary subtylostyle/ quasistrongyle (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Holotype. forming tangled mass, attached directly to sub- strate. Colour. Live colouration unknown, grey-brown in dry state, Oscules. Not observed. Texiure and surface characteristics. Firm, harsh in dry stale; surface fibres flattened, paper-like, porous reticulate with fibre-branches forming wide-meshed reticulation up to 3mm diameter. Ectasome and subectosome. Membraneous, stretched across adjacent surface fibre-branches, with tangential auxiliary megascleres sparsely dispersed within ectosome, singly or less fre- quently in paucispicular bundles, and protruding echinating principal styles protruding through surface; choanosomal fibres immediately below ectosome. Choanosome. Irregularly reticulate skeletal structure; primary spongin fibres large, heavy, 230-360,.m diameter, irregularly interconnected by relatively thinner secondary fibres, 95-145,.m diameter, forming cavernous meshes, 430- 1090m diameter; all fibres cored by multi- spicular tracts of auxiliary megascleres 516 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 277. Echinochalina (Echinochalina) anomala Hallmann (holotype AMG10548). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal spicule. E, Auxiliary subtylostyle. F, Ends of quasi-diactinal auxiliary spicule. occupying most of fibre diameter; fibres moderately heavily echinated by principal styles dispersed evenly over surface; mesohyl matrix light, containing numerous dispersed subec- tosomal auxiliary spicules; choanocyte chambers not observed (dry material). Megascleres. Principal styles echinating fibres relatively thick, straight or slightly sinuous, with smooth, slightly subtylote bases, fusiform points. REVISION OF MICROCIONIDAE 517 FIG. 278. Echinochalina (Echinochalina) australiensis (Ridley) (holotype BMNH1881.10.21.299). A, Principal subtylostyles (echinating fibres). B, Auxiliary subtylostyle/ tornostyles (coring fibres and interstitial). C, Section through peripheral skeleton. D, Australian distribution. E, Holotype. Length 160-(189.9)-2361 m, width 7-(9.3)- llum. Auxiliary megascleres coring fibres and found interstitially long, thin, straight, entirely smooth, varying from nearly symmetrical quasi-stron- gyles to asymmetrical tornostyles with slightly subtylote bases and rounded points. Length 165- (215.4)-277um, width 2-(4.3)-6.5 jum. Microscleres. Absent. REMARKS. Burton (1934a) assigned a specimen from Eagle I. (Cairns section, Great Barrier Reef) to E. (E.) anomala but his material 518 was not found in the BMNH. Moreover, Burton's figure does not resemble the holotype so that his record is dubious. Skeletal fibre reticulation of E. (E) anomala is reminiscent of Echinodic- tyum (Raspailiidae), with very widely spaced fibre- meshes, and with fibres of large diameter which are heavily cored by megascleres. Unlike most Echinodictyum species, however, which have a greater emphasis on the mineral skeleton over the or- ganic fibres, and which have true diactinal coring spicules, the fibres of E. (E.) anomala are prominent and heavily cored by quasi-diac- tinal megascleres. The honeycomb reticulate growth form of E. (E) anomala is similar to E. (E.) barba (both reminiscent of Holopsamma) | differing mainly in the geometry of its principal megascleres, dif- ferences in fibre construc- tion and spicule dimensions. Echinochalina (Echinochalina) australiensis (Ridley, 1884) (Figs 278-279) Ophlitaspongia australiensis Ridley, 1884a: 442-443, pl.42, figs c,c’. Echinochalina australiensis; Thiele, 1903a: 961-962; Hooper & Wiedenmayer, 1994: 276, MATERIAL. HOLOTYPE: BMNH1881.10. 21.299: Port Molle, Qld, 20°16'S, 148?42'E, L.v.1881, HMS ‘Alert’ (dredge). OTHER MATERIAL: SMFI855 (fragment MNHNDCL2265). HABITAT DISTRIBUTION. Rock and coral rubble substrata; 24m depth; Airlie region (NEQ) (Fig. 278D); also Moluccas, Indonesia. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 279, Echinochalina (Echinochalina) australiensis (Ridley) (holotype BMNH188]1.10.21.299). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal spicule. E, Auxiliary spicule. F, Ends of quasi-diactinal auxiliary spicule. DESCRIPTION. Shape. Massive, branching, clathrous, honeycomb reticulate growth form, 55mm diameter, fibre-branches flattened, up to 12mm long, 5mm wide, with cavernous, angular meshes up to 10mm diameter. Colour. Pale brown dry. Oscules. Small, up to 4mm diameter, between adjacent fibre bundles. 25um 519 REVISION OF MICROCIONIDAE FIG. 280. Echinochalina (Echinochalina) barba (Lamarck) (AMZ44), A, Principal style (echinating fibres). B, Auxiliary subtylostyle/ quasistrongyle (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Paralectotype MNHNDT341 1. F, Lectotype of S. favosa. G, Holotype of E. glabra. MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 281. Echinochalina (Echinochalina) barba (Lamarck) (QMG300274). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal spicule. E, Auxiliary spicule. F, Ends of quasi-diactinal auxiliary spicule. REVISION OF MICROCIONIDAE Texture and surface characteristics. Firm, harsh dry; canulose surface produced by anastomosing fibre bundles ('lacunae'), interconnected by translucent ectosomal membrane stretched be- tween adjacent conules. Ectosome and subectosome. Membraneous, with thinner choanosomal, undulating fibres lying im- mediately below ectosome, approximately 53m diameter, cored by paucispicular tracts of auxiliary megascleres, and with irregular layer of auxiliary megascleres scattered tangential to sur- face. Chaanosome. Vaguely regularly reticulate skeleton, composed of primary, distinctly ascend- ing, multispicular fibres. 100-140um diameter, interconnected ai irregular intervals by numerous transverse, paucispicular, secondary fibres. ap- proximately 70pm diameter; fibre anastomoses form rounded or elongate, narrow meshes up to 270pm diameter; fibres cored by auxiliary megascleres, sparsely echinated by principal sub- tylostyles; mesohy! matrix heavy but only lightly pigmented, with sparsely dispersed auxiliary megascleres between fibres; choanocyte. cham- bers ovoid, 20-25,4cm diameter, Megascleres. Principal subtylostyles (echinating fibres) straight, smooth, tapering, rounded or slightly subtylote bases, fusiform or slightly tele- scoped points. Length 83-(111.2)-I31um, width 6-(7.2)-9.5pm. Auxiliary megascleres (conng fibres and inter- stitial) straight, thin, subtylostyles. tórnostyles or quasi-stongyles, with smooth, rounded, or very slightly subtylote bases, tapering-rounded points. Length 147-(178.2)-[92um, width 1.5-(3.4)- 4 5um. Microscleres. Absent REMARKS. Ridley (18843) reported toxas (up to 42pm long, 2m wide) but these were not found in either the holotype or in Thiele's (1903) material and they are probably contaminants. E. (E.) australiensis, like most members of the genus, has few diagnostic characteristics, super- ficially resembling Echinodictyum cavernosum (Raspaiiidac) and C. (Thalysias) vulpina in growth form, with a heavy, well developed reticu- late, Clathria-like, horny fibre skeleton cored by auxiliary spicules and echinated by principal spicules. It differs from other Echinochalina in spicule dimensions, well-developed skeletal Structure and these fibre characteristics. 521 Echinochalina (Echinochalina) barba (Lamarck, 1814) (Figs 280-281, Plate 12A) Spongia barba Lamarck,1814: 372, 354; Topsent, 1930- |5-16, pl.4,hg.6. Erhinochalina barba; Hooper & Wiedenmayer, 1994: 276. Spongia favosa Lamarck, 1814: 373; Topsent, 1930; 19, pt.1, fig.5. Echinoclathria glabra Ridley & Dendy, 1886: 476; Ridley & Dendy, 1887. 163, pl.29, fig.1l, p31, fig.2; Dendy, 1896: 40. Tablis glabra; de Laubenfels, 19362: 76. Echinochalina glabra; Thiele, 1903a: 962; Hallmann, 1912: 275, 290-292, text-hg.67; Burton, 19342: 600; [2] Thomas, 1977: 115-116, text-figs 1F-G. Not Echinochalina glabra; Whitelegge, 1907; 504. MATERIAL, LECTOTYPE: MNHNDT561: un- known locality. PARALECTOTYPE: MNHN- DT3411: unknown locality. LECTOTYPE of 5 Javasa: MNHNDTS557 (fragment BMNH1954,2.20.9): Near King J., Tas, 39°50'S, 144°00'E, 1803, Peron & Lesueur collection. PARALECTOT YTE of S. favosa: MNHNDT 3401 (fragment BMNH1954.2. 20,37): same locality, HOLOTYPE of E. glabra: BMNH 1887.5.2.63: Bass Strait, Tas, 39°10.3°S, 146^37' E, 76m depth, 2.iv.1874, coll. HMS "Challenger" (dredge), OTHER MATERIAL: OLD-QMGL711 (fragment NTMZIS31). TAS- QMG300274 (NCIQ66C-3543-R) (fragment NTM23849), OMG311312 (NCIQ66C3542-Q) (fragment NTMZ3853), $ AUST- AMZ44, HABITAT DISTRIBUTION. Sand, shell grit, rock or coral rubble substrata, kelp beds; 0-76m depth; Mon- coeur I, Port Phillip Bay (Vic); King L, Deal 1., Bass Strait (Tas); Kingston SE. (SAY; Flinders [s (FNQ) (Fig. 281D). Thomas' (1977) reported locality of An- daman Sea is dubious from his description. DESCRIPTION. Shape. Massive, subcylindical, honeycomb reticulate sponge, upto [55mm long, 75mm wide, consisting of thin, flattened, ir- regularly anastomosing, reticulate fibre bundles; fibre anastomoses form meshes 3-Smm diameter, Colour. Light grey-brown exterior, beige interior alive (Munsell 7.5Y8/6), yellowish brown in ethanol. Oscules. Small, up to 3mm diameter, recessed, dispersed between surface fibre bundles. Texture and surface characteristics. Spongy, slightly rubbery, compressible, difficult to tear; surface of fibre bundles smooth, even, glabrous, porous and reticulate elsewhere, surface membrane stretched across adjacent fibre bundles. Ectosome and subectosome. Thinly membraneous, with irregularly dispersed 522 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 282. Echinochalina (Echinochalina) felixi sp.nov. (holotype QMG304741). A, Auxiliary subtylostyle (coring fibres and interstitial). B, Principal subtylostyles (echinating fibres). C, Wing-shaped and accolada toxas. D, Palmate isochelae. E, Section through peripheral skeleton. F, Known Australian distribution. G, Holotype. REVISION OF MICROCIONIDAE auxiliary megascleres lying tangential to surface and irregular, paucispicular, plumose brushes of principal spicules erect on peripheral fibres, bare- ly protruding through ectosome: choanosomal fibres immediately subdermal. Choanosome. Irregularly reticulate skeletal Structure, with primary, vaguely ascending, dis- tinctly meandering paucispicular fibres, 52- 120p m diameter, interconnected by aspicular or less frequently paucispicular, irregularly anas- tomosing, secondary fibres, 37-75,.m diameter; fibres cored by auxiliary megascleres, moderate- ly echinated by principal subtylostyles especially on peripheral fibres; fibre anastomoses form nearly regular regular oval meshes, 165-3704. m diameter; mesohyl matrix heavy, moderately lightly pigmented, with numerous auxiliary megascleres dispersed between fibres; choanocyte chambers oval to eliptical. 38- 145m diameter. Megascleres. Principal subtylostyles (echinating fibres) short, thick, straight or very slightly curved, with smooth, tapering, distinctly sub- tylote bases, fusiform points. Length 62-(94.3)- 15pm, width 5-(7.1)-8.5jum. Auxiliary megascleres (coring fibres and inter- stitial) straight, quasi-diactinal subtylostron- gyles, rarely strongyles. with smooth, slightly subtylote bases and rounded points. Length 147- (189.2}-214p.m, width 1.5-(3.3)-4.5 um. Microscleres. Absent. REMARKS, Echinochalina borba has a honeycombed reticulate construction similar to that of E. (E) anomala and reminiscent of Holop- samma species, hut in which the surface is very smooth and fleshy (superficially resembling an Acantheila (Axinellidae)). Both E. ano mala and E. barba also have asymmetrical (quasi-diac- tinal) auxiliary megascleres coring fibres. whereas in most other Echinochalina these are more obviously monactinal.In E. (E.) barba fibre anastomoses are relatively closely compacted and principal styles (echinating fibres) are short, tapering, subtylote, whereas E. (E.) anomala has a cavernous construction and long, more-or-less rounded principal megascleres. Echinochalina (Echinochalina) felixi sp. nov. (Figs 282-283) MATERIAL, HOLOTYPE: QMG304741: NE of Eagle 1., E. of Turtle [s group, Cairns section Great Barrier Reef, 14^39,8' S, 145°19.2°E, 3.ix.1994, 19m depth, coll. J. Kennedy, DP] RV "Gwendolyn May". trawl, HABITAT DISTRIBUTION. Soft substrata iater-reet region; 19m depth; Turtle Is (FNQ) (Fig. 282F). DESCRIPTION, Shape. Long, thinly cylindrical, fragile digit, 180mm long, up to 60mm diameter japenng towards base; embedded in soft substrata by small, cylindrical, basal root-like processes, up to 21mm long, 3mm diameter. Colour. Beige-grey alive (Munsell 2.5Y 7/2), light brown im ethanol. Oscules. Not observed. Texture and surface characteristics, Firm, com- pressible; surface fibrous, macroscopically even, microscopically porous, honeycomb reticulate. Ectosome and subectosome.Fibrous, with peripheral spongin fibres tangential to surface, forming circular meshes up to 2.5mm diameter, membranecus ectosome stretched between ad- jacent fibres; outer surface of peripheral fibres with small blind fibre endings and protruding spicules, both visible under low magnification; auxiliary subtylostyles scattered on membrancous ectosome, singly or in small bundles; subectosomal skeleton undifferentiated from choanosome, Choanosome. Reticulate skeletal structure: fibres more-or-less homogeneous, 40-85ym diameter cored hy auxiliary subtylostyles in multispicular tract of up to 6 spicules abreast: fibres long, generally curved or sinuous, anastomosing, regularly bifurecare; shorter connecting fibres uni- or paucispicular; fibre meshes cavernous oval or eliptical, 170-490jcm. diameter; fibres sparsely echinated by smaller principal subtylostyles; mesohyl matrix heavy, granular, cored by few microscleres and auxiliary megascleres; choanocyte chambers small. oval, 20-35pm diameter. Megascleres. Principal subtylostyles (echinating fibres) short, slender, straight, subtylote, smooth bases, tapering fusiform points. Length 207- (242.2)-2781.m, width 3-(3.3)-4.54.m. Auxiliary subtylostyles (coring fibres and dis- persed in mesohyl) long. very slender, straight, slightly subtylote, smooth bases, tapering fusiform or sometimes bluntened points, Length 285-(314.8)-345pm, width 3.5-(4.3)-5j.m, Microscleres. Palmate isochelae unmodified, front and lateral alae approximately equal length, lateral alae completely fused to shaft, front alae partially fused to lateral alae, straight shaft. Length 6-(10.1)- I2u.m. Toxas include both wing-shaped and slightly accolada forms, ranging from thin to raphidiform. often broken in preparations. Length 40-(65.8)- pum. width 0,8-(1.7)-2,5jum. 524 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 283. Echinochalina (Echinochalina) felixi sp.nov. (holotype QMG304741). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyles. D, Ends of principal spicule. E, Auxiliary subtylostyle. F, Ends of auxiliary spicule. G, Palmate isochelae. H, Fragments of wing-shaped and accolada toxas. 525 REVISION OF MICROCIONIDAE 25um FIG. 284. Echinochalina (Echinochalina) gabrieli (Dendy) (holotype NMVG2359). A, Principal subtylostyles (echinating fibres). B, Auxiliary subtylostyle (coring fibres and interstitial). C, Oxeote toxa. D, Section through peripheral skeleton. E, Holotype. F, Known Australian distribution. ETYMOLOGY. For Dr Felix Wiedenmayer, Naturhis- REMARKS. Echinochalina felixi is unusual torisches Museum Basel, for his contributions to having very thin megascleres and a cavernous Australian sponges. ; A Y A k reticulate construction, similar to E. digitata al- 526 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 285, Echinochalina (Echinochalina) gabrieli (Dendy) (holotype NMVG2359). A, Choanosomal skeleton. B, Fibre characteristics (x419). C, Principal style. D, Ends of principal style. E, Auxiliary style. F, Ends of auxiliary spicule. G, Oxeote toxa. H, Point of toxa. though in that species fibres are both cored and auxiliary styles core fibres, growth form is echinated by principal styles, whereas in E. felixi | cylindrical digitate, and toxas are present. REVISION OF MICROCIONIDAE Echinochalina (Echinochalina) gabrieli (Dendy, 1896) (Figs 284-285) Ophlitaspongia gabrieli Dendy, 1896: 38-39, Echinochalina gabrieli; de Laubenfels, 1936a: 119; Hooper & Wiedenmayer, 1994: 277. MATERIAL. HOLOTYPE: NMVG2359 (fragment BMNH1902.10.18,341): Port Phillip Bay, Vic, 38°09'S, 144°52’E, 11-13m depth, coll. J.B. Wilson (dredge). HABITAT DISTRIBUTION. Encrusting of shell frag- ments, 1 i-|3m depth; Port Phillip (Vic) (Fig. 284F). DESCRIPTION. Shape. Massive, lobate, semi- encrusting on detritus, 130mm long, 65mm wide, up to 45mm thick, with low bulbous lobes scat- tered over surface. Colour, Brown in ethanol. Oscules. Single. large apical oscule, up to 3.5mm diameter, on each surface lobe. Texture and surface characteristics. Firm, com- pressible, easily torn; surface lobate-bulbous, un- even, microscopically rugose, reticulate, granular. Ectosome and subecrosome. Membraneous, skin- like ectosomal skeleton, minutely hispid with points of auxiliary styles protruding through sur- face in pauci- or multispicular plumose brushes, some arising from ascending choanosomal skele- tal tracts, others unattached to peripheral fibres and free within mesohyl; sparse tangential tracts of auxiliary styles below surface. Choanosome, Irregularly reticulate skeleton, with thin but well developed spongin fibres form- ing primary, more-or-less ascending, and secon- dary connecting tracts; primary fibres, 40-70,.m diameter, cored by multi- or paucispicular tracts of auxiltary styles in plumose or disorganised arrangement; secondary connecting fibres, 15- 32um diameter aspicular or unispicular, curved, meandering throughout core of choanosomal skeleton producing oval, elongate, cavernous meshes, 180-3101.m diameter, becoming more regular, rectangular and cavernous towards peripheral region of skeleton, with meshes 180- 360j.m diameter; peripheral fibres slightly more compacted; spicule tracts heavier in subec- tosomal skeleton than at core; primary fibres sparsely echinated by principal styles; mesohyl matrix heavy, moderately heavily pigmented, numerous principal styles dispersed between fibres; choanocyte chambers elliptical, 62- 135um diameter. Megascleres. Principal styles echinating fibres straight, short, relatively thick, with smooth, rounded or slightly constricted, subtylote bases and nearly hastate points. Length 102-(117.7)- 138m, width 3,8-(5.5)-6.5j.m. Auxiliary styles, coring fibres and interstitial, straight or slightly curved, thin, relatively long, fusiform, with smooth, rounded or very slightly subtylote bases. Length 142-(213.1)-2644um, width 1.8-(2 7)-4.5jum. Microscleres. Isachelae absent. Toxas moderately uncommon, oxeote, with slight angular central curvature, straight points. Length 90-(116.9)-135j1.m, width 1.5-(2,2)- 2.5m, Larvae. Visiparous, oval-elongate parenchymel- la, 225-375x 190-210,um, dark orange, heavily collagenous. containing oxeote toxas und juvenile styles. REMARKS. De Laubenfels (19362) referred this species to Echinochalina (sensu Hallmann, 1912) having fibres cored hy megascleres geometrically equivalent to auxiliary spicules, whereas styles echinating fibres have a geometry equivalent tù principal megascleres of other Microcionidae. Dendy's (1896) descnption differed in spiculc dimensions from my observations of the type. Oxeote toxas, much smaller than those described by Dendy, are predominantly in association witli numerous larvae, and it is possible that those spicules are merely juvenile quasidiactinal morphs of (auxiliary) coring megascleres. The species differs from others in its spicule geometry and size. in particular the toxa morphology, und skeletal structure whereby spicule tracts increase in density towards the peripheral skeleton. Echinochalina (Echinochalina) intermedia (Whitelegge, 1902) (Figs 286-287, Table 44, Plate 12B) ? Echinoclathria viminalis, in part, Whitelegge, 1901: 87-88. Not Thalassodendron viminulis Lendenfeld, 1888: 2285. Echinoclathria intermedia; in part, Whitelegge, 1902a- 214-215; Dendy, 1922: 71, pl.2, fig.8. Echinochalina intermedia; Halimann, 1912; 294-295, text-fig.69; Shaw, 1927: 427; Burton, 19348; 563; Hooper & Lévi, 19932: 1273-1277, figs 27-28, table 12: Hooper & Wiedenmayer, 1994; 277. MATERIAL. HOLOTYPE: AM: Lendenfeld's No. 365 (missing). NEOTYPE: QMG300025: Inner Gneerings, off Mooloolaba, 26°39°S, |53*10" E, 25m depth, 10.x:t 1991. 528 MEMOIRS OF THE QUEENSLAND MUSEUM 25um FIG. 286. Echinochalina (Echinochalina) intermedia (Whitelegge) (neotype QMG300025). A, Principal styles (echinating fibres). B, Auxiliary subtylostyle/ tornostyle (coring fibres and interstitial). C-D, Principal and auxiliary subtylostyle of Hallmann (1912) specimen. E, Section through peripheral skeleton. F, Australian distribution. G, Neotype. OTHER MATERIAL: (see Hooper & Lévi, 1993, for QMG300834, QMG300865. INDONESIA - a list of additional material used in this study); QLD- © NCIOCDN-1306-F (fragment QMG303688). QMG304757, QMG304293, QMG304388, QMG303863, QMG304025, QMG303984, HABITAT DISTRIBUTION. Sand, rock and coral rubble substrata; 8-50m depth; Lizard I., Day Reef, REVISION OF MICROCIONIDAE (Echinochalina) 287. Echinochalina QMG300834). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal style. D, Ends of principal style. E, Auxiliary subtylostyle/ tornostyle. F, Ends of auxiliary spicule. FIG. Direction Is, Howick Is, Snake Reef, Frankland Is, Sudbury Reef, Stanley Reef, Gulf of Carpentaria (FNQ); Hook Reef (NEQ): Mooloolaba, Moreton Bay (SEQ), Illawarra (NSW), E. coast (Tas); Dampier Ar- chipelago, Northwest Shelf (WA) (Fig. 286F); also New Caledonia (Hooper & Lévi, 1993a), Cargados Carajos, Indian Ocean (Dendy 1922); Sulawesi, In- donesia (present study), DESCRIPTION OF NEOTYPE. Massive, lobate, bulbous-digitate, 105mm high, 65mm maximum breadth; convoluted bulbous, cavern- ous surface in life, collapsing to form sharp ridges intermedia (Whitelegge) and conules upon preserva- tion; texture soft, compres- sible, mucusy; pale orange-brown alive, brown in ethanol. Ectosome membraneous, with tangen- tial and paratangential tracts of auxiliary spicules scat- tered, barely protruding through surface; ectosome heavily collagenous, darker than choanosomal mesohyl; choanosomal skeleton ir- regularly reticulate, with long primary fibres, 90- 1304m diameter, meander- ing through choanosome, becoming more plumo- reticulate near periphery, cored by multispicular tracts of whispy auxiliary spicules, interconnected by short. aspicular otl paucispicular secondary fibres, 40-70j.m diameter, producing elongate or elip- tical, cavernous meshes up lo 450j.m diameter; fibres sparsely echinated by prin- cipal spicules; mesohyl matrix heavy, with abundant auxiliary spicules scattered between fibres; choanocyte chambers large, oval, up to 604m diameter. Principal styles short, thick, straight, smooth tapering or slightly subtylote bases, hastate or slightly telescoped points (72-95x2.5-4,5 zm). Auxiliary spicules subtylos- tyles or tornostyles, long, slender, straight, slightly subtylote smooth bases, rounded points (178-206x2-3.5pm). Microscleres absent. DESCRIPTION. (See Hooper & Lévi, 1993a). DIAGNOSIS (Table 44 for spicule dimensions). Lobo-digitate with flattened branches and fibre bundles on surface. forming vaguely "honeycombed' clathrous, reticulate growth form; pale orange-brown alive; large oscules mainly on apex of surface bulbs; surface conulose with low ridges and grooves, paler wanslucent 530 TABLE 44. Compatison in spicule dimensions be- tween populations of Echinochalina (Echineckaline) intermedia (Whitelegge), Measurements in pm (N=25). Comparative data from material examined here and Hooper & Lévi (1993a), Auxiliary subtylostyles Principal styles 178-(190,7)-206 x 2- | PES HSB RT | (2.63.5 Birds | 169-(175.3)-189 x LŁ(L9)-25 (2.8)-4.0 52) Population (Latitude) Neoty 500785 S 26°S) New Caledonia (22°S) L 184-(193,8)-205 x 2,.0-(3.1)-5.0 Great Barrier | Reef (17-19°S) Southern Qld. 184-(196.2)-208 x | 72-(88.3)-101 x 25- (26^8) 15-0.2)3 254 NW Shelf WA 89-(148.2)-180% | 92-(12555-142 x 4- | E S) 1,5-(2.5)-4.0 (5:3 S ^ cub TTE 200x 3 filawarra, NSW? 34°S) 140-185 x4 Bü-150 x 9 177-(183.2)-198 x. | 60-(82.7)-113 x 20- 1.54(1.9)-2.5 32)4 1922). 2. Hallmann (1912). — Sulawesi, Indonesia (1°N) | Source: 1. Dendy ( skin-like membrane covering, darker below; ec- tosome membraneous with tangential or oc- casionally paratangential sparse tracts of auxiliary spicules, lightly arenaceous, and sparse plumose brushes of auxiliary megascleres from ascending choannsomal fibres protruding short distance through surface predominantly on sur- face conules; choanosomal skeleton irregularly plumo-reticulate, spongin libres well developeu; primary fibres lightly cored by multispicular tracts of auxiliary megascleres becoming increas- ingly plumose towards penphery; secondary con- necting fibres uni-. pauci- or aspicular; all fibres irregularly echinated by sparse pnncipal styles, mainly on primary fibres; fibre anastomoses form irregular elongate, oval, eliptical or rarely rectan- gular meshes, meshes more cavernous in periphery than at core; mesohyl matrix heavy, numerous auxihary spicules and detritus scat- tered between fibres; principal styles echinating fibres short, thick, straight, almost hastate, with smooth, rounded, tapering, or less frequently with very slightly subtylote bases; auxiliary megascleres coring fibres and dispersed wilhin mesohyl long, thin, mostly straight, with rounded smooth bases, varying from styles, tornostyles or less often strongyles, usually with blackened axial canals, with rounded or slightly hastate points; microscleres absent. MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. This species was redescribed ihain- ly from New Caledonian material (Hooper & Lévi (19933). It is very characteristic in the field with a clathrous, *"honeycombed' reticulate growth form and pale orange-brown colour, and differs from other species in having an irregular plumo-reticulate skeletal architecture, poorly developed spongin fibres cored by a dense mul- tispicular axis of primary spicule tracts and light pauci- or aspicular tractsin secondary fibres, and hastate choanosomal (auxiliary) styles core fibres. Hooper & Lévi (1993a) note that the geometry of auxiliary spicules in this species varies. between regional populations, ranging from exclusively styles in the original type material (now lost; Whitelegge, 1901, 19022) and the Northwest Shelf of WA (Hooper & Lévi, 19933), to Virtually all strongyles in material from Mooloolabah including the neotype (present study) and Cargados Carojos (Dendy, 1922), New Caledonian population differs from others in having more abundant detritus and much more slender echinating styles, whereas growth form, skeletal architecture and spicule geometry are relatively homogeneous for the species. Echinochalina intermedia is used in the sense of Hallmann (1912) and Dendy (1922), because the synonymy of this species is sull confused, and the validity of type material of Thalassodendran viminalis and Echinochalinaü intermedia require further clarification. Lendenfeld (1888: 225) crected T. viminalis for a specimen from Illawar- ra. NSW, whereas Whitelegge (1901: 87) found the onginal description incorrect based om a misidenü fied cylindrical branching specimen belonging to Echinoclathria subhispida. Whitelegge (1901) redescribed the species based on a clathrous specimen (No. 365, confirmed missing from AM collections), which he sub- sequently designated holotype of intermedia (Whitelegge, 1902a: 214). However, he also redescribed a second specimen for the species (Lendenfeld's No, 332), which unfortunately belongs to E. subhispida. Both Lendenfeld's and Whitelegge's actions have created confusions, and, moreover, Lendenfeld's manuscript names and numbers on slides and specimens in the AM and BMNH do not completely correlate with the puhlished descriptions (see remarks for E. sub- hispida). | accept E. intermedia in the sense of Hallmann (1912), and designate a neotype for this species from the material described above, the most "typical being QMG300025 from Mooloolaba, SE. Qld. 25um REVISION OF MICROCIONIDAE s FIG. 288. Echinochalina (Echinochalina) reticulata Whitelegge (holotype AMZ950). A, Principal style (echinat- ing fibres). B, Auxiliary subtylostyle/ tornostyle (coring fibres and interstitial). C, Section through peripheral skeleton. D, Australian distribution. E, Neotype. 532 Echinochalina (Echinochalina) reticulata Whitelegge, 1907 (Figs 288-289) Echinochalina reticulata Whitelegge, 1907: 506, pl.45, fig.25; Hallmann, 1912: 289-290, pl.30, fig.2, text-fig.66 (et var.); Hooper & Wiedenmayer, 1994: 278. Tablis reticulata; de Laubenfels, 1936a: 76. MATERIAL. LECTOTYPE: AMZ950: Off Wollon- gong, NSW, 34?26'S, 150?53'E, 110-112m depth, coll. FIV *Thetis' (dredge). PARALECTOTYPES: AMZ715: same locality. AMZ23 (dry): E. coast of Flinders I., 40°01°S, 148°02’E, 29.vii.1909, coll. FIV ‘Endeavour’ (dredge; 'cotype' of var. crassa). AME646 (dry): same locality (‘cotype’ of var. crassa). (Other material presently missing: AMZ153, 716, 717). HABITAT DISTRIBUTION, Soft substrata; 110- 112m depth; Wollongong (NSW); Flinders I. (Tas) (Fig. 288D). DESCRIPTION. Shape. Massive, rounded, globular, lobo-digitate to thickly flabellate, up to 240mm long, 255mm wide, 150mm thick, with honeycomb reticulate construction produced by closely anastomosing, very thin membraneous, lamellate, apically pointed, fibre bundles. Colour. Brown to grey-brown in ethanol. Oscules. Large, up to 5mm diameter, scattered within lacunae of surface honeycomb reticula- tion. Texture and surface characteristics. Firm, flexible, harsh in dry state; surface honeycomb reticulate, lamellae very thin, membraneous and uneven. Ectosome and subectosome. Membraneous, tan- gentially disposed auxiliary megascleres lying below surface, in pauci- or multispicular tracts, pierced by sparse plumose bundles of auxiliary spicules from ascending choanosomal tracts. Choanosome. Irregularly reticulate skeletal structure, verging on subisodictyal, with differen- tiated primary and secondary spongin fibre sys- tems; fibres relatively light, mostly flattened, meandering or sinuous; primary fibres, 65- 112,.m diameter, pauci- or multispicular, running longitudinally through lamellae, intersected at regular intervals and obtuse angles by secondary fibres, 22-40j.m diameter, mostly uncored, oc- casionally unispicular; fibres anastomoses form cavernous eliptical meshes, 175-570um diameter; fibres cored by auxiliary megascleres, moderately heavily echinated by principal spicules; mesohyl matrix heavy but only lightly pigmented, with numerous auxiliary megascleres MEMOIRS OF THE QUEENSLAND MUSEUM scattered between fibres; choanocyte chambers large, oval, 45-75jum diameter. Megascleres. Principal styles echinating fibres straight, thick, with smooth, tapering, rounded or slightly subtylote bases, fusiform points. Length 108-(139.8)-196um, width 6-(8.9)-11 pm. Auxiliary megascleres coring fibres straight, slightly curved or sinuous, thin, ranging from hastate subtylostyles to asymmetrical tornostyles or sometimes strongyles, with rounded or slightly subtylote bases, rounded or telescoped points. Length 146-(252.0)-336um, width 2.5-(4.1)- 5.5m. Microscleres absent. Larvae. Viviparous, small, oval parenchymella, 80-110j.m diameter, with heavy matrix. REMARKS. In growth form, particularly its flat- tened spongin fibres, the holotype resembles the Caribbean Pandaros acanthifolium (Duchassa- ing & Michelotti), whereas Hallmann's variety crassa is more reminiscent of honeycomb reticu- late species of Holopsamma. However, in spiculation and skeletal architecture both morphs are identical and conspecific. Pandaros and Echinochalina differ substantially in their spicule geometries, whereby Pandaros has smooth or acanthose principal styles coring and echinating fibres, equivalent to those of Holopsamma, E. reticulata has auxiliary styles coring fibres and principal styles echinating fibres, typical of the Echinochalina condition. The species differs from other Echinochalina in spicule geometry, particularly the endings of auxiliary subtylos- tyles, spicule size, possession of flattened fibres, and a nearly isodictyal skeletal architecture, whereas in growth form it is similar to E. (E.) barba. Echinochalina (Echinochalina) ridleyi (Dendy, 1896) (Figs 290-291) Echinodictyum ridleyi Dendy, 1896: 44-45. ? Clathrissa or Stylotellopsis ridleyi; Hallmann, 1912: 151. Echinochalina ridleyi; Hooper, 1991: 1348; Hooper & Wiedenmayer, 1994: 278. MATERIAL. LECTOTYPE: NMVG2409: Port Phil- lip Bay, Vic, 38?09' S, 144°52’E, 40m depth, coll. J.B. Wilson (dredge). PARALECTOTY PES: NMVG2410: same locality, 38m depth. BMNH1902.10.18.363: same locality, 38m depth. BMNH1902.10.18.364, 57: same locality, 40m depth. HABITAT DISTRIBUTION. Substrate unknown; 38- 40m depth; Port Phillip (Vic) (Fig. 290D). REVISION OF MICROCIONIDAE 533 FIG. 289. Echinochalina (Echinochalina) reticulata Whitelegge (holotype AMZ950), A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal spicule. E, Auxiliary subtylostyle/ tornostyle. F, Ends of auxiliary spicule. DESCRIPTION. Shape. Stalked, digitate, club- composed of more-or-less flattened, fused lamel- shaped, 85-165mm long, 50-80mm maximum width, with thick cylindrical stalk, 18-32mm long, 12-17mm diameter, bulbous digitate apex flattened flabelliform or claviform growth. lae, up to 18mm wide, together producing bushy, 534 MEMOIRS OF THE QUEENSLAND MUSEUM C XS N SN RN N - NS : | Y WA EN J y 500 um FIG. 290. Echinochalina (Echinochalina) ridleyi (Dendy) (lectotype NMVG2409). A, Principal acanthostyle (echinating fibres). B, Auxiliary tornostyle/ oxeote (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. REVISION OF MICROCIONIDAE FIG. 291. Echinochalina (Echinochalina) ridleyi (Dendy) (lectotype NMVG2409). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal acanthostyle. D, Ends of principal spicule. E, Auxiliary tornostyle/ oxeote. F, Ends of auxiliary spicule. 336 Colour. Reportedly pale red-brown to red-orange in life, pale brown in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, com- pressible, flexible; surface optically smooth with distinct skin-like dermis. Ectosome and subectosome. Membraneous, microscopically hispid with multispicular, erect and paratangential tracts of auxiliary spicules. singly or in sparse plumose brushes, arising from peripheral choanosomal fibres and protruding through surface, choanosomal fibres immedi- ately below ectasome. Choanosome. Plumo-reticulate skeletal struc- ture, with incompletely differentiated, meander- ing, primary and secondary spongin fibres; primary fibres heavy, 52-116jm diameter, vaguely ascending, multispicular, spicules ar- ranged in loose whispy tracts occupying most of fibre diameter, becoming increasingly plumose towards periphery; secondary, connecting fibres, 18-43,.m diameter, pauci-, uni- or aspicular. producing cavernous, oval or elongate meshes, 75-365um diameter; fibres cored by auxiliary megascleres, sparsely echinated by principal spicules; mesohyl matrix heavy but only lightly pigmented, with numerous auxiliary megascleres dispersed between fibres; choanocyte chambers large, oval to elongate, 90-120p.m diameter. Megascleres. Principal acanthostyles echinating fibres small, straight, relatively thick, with taper- ing rounded bases. spined apical and basal ex- tremities, fusiform points; spines small, conical, on Length 87-(104.1)- I1 6m, width 5-(5.8)- p.m. Auxiliary spicules coring fibres long, thin, straight, slightly curved or sinuous, sometimes raphidiform, with variable terminations ranging from hastate oxeas, tornotes, quasi-monactinal tornostyles, or suhtylostyles. Length 184- (261.3)-314 um, width 2-(4.1)-6.5jum. Microscleres Absent. REMARKS. Dendy (1896) initially assigned this specta to Echinodictyum (Raspailiidae) having iactinal coring. monactinal echinating megascleres, extra-axial styles, and a more-or- less reticulate architecture. However, the spicules which actually core these fibres range from true diactinal to true monactinal forms, without ob- vious axial and extra-axial differentiation in their distribution. The skeletal architecture verges on plumose, not prominently reticulate as is found in most Echinodictyum species, and fibres are much heavier than those noremally found in species of MEMOIRS OF THE QUEENSLAND MUSEUM that genus (see Heoper, 1991). Includmg this species in Echinochalina requires broadening the generic definition to allow for the inclusion of spined echinating styles, but these spicules ap- pear to be true principal megascleres (as opposed to a special category of echinating acanthostyles such as those found in Clathria, for example). In any case, Simpson (1968a) and others have shown that acanthose verses smooth megascleres may be of minor consequence at higher sys- tematic levels in Microcionidae, and in other respects the species fits well with the present concept of Echinochalina. The species has closest affinities 10. E. spongiosa, also having acanthose principal spicules, though differing substantially in growth form, spicule geometry and spicule dimensions. Echinochalina (Echinochalina) spongiosa (Dendy, 1896) (Figs 292-293) Echinodictyum spongiosum Dendy, 1896: 45; Hallmann, 1912; 151: Echinachalina spongiosum; Hooper, 1991. 1348; Hooper & Wiedenmayer, 1994: 278. MATERIAL. LECTOTYPE: NMVG2432: Sorrento Jetty, Port Phillip Bay, Vic, 38°21°S, 144°42°E, 20m depth, coll, J.B. Wilson (dredge), PARALEC- TOTY PE: BMNH1902.10.18.58, 365: same locality. HABITAT DISTRIBUTION. Substrate unknown; 12m depth; Port Phillip ( Vic) (Fig. 292D). DESCRIPTION. Shape. Thickly encrusting, massive, subspherical sponge, 65mm diameter, attached directly to substrate. Colour. Reportedly grey in life, pale brown in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, com- pressible, spongy: surface shaggy, covered wilh large, mostly irregular, lobate papillae, low ridges and abundant striations. Ectosome and subectosome. Membrancous, lightly arenaceous ectosome, with irregular plumose brushes of auxiliary megascleres barely protruding through surface arising from peripheral choanosomal fibres. Choanosome, Plumo-reticulate skeletal struc- ture, verging on plumose-dendritic in periphery, without any obvious differentiation into primary or secondary fibres or tracts; fibres at core of skeleton light, whispy, multispicular, sinuous, reticulate, 22-54jum diameter, interconnected at irregular intervals by smaller multispicular fibres, 25um 537 REVISION OF MICROCIONIDAE FIG. 292. Echinochalina (Echinochalina) spongiosa (Dendy) (lectotype NMVG2452). A, Principal acanthos- tyles (echinating fibres). B, Auxiliary tornostyles (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. 538 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 293. Echinochalina (Echinochalina) spongiosa (Dendy) (lectotype NMVG2452). A, Choanosomal skeleton. B, Fibre characteristics (x475). C, Principal acanthostyle. D, Ends of principal spicule. E, Auxiliary tornostyle. F, Ends of auxiliary spicule. 8-18m diameter, forming irregular ovoid, rec- — paucispicular towards periphery, terminating on tangular or elongate meshes, 65-118pm or near surface as single spicules, skeletal tracts diameter; fibres increasingly plumose, more in periphery occupy most of fibre diameter, rang- REVISION OF MICROCIONIDAE 539 50 um FIG. 294. Echinochalina (Echinochalina) tubulosa (Hallmann) (lectotype BMNH1925.11.1.568). A, Principal styles (echinating fibres). B, Auxiliary styles (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Paratype. F, SAMTS4018. ing from 12-35jum diameter, fibre meshes cav- ernous, elongate, 240-6604.m diameter; all fibres cored by auxiliary megascleres, sparsely echinated by principal spicules; mesohyl matrix very heavy, dark yellow-orange pigmented, with moderate numbers of auxiliary spicules scattered between fibres; choanocyte chambers small, oval, 15-544.m diameter. Megascleres. Principal styles echinating fibres straight, relatively thin, subtylote, with light 540 spines concentrated mainly on base and near points, fusiform, bare points. Length 82-(92.3)- 98y.m, width 2.5-(4.4)-5.5j.m. Auxiliary spicules coring fibres thin, mostly straight, sometimes slightly curved or sometimes sinuous, varying from tornostyles, strongyles or tornotes with rounded or subtylote extremities. Length 166-(187.6)-214j.m, width 1.5-(3.2)- 4.2um. Microscleres absent. REMARKS. Echinochalina (E.) spongiosum was originally assigned to Echinodictyum, like E. (E.) ridleyi, having diactinal coring and monactinal echinating spicules, but E. (E.) spongiosum was even more atypical of Echinodictyum in its skele- tal architecture, having an almost completely plumose skeleton, with only vestiges of reticulate construction. The species is easily assigned to Echinochalina even though it has acanthose (ver- sus completely smooth) principal styles echinat- ing fibres (see remarks for E. (E.) ridleyi), differing from E. (E.) ridleyi in growth form, spicule geometry and spicule dimensions. Echinochalina (Echinochalina) tubulosa (Hallmann, 1912) (Figs 294-295, Plate 12C) Ophlitaspongia tubulosa Hallmann, 1912: 272-275, pl.35, fig.3, text-fig.60. Echinochalina tubulosa; de Laubenfels, 1936a: 119; Hooper & Wiedenmayer, 1994: 278. Echinoclathria tubulosa; Wiedenmayer, 1989: 66, pl.6, fig.9, pl.25, figs 3-4, text-fig.45. MATERIAL. LECTOTYPE: BMNH1925.11.1.568: Westernport Bay, Vic, 38?26'S, 145°08’E, coll. J. Gabriel (dredge). PARALECTOTYPES: AME1271 (dry): Precise locality unknown, Illawarra region, NSW. AMZI55 (dry): same locality. OTHER MATERIAL: QLD- QMG304737, QMGL864 (frag- ment NTMZ1528), QMGL2759 (fragment NTMZ1566). TAS- QMG300265 (NCIQ66C-3558-J) (fragment NTMZ3783). S AUST- SAMTS4018 (frag- ments NTMZ1606, QMG300475). HABITAT DISTRIBUTION. Sand, rock and coral rubble substrata; 15-32m depth; Turtle Is, Agincourt Reefs (FNQ); Moreton I. (SEQ); Il- lawarra (NSW); Westernport Bay (Vic); Kent Is, Bass Strait (Tas); Kangaroo I. (SA) (Fig. 294D). DESCRIPTION. Shape. Tubulo-digitate, mas- sive, erect, up to 120mm high, attached directly to substrate, with erect, regularly cylindrical or irregular, stoloniferous, thin or thick, bulbous MEMOIRS OF THE QUEENSLAND MUSEUM digits tapering towards their extremities, up to 45mm high, 22mm diameter. Colour. Bright orange, yellow-orange, or ver- milion-red alive (Munsell 10R 6/10 - 2.5R 5/10), pale brown in ethanol. Oscules. Large, up to 10mm diameter, at apex or subapical on each digit. Texture and surface characteristics. Firm, rub- bery, compressible, flexible digits; surface even, porous, microscopically rugose fibre bundles. Ectosome and subectosome. Membraneous, with dense paratangential layer of auxiliary styles lying below, occasionally protruding through sur- face, in irregular plumose formations, with or without light arenaceous ectosomal layer; choanosomal fibres immediately below ec- tosome. Choanosome. Regularly isodictyal to irregularly reticulate skeletal architecture, with poorly dif- ferentiated primary and secondary fibres; primary ascending fibres thin, 42-93ym diameter, heavy, paucispicular, sometimes uni- or aspicular, occasionally meandering, sinuous, often ascending directly to periphery, intercon- nected at irregular intervals by thin, more-or-less transverse, uni- or aspicular secondary fibres, 18-35j.m diameter; secondary fibres branch and anastomose amongst themselves, forming oval, rectangular or triangular, relatively cavernous meshes, 124-650j.m diameter; fibres cored by predominantly longer and thinner auxiliary megascleres, never forming more than paucispicular tracts, usually becoming plumose towards periphery; fibres echinated by shorter, stouter principal spicules; mesohyl matrix heavy but only lightly pigmented, with numerous thin auxiliary spicules dispersed between fibres and occasionally also small quantities of inorganic debris; choanocyte chambers large, oval, 72- 113j.m diameter. Megascleres. Principal styles echinating fibres styles straight, short, relatively thick, with smooth, evenly rounded bases or less frequently slightly subtylote bases, almost hastate points. Length 74-(96.5)-1284.m, width 4-(6.6)-8.5 p.m. Auxiliary styles coring fibres and interstitial long, thin, straight or slightly curved hastate styles, occasionally modified to asymmetrical styloids, tornotes or strongyles, rarely sinuous. Length 108-(226.1)-305um, width 1-(2.2)- 4.5j.m. Microscleres absent. REVISION OF MICROCIONIDAE 541 FIG. 295. Echinochalina (Echinochalina) tubulosa (Hallmann) (QMG300265). A, Choanosomal skeleton. B, Fibre characteristics (x498). C, Principal styles. D, Ends of principal spicule. E, Auxiliary style. F, Ends of auxiliary spicule. Larvae. Incubated parenchymella larvae in 1 Associations. One specimen contained numerous specimen, oval, small 62-170j.m diameter, con- algal filaments within fibres, almost completely taining abundant collagen but no larval spicules. obscuring spicules within fibre skeleton; Wieden- 542 100 urn MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 296. Echinochalina (Protophlitaspongia) bispiculata (Dendy) (lectotype NMVG2319). A, Auxiliary oxeas (coring fibres and interstitial). B, Principal oxea (echinating fibres). C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. F, Fibre skeletal structure. mayer (1989) also noted microsymbionts in his material from Bass Strait. REMARKS. This species shows some variability, particularly in organisation of skeletal structure (ranging from regularly isodictyal fibre network to irregularly reticulate), palmate isochelae were abundant in 1 specimen from SA (but presumed to be contaminants), and position of oscules (terminally or subtermally on digits), otherwise there is no doubt that all these specimens belong to E. (E.) tubulosa. The longer, thinner styles coring fibres and shorter, thicker styles echinating fibres indicate that it belongs to Echinochalina. Interpretation of these features is of primary importance in generic placement. This species has been included with Ophlitaspongia (=Echinoclathria) (Hallmann, 1912), or Echinoclathria (=Holopsamma) (Wiedenmayer, 1989), based on misinterpretation of these genera and that megascleres coring and echinating fibres were geometrically identical. However, it is clear that longer, thinner auxiliary styles are most com- monly found inside fibres, whereas shorter, thick- er principal styles are most frequently found REVISION OF MICROCIONIDAE echinating fibres, even though few intermediate examples of both spicule types can be found coring or echinating fibres. Contrary to Hallmann (1912: 274), I infer that spicules coring fibres in this species are equivalent to auxiliary megascleres of the Microcionidae, and the thick- er echinating styles are derived from principal spicules, and thus the most appropriate placement is with Echinochalina. Hallmann (1912) remarked on the resemblance and possible close relationship between this species and £ (Protophlitaspongia) bispiculata, although spicule geometry is very different. OTHER SPECIES OF ECHINOCHALINA (ECHINOCHALINA) Echinochalina (Echinochalina) isochelifera (Uriz, 1988) Echinoclathria isochelifera Uriz, 1988: 89-90, pls 22b, 42c, 36b-d, text-fig.64 [Namibia]. MATERIAL. HOLOTYPE; ABIP7B-GA, SW. Africa. Echinochalina (Echinochalina) melana Van Soest & Stentoff, 1988 Echinochalina sp, Hartman in Lewis, 1965; in table. Echinochalina melana Van Soest & Stentoff, 1988; 123-125, pl.12, fig.1, text-fig. 60 [Barbados, West Indies]. MATERIAL. HOLOTYPE: ZMAPORS509, Caribbean. Echinochalina (Protophlitaspongia) Burton, 1934 Protophlitaspongia Burton, 19343: 562, [Not Protophlitaspongia; de Laubenfels, 1954: 96; Pulitzer-Final, 1986: 138]. TYPE SPECIES. Siphonochalina bispiculara Dendy, 1895: 246 (by original designation). DEFINITION. Diactinal or quasi-monactinal auxiliary megascleres core fibres. with diactinal or quasi-monactinal principal spicules echinating fibres, REMARKS. In this subgenus structural megascleres (conng and echinating fibres) ap- pear to be diactinal, unlike all other microcionids, but they are interpzeted here as being highly modified monactinal spicules allowing its in- clusion in the Microcionidae. This interpretation is supported by the more-or-less plumose as- cending, primary spicule tracts, true echinating megascleres, isochelae and toxa microscleres in several species, and obvious (i.e., less modified) monactinal ectosomal spicules in several species, indicating affinities with Echinochalina. The 543 coring and echinaling megascleres are equated here with auxiliary and principal spicules, respec- tively, of typical Echinochalina. Dendy (1896) included the type species in the Haplosclerida, but remarked on its unique spicule arrangement, particularly the ectosomal structure and fibre echination. Burton (19343) assigned the type to Microcionidae, for similar reasons as those oul- lined above, whereas de Laubenfels (1936a] referred jt to the Desmacididae because, he sug- gested, the hastate diactinal megascleres resembled those of Guitarra and Liosina, al- though it lacked poecilosclerid microscleres. Echinochalina ( P.) bispiculata is included in this subgenus since it has a paratangential ectosomal skeleton composed of auxiliary megascleres, reticulate spongin fibres and echinating principal spicules, whereas other species described by de Laubenfels (19362) (P. aga, P. ada and P. antil- lana) are more appropriately. placed in Des- macididae and Haplosclerida (see remarks for Protophlitaspongia under Genera Included). Eight species are now included in the subgenus, 2 known exclusively from New Caledonia and the remainder from eastern Australia, (Table 46; Hooper & Lévi, 1993a: 1279). Echinochalina (Protophlitaspongia) bispiculata (Dendy, 1895) (Figs 296-297, Table 45} Siphonochalina bispiculata Dendy,1895; 244. Diplodermia bispiculata; Hallmann, 1912; 255. Protophlitaspongia bispiculata; Burton, 1934a: 562. Echinochalina bispiculara; Hooper & Lévi, 1993a: 1279; Hooper & Wiedenmayer, 1994; 277. MATERIAL. LECTOTYPE: NMVG2319: 5-6km from Point Lonsdale, Port Phillip Heads, Vie, 38°20.5'S, 144°35.6'E, 34-38m depth, 1894, coll. J.B. Wilson (dredge). PARALECTOTYPE: NMVG232( (fragment BMNH1902,10.18.14.110): same locality, OTHER MATERIAL: VIC - QMG304102, HABITAT DISTRIBUTION. Rock reef; 34-38m depth; Port Phillip (Vic) (Frg. 296D). DESCRIPTION. Shape. Massive, subspherical, imegularly lobate, 9Üümm maximum diameter, with short bulbous surface lobes, or subcylindri- cal, flattened digitate sponge. 160mm long, 28mm wide, | 5mm thick. Colour. Pale yellow-brown in ethanol. Oscules. Small, 2-3mm diameter, scattered over surface, particularly on apex of surface lobes. 544 Texture and surface char- acteristics. Soft, compres- sible, slightly rubbery; surface optically even, minutely reticulate. Ectosome and subec- tosome. Microscopically hispid ectosome with scat- tered plumose brushes of long, thin auxiliary oxeas, arising from the points of peripheral fibres and paratangential to surface; choanosomal fibres imme- diately below surface al- though spicule tracts more plumose in peripheral skeleton than at core. Choanosome. Regularly reticulate to subisodictyal skeletal structure, with well developed fibres divided into primary and secondary elements; primary fibres ascending, multispicular, 60-95p.m diameter, form- ing distinctly plumose tracts within core of fibre, interconnected at more-or- less regular intervals by slightly thinner secondary fibres, transverse, uni-, pauci- or aspicular fibres, 20-43pm diameter; fibres relatively light, cored by long, thin auxiliary oxeas and echinated by short, stout principal oxeas; fibre anastomoses form cavern- ous ovoid meshes, 180- 540p.m diameter; mesohyl matrix heavy but only light- ly pigmented, with moderate quantities of auxiliary spicules dispersed between fibres; choanocyte chambers large, oval to elongate, 60- 130jum diameter. Megascleres (Table 45). Principal megascleres echinating fibres straight, short, thick, invariably smooth, hastate oxeas, with abruptly pointed or telescoped ends. Auxiliary megascleres coring fibres long, thin, FIG. 297. straight, hastate or rarely fusiform oxeas. Microscleres absent. Echinochalina MEMOIRS OF THE QUEENSLAND MUSEUM (Protophlitaspongia) bispiculata (Dendy) (QMG304102). A, Choanosomal skeleton. B, Fibre characteristics. C, Auxiliary oxea. D, Principal oxea. E, Ends of oxeas. Larvae. Incubated parenchymella larvae small spherical, with heavy mesohyl matrix, 210- 240m diameter, no larval megascleres. REMARKS. Of all E. (Protophlitaspongia) this species is the most reminiscent of Haplosclerida in skeletal characteristics (e.g., three dimensional ectosomal architecture like Hemigellius (Niphat- idae); choanosomal fibre construction similar to Callyspongia (Callyspongiidae)). Echinochalina (P.) bispiculata is slightly similar to E. (P.) oxeata but that species lacks true geometric differentia- tion between coring and echinating spicules and has a different growth form (Table 46). REVISION OF MICROCIONIDAE 545 Iri Li * | 50 um FIG. 298. Echinochalina (Protophlitaspongia) collata sp.nov. (holotype QMG304120). A, Principal oxea (echinating fibres). B, Auxiliary oxeas (coring fibres and interstitial). C, Ectosomal subtylostyle/ anisoxea. D, Wing-shaped toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype. Echinochalina (Protophlitaspongia) collata HABITAT DISTRIBUTION. Coral pinnacles; 9m depth; Lizard I. (FNQ) (Fig. 298F). sp. nov. Figs 298-299, Table 45, Plate 12D . i (Hig ) DESCRIPTION. Shape. Thickly encrusting, up MATERIAL. HOLOTYPE: QMG304120: Blue 19. 12mm thick, bulbous in places on surface Lagoon, Lizard I., Cairns section, Great Barrier Reef, following contours of substrate. f Qld, 14°41.0'S, 145?27.5' E, 9m depth, 3.iv.1994, coll. Colour. Dark red alive (Munsell 2.5R 4/10), light J.N.A. Hooper & S.Cook (SCUBA). brown in ethanol. 546 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 299. Echinochalina (Protophlitaspongia) collata sp.nov. (holotype QMG304120). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Ends of auxiliary anisoxea and style. E, Ends of principal oxea. F-G, Ends of ectosomal anisoxea and style. H, Wing-shaped toxas. REVISION OF MICROCIONIDAE 547 TABLE 45. Comparison in spicule dimensions between species of Echinochalina (Protophlitaspongia). Meas- urements in micrometres (N=25); comparative data from Hooper & Lévi (1993). EAP.) labourei EAP.) bargibantr Hooper & Hooper & favulosa Lévi Lévi 5p.noV. (QMG300685) (QMG301270) (OMGL2166) (New (New (SE Qld) Caledonia) Caledonia) 52-(75.1)- | 55-(73.3)-98 108 x 1.8- x 0.8-(1.9)- (2.9)-4.0 3.0 1134156.2)- | 144-(216.1)- auxiliary 194 x 1.0- 278 x 1.0- styles (L7)-2:5 (24-35 Principa 28-(43.5)-42 | 32-(46.7).58 (echinating) »2.0-2.5)- | a L0-(2.0)- cou mM 0 25 eum [Chele — | 14- | 13-05.53-21 | 5)21 E(P.) 101-(121.2)- J66 x 3.5. (6.1)-7.0 | Auxiliary | (coring) | oxeas Ectosomal 57(79.7)- 12244,5- prom 9-(10. ETT EAP.) isuuci 232-(265.2)- 287 x 3-(3.6)- 212-(234,0)- 262 x 2-(2.8)- ] T8-(142,7)- 162x rh Ay absent | absent | nt E. tuberosa p.nov, (digitale morph) E. tuberosa sp.nov. (typical EAP.) oxeata (Burton) E. collata (SE. Qld.) 128-(132)164 Ups 102-(123.9}- x 35(42)70 [CC [158x 3-446)-7 212-(237.2)- | 14-(187.7)- 264 X 1.5- wy S (2.2)-3 94-(148,0)- 1B8 x | 5- (4. mM 6 (SE. Qld.) 94-(101 7)- 163 x 15- (G.8)-7,5 j42-(181,9)- 228 x 1.5- (2.2)3 54-(71.2)-84 Pu x2-(2. cL | absent | 14-(73,2)- m -re x Ü.5-(0.9} x 0.5-{1.1}- 2 ; . x» Tiai nm | ss Tees ae a = 2 ats 15 X? Oscules. Very small, 1-2mm diameter, on apex of larger surface bulbs, surrounded by radiating sub- ectosomal drainage canals. Texture and surface characteristics. Hàrsh, firm, compressible; surface microconulose, minuiely shaggy, conules close-set, Jess than 2mm high, surface silty in situ with radiating drainage canals clear of silt, Ectosome ana subeciosome. Slightly plumose, multispicular brushes of auxiliary spicules from primary choanosomal tracts protrude through surface mainly on ends of conules; bundles of ectosomal auxiliary styles tangential to surface; abundant detritus and collagen in ectosomal layer. Choanosome. Irregularly reticulate skeletal structure, very heavy spongin fibres relatively homogeneous throughout skeleton, 45-120. diameter, forming oval or elongate meshes, 120- 440jum diameter, without marked difference in mesh size between periphery and core of skeleton; ascending primary fibres cored by mul- tispicular tracts of auxiliary spicules but occupy- ing only about 50% of fibre diameter, interconnected by uni- or paucispicular secon- dary, transverse fibres; fibres sparsely echinated by thinner oxeas (? principal spicules) protruding at acute angles through fibres; mesohyl matrix heavy, with moderate numbers of auxiliary megascleres and toxa microscleres dispersed be- tween fibres; choanocyle chambers large, elon- gate, 30-60j.m diameter. Megascleres (refer to Table 45 for dimensions). Principal (7) spicules echinating fibres shon oxeis, thin, straight or slightly curved at centre, with telescoped points. Auxiliary spicules coring fibres vary from true oxeas to true styles, with many intermediates, long, thick, straight or slightly curved at centre, points usually telescoped. Ectosomal auxiliary styles or anisoxeas long, thin, whispy, straight or slightly curved, with rounded or spiked base and fusiform or tele- scoped points. Microscleres (Table 45). Toxas intermediate be- tween oxhem and wing-shaped, thin, slightly curved at centre, slightly reflexed points. ETYMOLOGY Latin collatus (L.), extended, diffuse: for the growth Form. REMARKS, This species is unusual in its thickly encrusting growth form, conulose surface, multi- spicular tracts occupying only about 50% of fibre diameter, prevalence of telescoped points and asymmetrical spicules. It is most similar to, and probably a sibling species of, E. (F) tuberosa in having a special category of ectosomal styles and oxhorn toxas, although spicule geometry and spicule sizes are substantially different berween these species. Further comparisons between species of E (Protophlitaspongia) are given in Table 46. 548 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 300. Echinochalina (Protophlitaspongia) favulosa sp.nov. (holotype QMGL2166). A, Principal oxea and style (echinating fibres). B, Auxiliary oxea and style (coring fibres and interstitial). C, Palmate isochelae. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. Echinochalina (Protophlitaspongia) favulosa sp. nov. (Figs 300-301, Table 45) MATERIAL. HOLOTYPE: QMGL2166: Off Noosa Heads, Qld, 26?31'S, 153?48'E, 480m depth, 13.ix.1980, coll. QFS Craigman Survey (trawl). HABITAT DISTRIBUTION. Sand and shell grit sub- strata; 480m depth; Noosa region (SEQ) (Fig. 300E). DESCRIPTION. Shape. irregularly branching, cylindrical sponge, 195mm long, up to 33mm diameter, without apparent basal attachment; branches bifurcate few times. Colour. Live colouration unknown, grey-brown ethanol. Oscules. Large, up to 5mm diameter, on lateral margins and ends of branches. REVISION OF MICROCIONIDAE 549 FIG. 301. Echinochalina (Protophlitaspongia) favulosa sp.nov. (holotype QMGL2166). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal oxea and style. D, Ends of principal spicules. E, Auxiliary style and oxea. F, Ends of auxiliary spicules. G, Palmate isochela. Texture and surface characteristics. Soft, com- pressible; surface ‘honeycomb’ reticulate, highly porous, uneven, conulose. Ectosome and subectosome. Membraneous, minutely conulose surface with erect, pauctspicular brushes of auxiliary megascleres ansing from choanosomal spicule tracts protrud- ing into surface conules but rarely through sur- face; some detritus scattered over surface; choanosomal fibres immediately below ec- tosomal skeleton. Choanosome. Regularly reticulate, subisodictyal skeletal archilecture composed of more-or-less ascending, mühi- or paucispicular, primary fibres, 22-43,.m diameter, interconnected at ir- regular intervals by transverse. uni-, pauci- or aspicular fibres, 14-25 um diameter; fibres cored hy longer auxiliary megascleres and moderately echinated by shorter, principal spicules; fibre anastomoses form relatively wide, clongate or hexagonal meshes, 92-2361.m diameter; mesohyl matrix heavy, relatively darkly pigmented, granular, with auxiliary and principal megascleres dispersed between fibres; choanocyte chambers small ovoid to elongate, 22-46 .m diameter. Megascleres (Table 45). Principal megascleres echinating fibres relatively short, thick, straight, hastate oxeas, sometimes slightly telescoped, oc- casionally quasi-monactinal (asymmetrical ends), rarely styles, only marginally shorter than coring spicules. Auxiliary spicules coring fibres long, slender, straight, hastate oxeas, often with telescoped points, occasionally modified to quasi-monac- tinal forms, rarely tme styles, longer forms generally thinner. Mierascleres (Table 45). Palmate isochelae abun- dant, small. with approximately 15% twisted forms. lateral alae completely fused to shafi, front ala partially detached from lateral akie, shalt straight: Toxas absent, ETYMOLOGY, For its honeycomb reticulate growth form, REMARKS. This species is unusual for its Holopsamma-like honeycomb reticulate growth form, having palmate isochelae, both principal and auxiliary spicules ranging in geometry from (predominantly) oxeas to styles. with asymmetri- cal intermediate geometries, and corimg and echinating spicules differing only marginally iri their length and thickness. The possession of isochelae microscleres in the present species sup- MEMOIRS OF THE QUEENSLAND MUSEUM ports the inclusion of the genus in the Poecilosclenda, although it is still feasible that this species, and Protophlitaspongia, may be more appropriately assigned to the poecilosclerid Desmacididae (de Laubenfels (1936a) and fol- lowed by Pulitzer-Finali (1986)), but little sup- port for its inclusion in the haplosclerid Niphatidae (as proposed for Isodictya by Hajdu et al., 19945). Echinochalina (Protophlitaspongia) isaaci sp. nov. (Figs 302-303, Table 45, Plate 12E) MATERIAL. HOLOTYPE: QMG305051: W side Oyster Reel, Cairns section, Great Barrier Reef, Qld, 16°38.4S 145°54.7°E 20m depth, 21.41.1995, coll. J.N.A. Hooper (SCUBA). PARATYPE; QMG 305464: W: side Gannett Cay, Swain Reef, Mackay section, Great Barrier Reef, Qld., 21*58.68' S, 152°28.34°E, 22m depth, 24.vii. 1995, coll. J. N.A. Hooper & P. Tüm- kins (SCUBA),OTHER MATERIAL. QLD- QMG305398, QMG305430, QMG305504. HABITAT DISTRIBUTION. Fringing coral reefs, coral pinnacles, outer reef slopes, on dead coral, 20- 33m depth; Oyster Reef (FNQ); Swain Reefs (MEQ) (Fig. 302E). DESCRIPTION. Shape. Arborescent, digitale, branching. up to 150mm high, with conical cylindrical branches, up to 80mm long, 15mm diameter, hifurcate at tips and expanded towards apex of branches in life, collapses producing flat- tened branches in ethanol; basal stalk upto 25mm long, 12mm diameter, with expanded basal at- tachment. Colour. Pale orange-brown alive (Munsell 5R 7/8), light brown in ethanol. Oscules. Large, up to 13mm diameter alive, on apex of each digit, surrounded by raised membraneous lip, collapses completely in ethanol leaving no visible trace of oscule on external surface. Texture and surface characteristics. Soft, spong i alive. firm, compressible, rubbery in ethanol; fleshy alive, with longitudinal striations and ridges running from terminal oscule along sides of digits, collapsing when preserved producing concave striations and pits on sides of digits. Ectosome and subectosome. Well developed, dis- crete brushes of ectosomal auxiliary subtylos- tyles arising from ends of ascending choanosomal tracts; ectosome more heavily col- lagenous than choanosome, lightly pigmented; subectosomal skeleton slightly cavernous, with widely spaced peripheral fibres and skeletal tracts REVISION OF MICROCIONIDAE 551 AW 77 WE 7 4 FIG. 302. Echinochalina (Protophlitaspongia) isaaci sp.nov. (holotype QMG305051). A, Principal oxea (echinating fibres). B, Auxiliary oxea (coring fibres and interstitial). C, Ectosomal auxiliary subtylostyles. D, Section through peripheral skeleton. E, Australian distribution. F, Holotype. 552 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 303. Echinochalina (Protophlitaspongia) isaaci sp.nov. (paratype QMG305464). A, Choanosomal skeleton. B, Fibre characteristics. C, Point of principal oxea. D, Point of auxiliary oxea. E, Ends of auxiliary anisoxea. becoming progressively more plumose towards periphery. Choanosome. lrregularly reticulate, wise- meshed fibre skeleton, cavernous choanosomal structure; fibres relatively homogeneous in size and distribution, 40-70.m diameter, without any marked differences between primary and secon- dary fibres; fibres light; fibre meshes wide, oval to elongate, up to 650j.m diameter, slightly more cavernous in peripheral skeleton than at core; REVISION OF MICROCIONIDAE ascending primary fibres cored by pauci- or mul- tispicular tracts of auxiliary oxeas; smaller con- necting secondary fibres, predominantly transverse through skeleton, cored by unispicular tracts of auxiliary oxeas, occasionally aspicular; fibres at core of skeleton sparsely echinated by principal oxeas; mesohy] matrix light, virtually unpigmented, with sparse auxiliary oxeas scat- tered between fibres; choanocyte chambers large, oval. up to 6üium diameter. Megascleres (Table 45). Principal oxeas echinat- ing fibres short, slender, straight, usually with telescoped points. Auxiliary oxeas coring fibres long, slender, straight, usually with telescoped points. Ectosoma! auxiliary subtylostyles long. slender, straight, with slightly swollen bases, sometimes pointed at apex, with fusiform or slightly telescoped points. Microscleres. Absent. ETYMOLOGY. In memory of Isaac Cook, son of pepe Cook. collector of many of the samples studied TÉ. REMARKS. This species has a cavernous, ir- regularly reticulate skeletal structure. with rela- tively poorly developed fibre system and mostly dominated by more-or-less plumose tracts coring major fibres. The ectosomal skeleton is par- ticularly well developed, moreso than other species of E. (Protophliraspongia), whereas the echinating spicule skeleton is very sparse, nearly vestigial. The species is most closely related to £. {P} bargibanti in its growth form and aspects of its skeletal structure but they differ substantially in their respective live field characters, coring spicule skeleton, and spicule sizes. The species is compared further with other £. ( Prorophilitaspon- gia) m Table 46. Echinochalina (Protophlitaspongia) oxeata (Burton, 1934) (Fig. 304, Table 45) Protophlitaspongia oxeata Burton, 1934a: 562, text- fig.128. Echinochalina oxeata, Hooper & Lévi, 19933; 1275; Hooper & Wiedenmayer, 1994: 278. MATERIAL. HOLOTYPE: BMNH1930,8, 13.45: Papuan Pass, Cooktown region, Great Barrier Reel, Qld, 15°47'S, 145^48' E, 40m depth, coll. GBR Ex- pedition (dredge). HABITAT DISTRIBUTION. On foraminifera and coral fragments, 40m depth; Cooktown (FNQ) (Fig. 304C). 553 DESCRIPTION. Shape. Branching, cylindrical sponge 3-Smm diameter, now fragmented: branches bifurcate near ends. Colour. Light brown in ethanol. Oscules. Small, 1-2mm diameter, on lateral sides of branches. Texture and surface characteristics. Tough, elas- tic; surface smooth, even. perous. Ectosome and subectosome. Membraneous, with Sparse. more-or-less erect bundles of auxiliary oxeas from ascending choanosomal tracts protruding through surface; choanosomai fibres immediately below ectosome. Choanosome. Irregularly reticulate skeleton wilh differentiated primary and secondary spongin fibres; primary fibres ascending, pauci- or multi- spicular, 35-S5ym diameter, interconnected by transverse, uni- or aspicular secondary fibres, up a 224m diameter; fibre anastomoses produce large, cavernous, oval meshes in peripheral skeleton, up to S50um diameter, smaller, oval close-meshed reticulation at core of skeleton, up to [45m diameter; all spicule tracts occupy only small proportion of fibre diameter: differentiated echinating megascleres absent although auxiliary spicules may protrude through fibres at acute angles; mesohyl matrix heavy but only lightly pigmented, containing numerous auxiliary spicules scattered between fibres; choanecyle chambers large, oval 45-95y:m diameter. Megascleres (Table 45). Principal oxeas coring fibres short, slender, straight or slightly curved at centre, with hastale or slightly telescoped points, occasionally modified to styles. Microscleres absent. REMARKS. This species has reduced spicula- tion, lacking either echinating (principal) spicules or ectosomal styles. In spongin fibre structure. distnbution of spicules within fibres, ectosomal spiculation, it is close to E. (P.) bispiculata, Hooper & Lévi's (1993a: 1279) E. (P.) oxeata from Moreton Bay is not conspecific with the holotype of E. (P.) tuberosa, whereas E. (P) oxeata is restricted to the type specimen. Echinochalina (Protophlitaspongia) tuberosa sp. nov. typical growth form (Figs 305-306, Table 45 Plate 12F) MATERIAL, HOLOTYPE: QMG2300039: Inner Gneéerings Shoals, off Mooloolaba, SE. Qld, 26^38.58, 153709.5 E, 10m depth, I0.xii. 1991, coll. J.N.A Hooper & S.D. Cook (SCUBA). PARATYPE; QMG300082: Shag Rock, N. Stradbroke 1., Moreton 554 100 um MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 304. Echinochalina (Protophlitaspongia) oxeata (Burton) (holotype BMNH1930.13.45a). A, Auxiliary oxeas. B, Section through peripheral skeleton. C, Australian distribution, D, Fibre characteristics. E, Choanosomal skeleton. F, Holotype. Bay, SE. Qld, 27?25.0'S, 153?31.4'E, 15m depth, 05.11.1992, coll. J.N.A. Hooper & J. Wilkinson (SCUBA).OTHER MATERIAL: QLD- QMG304015. HABITAT DISTRIBUTION. Rock and coral reef, usual- ly sheltered under ledges; 9-25m depth; Stradbroke 1., Moreton Bay, Mooloolaba (SEQ) (Fig. 305F). DESCRIPTION. Shape. Thickly encrusting with bulbous-digitate surface projections forming lobate-digitate or bulbous sponge, up to 120mm long, 70mm wide, 55mm thick. Colour. Bright red to pale red alive (Munsell 2.5R 4-5/10), brown in ethanol. Oscules. Small, up to 3mm diameter, scattered at base of surface bulbs, between adjacent bulbs, each with prominent membraneous lip. Texture and surface characteristics. Firm, slight- ly hispid; surface uneven with prominent bulbous REVISION OF MICROCIONIDAE 555 FIG. 305. Echinochalina (Protophlitaspongia) tuberosa sp.nov., typical morph (holotype QMG300039). A, Principal oxeas (echinating fibres). B, Auxiliary oxeas (coring fibres and interstitial). C, Ectosomal auxiliary subtylostyle. D, Oxhorn toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype. 556 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 306. Echinochalina (Protophlitaspongia) tuberosa sp.nov., typical morph (holotype QMG300039). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal oxea. D, Base of principal spicule. E, Auxiliary oxea. F, Base of auxiliary spicule. G, Ectosomal auxiliary style. H, Ends of ectosomal spicule. I, Oxhorn toxa. protruberances, up to 9mm high, 5mm diameter, auxiliary subtylostyles perched on surface, over- inflated at ends. Ectosome and subectosome. Relatively thick paratangential brushes composed of ectosomal ascending choanosomal skeletal tracts; lying protruding principal oxeas from peripheral, REVISION OF MICROCIONIDAE choanosomal fibres immediately below ec- tosome. Choanosome. Regularly reticulate skeletal struc- ture, without any marked compression at core, with nearly renieroid fibre skeleton and plumo- reticulate spicule tracts diverging towards periphery; spongin fibres heavy, without notice- able size differences between primary or secon- dary fibres, 20-60j.m diameter; all fibres cored by auxiliary oxeas and moderately heavily echinated by principal oxeas, particularly at fibre nodes; primary fibres with ascending multi- spicular tracts of auxiliary oxeas terminating in sparse bundles at surface; secondary fibres pauci- or unispicular in peripheral skeleton, uni- or bispicular at core of skeleton; fibre anastomoses produce square or round, relatively even meshes throughout skeleton, 70-150pm diameter; mesohyl matrix moderately heavy, with few auxiliary spicules scattered between fibres; choanocyte chambers small, oval, 25-404m diameter. Megascleres (Table 45). Principal oxeas echinat- ing fibres short, thick, straight, with fusiform or slightly telescoped points. Auxiliary oxeas coring fibres, long, slender, straight or slightly curved at centre, with tapering fusiform or slightly telescoped points. Ectosomal auxiliary subtylostyles long, very slender, whispy, usually curved at centre, base subtylote, usually microspined, point hastate. Microscleres (refer to Table 45 for dimensions). Toxas oxhorn, short, thick, wide central curva- ture, points only slightly reflexed. Digitate growth form (Figs 307-308, Table 45) MATERIAL. QMG300030: Outer Gneerings Shoals, off Mooloolaba, SE. Qld, 26?39'S, 153?10'E, 25m depth, 10.xii.1991, coll. J.N.A. Hooper & S.D. Cook (SCUBA). DIAGNOSIS. Shape. Arborescent, tubulo- digitate growth form, 65mm high, 78mm wide, cylindrical or slightly flattened, repeatedly bifur- cate, occasionally anastomosing branches, 3- 5mm diameter, slightly swollen at extremities; sponge attached to substrate by expanded, short basal attachment. Colour. Same as typical form. Oscules. Small, up to 2mm diameter, on lateral sides of branches. Texture and surface characteristics. Firm, com- pressible, slightly rubbery; surface of branches smooth, macroscopically even, microscopically slightly hispid. Ectosome and subectosome. Thick, erect bundles of ectosomal auxiliary subtylostyles forming dis- crete plumose brushes on surface, usually at ends of ascending choanosomal spicule tracts, inter- mingled with protruding auxiliary coring oxeas; subectosomal region relatively thick, spiculose, with no fibre component. Choanosome. Regularly reticulate skeletal struc- ture without marked compression of axis. Mega- and Microscleres. As for typical form. ETYMOLOGY. For the bulbous encrusting form. REMARKS. There are several notable differen- ces between the bulbous-encrusting, shallower water (typical) morph and the deeper water, digitate morph that warrant their separate description above (i.e., growth form, ectosomal development), but there is little doubt that they are conspecific given similarities in their skeletal structure, fibre characteristics, spicule geometry, spicule dimensions. This species resembles E. (P) bispiculata in growth form (both bulbous, massive), fibre char- acteristics (regularly reticulate), and geometry of coring and echinating megascleres. However, E. (P.) tuberosa has a special category of auxiliary style forming a thick ectosomal skeleton (whereas E. (P.) bispiculata simply has protrud- ing auxiliary oxeas from the peripheral choanosomal fibre skeleton), toxa microscleres (versus no microscleres), and different spicule dimensions (see Table 46). The species is also related to E. (P.) laboutei, particularly the digitate morph, in having special auxiliary styles forming the ectosomal skeleton, and similar fibre charac- teristics with multispicular primary ascending fibres and uni-, pauci- or aspicular secondary fibres, whereas E. (P) laboutei lacks toxa microscleres, has a compressed central axial fibre skeleton and generally has aspicular fibres. OTHER SPECIES OF ECHINOCHALINA (PROTOPHLITASPONGIA) Echinochalina (Protophlitaspongia) bar- gibanti Hooper & Lévi, 1993 Echinochalina (Protophlitaspongia) bargibanti Hooper & Lévi, 1993a: 1280-1283, figs 31-32 |New Caledonia]. MATERIAL, HOLOTYPE: QMG301270. SW. Pacific. Echinochalina (Protophlitaspongia) laboutei Hooper & Lévi, 1993 MEMOIRS OF THE QUEENSLAND MUSEUM 558 2s \ i N / FIG. 307. Echinochalina (Protophlitaspongia) tuberosa sp.nov., digitate morph (specimen QMG300030). A, Auxiliary oxea (coring fibres and interstitial). B, Principal oxeas (echinating fibres). C, Ectosomal auxiliary subtylostyle. D, Oxhorn toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Digitate specimen. Echinochalina (Protophlitaspongia) laboutei Hooper Artemisina Vosmaer, 1885 & Lévi, 1993a: 1277-1280, figs 29-30 [New Artemisina Vosmaer, 1885a: 25; Ridley & Dendy, Caledonia]. MATERIAL. HOLOTYPE: QMG300685. SW. Pacific. 1887: 112; Topsent, 1894a: 12; Lundbeck, 1905: 110; Burton, 1930a: 501, 528-531; de Laubenfels, REVISION OF MICROCIONIDAE 559 FIG. 308. Echinochalina (Protophlitaspongia) tuberosa sp.nov., digitate morph (specimen QMG300030). A, Choanosomal skeleton. B, Fibre characteristics. C, Point of principal oxea. D, Point of auxiliary oxea. E, Ends of auxiliary style. F-G, Oxhorn toxas. 1936a: 117; Lévi, 1960a: 61, 83; Ristau, 1978: 585; TYPE SPECIES. Artemisina suberitoides Vosmaer, Van Soest, 1984b: 122, 130. 18852: 25 (by monotypy) (junior synonym of Suberites Artenisina; Burton, 1934b: 54 [lapsus]. arciger Schmidt, 1870: 47 (Burton, 1930a: 528)). Qasimella Thomas, 1974: 311. 560 DEFINITION. Without choanosomal fibres or indefinite fibres, whereas skeletal architecture vaguely ascending longitudinal tracts of spicules bound by abundant collagen, cored by smooth choanosomal principal subtylostyles in a more- or-less confused halichondroid reticulation of vaguely multispicular ascending and scattered transverse megascleres; echinating megascleres absent; subectosomal peripheral skeleton more radially arranged; ectosome membraneous, skin- like, with smooth styles of a single size category protruding through surface, forming paratangen- tial or erect, discrete spicule bundles; microscleres palmate or arcuate isochelae and toxas with smooth or spined points. Artemisina apollinis (Ridley & Dendy, 1886) (Figs 309-310) Amphilectus apollinis Ridley & Dendy, 1886: 350; Ridley & Dendy,1887:125, pl.19, fig.10, pl.25.fig.2. Artemisina apollinis; Landbeck, 1905: 114-116, pl.13, fig.4; Kirkpatrick, 1908b: 34, pl.20, fig.4; Hentschel, 1914: 70; Topsent, 1917: 62, pl.4, fig.6, pl.6, fig.7; Burton, 1929a: 431; Hentschel, 1929: 876, 939; Burton, 19302: 529; Burton, 1932a: 323; Koltun, 1964a: 73; Desqueyroux, 1975: 66, pl.4, figs 42-46; Koltun, 1976: 188; Hooper & Wieden- mayer, 1994: 256. Artemisina dianae Topsent, 1907:70-72; Topsent, 1908: 22, pl.3, fig.4, pl.5, fig.1; Vosmaer, 1935a: 630. MATERIAL. HOLOTYPE: BMNH1887.5. 2.140: Royal Sound, Kerguelen, 40-120m depth, HOLOTYPE of A. dianea: MNHN DT1666. HABITAT DISTRIBUTION. Rock reefs and soft sub- strata; 18-380m depth; widespread boreal and an- tiboreal: Antarctica - MacRobertson Land, Australian Antarctic Territory (Koltun, 1976), Graham Land, Wil- liam II Coast, Victoria Land (Desqueyroux-Faundez, 1975) (Fig. 309F), Gauss Station, Winter Quarters, South Georgia (Hentschel, 1914; Kirkpatrick, 1908b), Kerguelen Is (Ridley & Dendy, 1886, 1887; Koltun, 1976). Circum-polar (Koltun, 1964a). Arctic - East Greenland (Lundbeck, 1905), South Shetland Is (Top- sent, 1907, 1908), Iceland, North Sea (Hentschel, 1929). DESCRIPTION. Shape. Flattened, oval, cushion-shaped, up to 50mm long, 38mm wide, 19mm thick. Colour. Grey-brown preserved. Oscules. Not seen. Texture and surface characteristics. Soft, com- pressible, friable; surface uneven, folded, shaggy, conulose, cavernous, with skin-like ectosomal membrane stretched across adjacent conules. MEMOIRS OF THE QUEENSLAND MUSEUM Ectosome and subectosome. Variably developed spicule brushes forming more-or-less continuous palisade on surface, sometimes also tangential or paratangential; plumose brushes of subectosomal auxiliary spicules with few choanosomal prin- cipal spicules protruding through surface from underlying skeleton. Choanosome. Skeletal architecture irregularly reticulate or renieroid reticulate in places with rectangular and triangular meshes, up to 280j.m diameter, made of uni- or paucispicular tracts of choanosomal principal styles; skeletal tracts not definitely differentiated into primary or secon- dary elements although ascending tracts contain 2-4 spicules abreast and connecting, usually transverse tracts contain 1-2 spicules per row; fibres absent and spicules bound by light col- lagen; mesohyl matrix heavy, with numerous microscleres dispersed. Megascleres. Choanosomal principal styles long, slender, rounded smooth bases, occasionally slightly subtylote, with fusiform points, straight or slightly curved towards basal end. Length 352- (407.3)-458j.m, width 11-(14.0)-17 um. Subectosomal auxiliary styles long, slender, straight, with slightly subtylote, lightly microspined bases, occasionally smooth, taper- ing to fusiform points. Length 209-(244.6)- 293m, width 3-(4.4)-7jum. Microscleres. Palmate isochelae small, well silicified, unmodified. Length 8-(12.3)-161.m. Toxas wing-shaped, variable in length and thickness, with sharply angular, deep central cur- vature, long straight arms on larger spicules, or slightly recurved arms on smaller spicules; largest with microspined points. Length 24- (123.2)-264jum, width 0.8-(2.3)-5j.m. REMARKS. Ridley & Dendy (1887) stated that this species had a halichondroid reticulate skeleton whereas the type shows the skeleton to be much more regular, renieroid-reticulate in places. Synonymy of A. dianae and A. apollinis, proposed by Koltun (1964a, 1976) is doubtful, as is the assumption that the species is bipolar (i.e., possible two or more sibling species). Their respective type specimens are similar, with only slight differences in spicule sizes, spicule geometries and skeletal structures, and conse- quently Koltun's proposed synonymy is main- tained here. REVISION OF MICROCIONIDAE — — Le = X NY ns [7 1 v Ti `, E id BR ^e 2 at 8 i FIG. 309. Arremisina apollinis (Ridley & Dendy) (syntype BMNH1887.5.2.140). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Palmate isochelae. D, Wing-shaped toxas. E, Section through peripheral skeleton. F, Australian distribution. Artemisina jovis Dendy, 1924 (Fig. 311) Artemisina jovis Dendy, 1924a: 343, pl.12, fig.6; Bur- ton, 19302: 530; Koltun, 1964a: 74-75; Bergquist & Fromont, 1988: 119-120, pl.56, figs b-e; Dawson, 1993: 36; Hooper & Wiedenmayer, 1994: 257. MATERIAL. HOLOTYPE: BMNH1923.10. 1.112: 5km east of North Cape, New Zealand, 138m depth, PARATYPES: BMNH1923. 10.1.315, 316. HABITAT DISTRIBUTION. Hard benthos; 45-138m depth; Antarctica - Wilkes Land, AAT (Koltun, 1964a) (Fig. 31 1E); also North Cape and Three Kings Is, New Zealand (Dendy, 1924a; Bergquist & Fromont, 1988). DESCRIPTION. Shape. Flabellate, stalked, bifurcate digitate margins, 120-300mm high, 80- 110mm wide, 8mm thick lamellae. Colour. Greyish in ethanol. Oscules. Oscules up to 6mm diameter, with slightly raised margins, clustered on osculiferous face of sponge. 562 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 310. Artemisina apollinis (Ridley & Dendy) (syntype BMNH1887.5.2.140). A, Skeletal structure. B, Palmate isochelae. C, Wing-shaped toxas. Texture and surface characteristics. Firm, com- pressible; inhalant surface smooth, slightly un- dulating, with clearly visible porous ectosomal membrane stretched across pitted subectosomal region; exhalant surface osculiferous. Ectosome and subectosome. Erect plumose brushes of subectosomal auxiliary styles produc- ing continuous palisade at surface but markedly cavernous in subectosomal region. Choanosome. Nearly regularly renieroid reticu- late skeletal architecture composed of uni-, pauci- and multispicular tracts of choanosomal principal styles producing triangular meshes up to 3501m diameter, enclosed within moderately light col- lagen; fibres absent; spicule tracts relatively homogeneous, undifferentiated into primary or secondary elements; mesohyl matrix heavy, dark- ly pigmented, containing abundant micro- scleres. Megascleres. Choanosomal principal styles large, robust, straight or slightly curved near basal end, fusiform pointed, with smooth rounded or very slightly subtylote bases. Length 361- (390.3)-448jum, width 12-(19.2)-264.m. REVISION OF MICROCIONIDAE E E D N 563 FIG. 311. Artemisina jovis Dendy (holotype BMNH1923.10.1.112). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Accolada - wing-shaped toxas. D, Palmate isochelae. E, Australian distribution. F, Section through peripheral skeleton. Subectosomal auxiliary styles slender, short, fusiform pointed, straight or slightly curved near basal end, with smooth round bases. Length 234- (290.6)-375,1.m, width 3-(7.1)- 1Oj.m. Microscleres. Palmate isochelae unmodified, abundant, well silicified. Length 10-(12.5)- 16m. Toxas intermediate accolada-wing shaped, long, slender, with slightly rounded central] cur- vature, slightly reflexed arms, microspined points. Length 98-(155.2)-215y.m, width 1.5- (2.4)-4.m. REMARKS. The type material revealed some discrepancies from published descriptions of both Dendy (1924a) and Bergquist & Fromont (1988), indicating variability between specimens. In particular skeletal meshes, described as being differentiated into ascending and transverse com- ponents, are virtually homogeneous in the holotype, and skeletal structure described by Bergquist & Fromont (1988) as plumose is regularly renieroid reticulate with no plumose component in the holotype. Spicule sizes also differ marginally from those described by Dendy (1924a). Artemisina jovis is very similar to A. apollinis in its skeletal structure, differing only substantially in growth form, toxa size and to a certain degree toxa geometry, and size and spina- tion of other megascleres. Artemisina plumosa Hentschel, 1914 (Fig. 312) Artemisina plumesa Hentschel, 1914: 70-72, pl.4, fig.5, pl.6, fig.1 [et var. lipochela]; Burton, 1929a: 431 [Antarctica]; Burton, 1930a: 530; Koltun, 1964: 73, pl.11, figs 1-8; Koltun, 1976; 190; Hooper & Wiedenmayer, 1994: 257. Artemisina strongyla Hemschel, 1914: 72-73. pL6. fig.2. MATERIAL. HOLOTYPE: HM (fragments BMNH- 1933.7.20.3, ZMB4762): Off Wilhelm Il Coast, An- tarcica. HABITAT DISTRIBUTION. Soft substrata; 125- 2269m depth; Antarctica - Enderby Land, Sabrina Coast, Princess. Elizabeth Land, Wilhelm 1l. Coast, Australian Antarctic Territory (Koltun, 1976) (Fig. 312F); also Falkland Is, Adelie Coast (Koltun. 1964a)_ Gauss Station (Hentschel, 1914; Burton, 1929a). DESCRIPTION. Shape. Small, erect, bulbous, club-shaped, 28mm high, 25mm diameter. Colour. Grey-brown in ethanol. Oscules, Not observed. Texture and surface characteristics. Firm, harsh texture; irregulary reticulate conulose. Ectosome and subectosome. Plumose bundles of choanasomal principal styles protruding, form- ing nearl continuous palisade on surface; subec- tosomal auxiliary styles tangential or paratangentia] to surface, sparse, dispersed be- tween protruding bundles of choanosomal styles, Chounosome. Exlusively plumose near peripheral skeleton, slightly more reticulate, less organised in deeper parts of choanosome, with multispicular ascending tracts of choanosomal styles, without spongin fibres, most obvious near peripheral region of skeleton; length and width of spicule tracts reduced al core with diminished MEMOIRS OF THE QUEENSLAND MUSEUM skeletal organisation whereby only pauci- or smal] multispicular tracts form skeletal reticula- lion at core, with square or triangular meshes, up to Wum diameter; mesohy} matrix heavy, light- ly pigmented, smooth, containing very abundant microscleres throughout. Megascleres. Choanosomal principal styles long, robust, fusiform pointed, smooth, with rounded bases and typically slightly curved near basal end. Length 305-(378.2)-472pm, width 11-(13.7)- Bum. Subectosomal auxiliary styles or tornostyles shon; thin, straight, with slightly subtylote bases, smooth or microspined bases, rounded, hastate or mucronate points often with small spines, nr sometimes with single small terminal spine, or guasi-diactinal asymmetrical ends. Length 181- (196.7)-268j.m. width 3-(6.1)-Bjum. Microscleres. Palmate 1sochelae very abundant, moderately large, well silicified, occasionally contort. Length 8-(13.2)-165.m. Toxas intermediate accolada-wing shaped. varying from minute to very large, typically with large angular central curvature, slightly reflexed arms, microspined points. Length 36-(140.1)- 296m, width 0.8-(2.5)-5j.m. REMARKS. Arremisina plumosa differs from other species in having quasi-diactinal auxiliary meeascleres, including some with basal spins- tion. Its toxas are also much larger and mare prominently spined than the other 3 antarctic species described here, and skeletal architecture is heavily reticulate at core and exclusively plumose near periphery. Hentschel (1914) and Koltun (19644) recorded u second category of large choanosomal style, with similar geometry to principal spicules but nearly twice their length (800-2160x21-44yum), but these were not seen in the fragment of the holotype examined. Artemisina tubulosa Koltun, 1964 (Figs 313-314) Artemisina tabulosa Koltun, 19642: 74, text-fig. l8; Hooper & Wiedenmayer, 1994: 257. MATERIAL. HOLOTYPE: BMNH1963.7. 29.52. Mawson Coast, Australian Antarctic Territory, 100m depth HABITAT DISTRIBUTION, Hard benthos; [00m depth; Australian Antarctic Territory (Fig. 313), DESCRIPTION. Shape. Tubulo-digitate, hollow, cylindrical, 100mm long. 30mm diameter. REVISION OF MICROCIONIDAE 100 um 25um gJ € 565 FIG, 312. Artemisina plumosa Hentschel (fragment of holotype BMNH1933.7.20.3). A, Choanosomal principal styles. B, Subectosomal auxiliary styles/ tornotes. C, Accolada - wing-shaped toxas. D, Palmate isochela. E, Section through peripheral skeleton. F, Australian distribution. Colour. Beige-brown in ethanol, Oscules. Not seen. Texture and surface characteristics. Firm, harsh; surface microconulose, goose-flesh, pitted. Ectosome and subectosome. Hispid, with plumose bundles of choanosomal principal styles protruding through surface in mulüspicular bundles; subectosomal auxiliary styles tangential or paratangential, scattered over surface, also with detritus embedded in surface. Choanosome. Reticulate skeletal architecture, with large multispicular tracts of choanosomal principal spicules ascending to surface, and ir- regular uni-, paucispicular or smaller, transverse multispicular tracts interconnecting main tracts, together producing a nearly myxillid-like trian- gular reticulation, with meshes up to 800um diameter; spongin fibres absent, with spicule tracts bonded together by collagen; mesohyl matrix heavy but only lightly pigmented, with numerous microscleres scattered between fibres. Megascleres. Choanosomal principal styles thick, fusiform, rounded bases, smooth, slightly curved near basal end. Length 705-(778.8)- 874jum, width 18-(30.2)-391.m. Subectosomal auxiliary styles short, thin, mostly straight, with rounded or very slightly subtylote or occasionally polytylote bases, typi- 566 cally microspined, and with blunt hastate microspined points, Length 384-(456.3)494ym, width 6-(8.4)-11j.m. Microscleres. Palmate isochelae abundant, moderately large, well silicified, unmodified. Length 12-(15.8)-19j.m. Toxas intermediate accolada-wing shaped, lang, thick, angular central curvature, spined points. Length 602-(674.2)-824j.m, width 4- (6.4)-9.m. REMARKS. Artemisina tubulosa is similar to A. plumasa in skeletal architecture and spicule geometry, and the two are probably at least sib- ling species, Whereas A. tubulosa has microspined base on some of the auxiliary spicules it lacks diactinal modifications to those spicules as found in A. plumosa. Similarly, skele- tal structure is also prominently reticulate, with large spicule tracts clearly differentiated into primary and secondary elements, whereas those of A. plumosa are exclusively plumose in the peripheral skeleton. OTHER SPECIES OF ARTEMISINA Artemisina archegona Ristau, 1978 Artemisina archegana Ristau, 1978b: 585-587. text- figs 2F, 3P, 6C-D [Carmel, California]. MATERIAL. HOLOTYPE: USNM24528. Province: NE. Pacific. Artemisina arciger (Schmidt, 1870) Suberites arciger Schmidt, 1870: 47, pl.5, fig.6. Artemisina suberitoides Vosmaer, 1885a; 25-26, pl. 1, fig.16, pl.4, figs 11-14, pl.5, figs 51-55 [Arctic]; Ridley & Dendy, 1887: 112 [off Nova Scotia]; Fris- tedt, 1887: 430, pl.24, figs 15-17: Whileaves, 1901; 17; Lundbeck, 1905: 113. Artemisina arcigera; Lundbeck, 1905: 110-114, pl.1, figs 9-11, pl.13, fig.3; Arndt, 1913: 115; Topsent, 1913b: 47, phl, fig.6: Rezvoi, 1925: 197, [var. spiceps; Arche]; Burton, 1930a: 528-529 |Lype- species]: Hentschel, 1929; 876, 938 [Arctic]. Artemisina arciger; Burton, 1959h: 42 [Iceland]; Van Soest & Stone, 1986: 47 [note]. MATERIAL. HOLOTYPE; LMJG, lragment BMNH1870.5.3,90; holotype of suberitoides: ZMA PORAA3, fragments BMNHISO1.1.1604, 1351. Province; NW, Atlan- lic, NE, Atlantic, Arctic. Artemisina elegantula Dendy, 1924 Artemisina eleganiula Dendy, 19242: 344 |N.of New Zealand]; Burton, 1930a; 531; Bergquist & Fromont, 1988: 120; Dawson, 1993; 36 [index]; Lévi, 1994: 36, fig.206, pL7. fig.1 [S. of New Caledonia]. MATERIAL. HOLOTYPE: BMNH1923,10. 1,113. New Zealand, New Caledonia. MEMOIRS OF THE QUEENSLAND MUSEUM Artemisina erecta Topsent, 1904 Artemisina erecta Topsent, 1904u; 214-215, pl.5, fig.18, pl, 15, fig.10 [Azores]; Burton, 1930a: 530, MATERIAL. HOLOTYPE: MOM. NE. Atlantic. Artemisina hispanica Ferrer-Hernnádez, 1918 Artemisina hispanica Ferrer-Hemández, 1918: 536- 538. fig.3, text-fig.5 [Santander]; Lévi, 19603; 83 |Santander, Mediterranean]; Burton, 19302: 530. MATERIAL. HOLOTYPE: Madrid. Medilerranean. Artemisina indica (Thomas, 1974) Qusimella indica Thomas, 1974; 3] 1-312, text-fig.1 [Gulf of Manaar]. MATERIAL. HOLOTYPE: CMFRITS4/1, paratype CMFRIT84/2. N. central Indian Ocean. Artemisina melana Van Soest, 1984 Artemisinu melana Van Soest, 1984b; 122-124, pls, figs 7-8, text-fig.49 [Curacao]; Zea, 1987: 173, text- fig.61, pl. 1H, figs 3-4 [Columbian Caribbean). MATERIAL. HOLOTYPE; ZMAPOR4BS1, paratype ZMAPOR4882, 4883, Caribbean: Artemisina stipitata Koltun, 1958 Artemisina stipitata Koltun, 1958: 52-53, text-lig. 10 [Kurile Strait]. MATERIAL. HOLOTYPE: BMNH1963.7. 29.39. NW. Pacific. Artemisina transiens Topsent, 1890 Artemisina transiens Topsent, 1890b: 66 [namen nudum]; Topsent, 1892a: 94, pl.2, fig.5, pt.4, fip. 8. pl.9, fig.18 [Atlantic coast of France]; Ferrer» Hernández, 1914: 3; Burton, 19302: 529-530; Lévi, 19602: 83-84 [Asturies, Mediterranean |. MATERIAL. HOLOTYPE: MOM. NE, Atlantic, Mediter ranean. Artemisina vulcani Lévi, 1963 Artemisina vulcani Lévi, 1963: 64-65, pl.9F, text- lig.74 [South Africa]. MATERIAL. HOLOTYPE: MNHNDCLG622. South Africa. TRANSFERS List of other species described in Artemisina but now transferred to another genus, Paresperia intermedia Burton, 1930a; 501 [Norway |, Artemisina intermedia, Van Soest, 1984b: 130 [pos- sible generic synonymy]. Esperiopsis intermedia; Van Soest & Stone, 1986; 45 MATERIAL. HOLOTYPE: BMNHi1910.]. i.912. Nate: referred to Desmacididae, Experiopsis. Amphilectus pilosus Ridley & Dendy, 1886: 350; Ridley & Dendy, 1887: 126-127, pl. 19, fig, 5, pl.25, fig.3 [Kerguelen Is; Marion 1.]. Lissodendoryx pilosus; Topsent, 1901: 20. Megaciella pilosa; Hallmann, 1920: 772; Van Soest, 1984b: 128 [possibly Myxillidae]. REVISION OF MICROCIONIDAE 567 jf ZZ, A A 100 um 25um FIG. 313. Artemisina tubulosa Koltun (holotype BMNH1963.7.27.52). A, Choanosomal principal styles. B, Subectosomal auxiliary styles. C, Accolada - wing-shaped toxas. D, Palmate isochela. E, Section through peripheral skeleton. F, Australian distribution. Artemisina pilosus; Ristau, 1978: 585-586 [note on Artemisina tylota Boury-Esnault, 1973: 286-287, text-fig.48 affinities]. [Brazilian Basin]. MATERIAL. HOLOTYPE: BMNH1887.52. 124, Note: Cormulum tylota; Van Soest, 1984b: 124. referred to lophonidae, Megaciella. 568 MEMOIRS OF THE QUEENSLAND MUSEUM MATERIAL. HOLOTYPE: MNHNDNBE?943, Note: referred to lophonidae, Cornulum. Incertae sedis Pandaros Duchassaing & Michelotti, 1864 Pandaros Duchassaing & Michelotti, 1864: 88; Schmidt, 1870: 59; de Laubenfels, 1936a: 123; Wieden- EE mayer, 1977: 143; Van Soest, f 1984b: 127. TYPE SPECIES. Pandaros acan- thifolium Duchassaing & Michelotti, 1864; 90 (by subsequent designation ofde Laubenfels, 1936a; 123). DEFINITION. Well-developed choanosomal skeletal reticulation of spongin fibres woven into flat- tened anastomosing branches, or forming continuous sheets without | any regular architecture; fibres Ky cored by auxiliary styles, subtylos- MPs. tyles to tylostyles, predominantly [| smooth, distributed within fibres in criss-cross fashion; fibres § echinated by rare acanthostyles, rarely smooth styles; few anisotor- notes, verging on oxeas, scattered throughout mesohyl; sparsely dis- persed tangential subectosomal auxiliary spicules dispersed throughout peripheral skeleton; microscleres absent. REMARKS. No Australian species. SPECIES OF PANDAROS Pandaros acanthifolium Duchas- saing & Michelotti, 1864 Pandaros acanthifolium Duchassaing & Michelotti, 1864: 90, pl.20, fig.2 [St.Thomas, Virgin Is]; de Lauben- fels, 1936a; 123 [Florida]; Hechtel, 1965: 45; Wicdenmayer, 1977: 144, pl.30, figs 4-5, text-fig.148 |Bimini, Bermudas]; Schmitz et al, 1981: 853-865 [biochem.]; Van Soest, 1984b: 127-128, text-fig.51 [Curacao]; Pulitzer-Finali, 1986: 149, fig.68 [West Indics]; Zea, m 7 1987: 175, pl, fig.4, text-fig.62 FIG. 314. Artemisina tubulosa Koltun (holotype BMNH1963.7.27.52). {Columbian Caribbean]. A, Skeletal structure. B, Palmate isochelae and spined toxas. REVISION OF MICROCIONIDAE Pandaros arbusculum, in part, Duchassaing & Michelolti, 1864: 88, pl.18, fig.6 [St. Thomas, Vir- gin Is]. Thalyseurypon conulosa Hechtel, 1965: 44, pl.6, fig. l. text-fig.9 [Port Royal; Jamaica]; Alcolado, 1976; 5 [Cuba]. MATERIAL. LECTOTYPE: TMPORS7 (fragments USNM31020, MNHNDNBE!309, BMNHI1928.11.12.16). paralectorvpe: IZUGCE38.762 (fragment BMNH1928.11. 12.15); holotype of T. conulosa: YPM5042. Caribbean. Pandaros kasumiensis (Tanita, 1965), uncertain placement Raspailia kasumiensis Tanita, 1965: 67, text-figs 1-2 [Japan]; Hoshino, 1970: 23, text-figs 3(2), Udh; Hoshino, 1971: 24. Thalyseurypon kayumtensis; Hoshino, 1981: 159-161. text-fig.70 [Japan]. 7 Pandaros kasumiensis, Hooper, I988b; 353, 766. MATERIAL. HOLOTYPE: MMBSSIS 452. NW. Pacific. TRANSFERS List of other species described in Pandaros but now transferred to another genus. Pandaros angulosa Duchassaing & Michelotti, 1364: $9, pl.19, fig.4 [St. Thomas, Virgin Is]. Mycale (Aegogropila) angulosa; Van Soest, 1984b. 16-19, pl.], figs 5-6, texi-fig.3 [plus synonymy]; Pulitzer-Finali, 1986: 130-133, figs 52-55. Not Mycale angulosa. de Laubenfels, 19363; 116. Zygomycale parishii, in part; de Laubenfels, 1956; 3; Hechtel, 1965: 48, pl.5, fig.3; Alcolado, 1980: 4. MATERIAL. LECTOTYPE: TMPORS4 (fragments USNM31021, MNHNDNBEI310, BMNH1928.1 1.12.20). referred to Mycalidae, Mycale, Pandaros arbusculum Duchassaing & Michelotti, IRGA: 88, pl.18, fig.6 (St. Thomas, Virgin ls]. Pulocaulis sp. aff. spiculifer (Lamarck, 1814); Wiedenmayer, 1977: 153,251. Piilocaulis arbusculum; Van Soest et al., 1983; 204. MATERIAL. LECTOTYPE: ZMAPOR 1728, paralectotype ZMAPORI729. referred to Axinellidae, Prilocaulis, Pandaros lugubris Duchassaing & Micheloni, 1864: 89 (St. Thomas, Virgin Is]. Echinodictyum lugubre; Wiedenmayer, 1977: 255; Pulitzer-Finali, 1986: 106-107. MATERIAL. LECTOTYPE: TMPORSS (fragments USNM31023, MNHNDNBE1312, BMNHI928.1 1.12.21). referred to Raspailiidae. Pandaros pennata Duchassaing & Michelotti, 1864: RB. pl.20, fig.3 [St, Thomas, Virgin Is]. Echinodictyum pennatum, Wiedenmayer, 1977: 251, 255. MATERIAL. LECTOTYPE; TMPORSS (fragments USNM31024, MNHNDNBEI313, BMNHI928.11,12.22), paraleciotype: ZMA PORI731. teferred to Raspailiidae, Echinodictyum. 569 Pandaros walpersii Duchassaing & Michelotti, 1864; 90, pl.20, fig.4, text-fig-2D [St Thomas, Virgin ls), Piilocaulis walpersi; Wiedenmayer, 1977; 255. Spongia spiculifera Lamarck, 1814: 449. Piilocaulis spiculifera; Pulitzer-Finali, 1986: 103-104 [ West Indies]. MATERIAL. LECTOTYPE: TMPORS6 (fragments USNM31025, MNHNDNBE!314, BMNHI1928.1 1.12,17). paralectotype: IZUGCE 38.763. referred 10 Axinellidae, Prilocaulis. DISCUSSION Classification. Microcionidae contains 7 genera, 12 subgenera and 459 valid species of which 148 are recorded in the Australian fauna. The present work describes 3] new species from Australia, and 3 from other localities for misidentified taxa, 18 new names for preeccupied taxa, and 162 new taxonomic combinations. Many species are also new records for Australiaand many are illustrated for the first time. Nevertheless, there are likely many more species still undescribed in this region considering that the encrusting fauna has not yet been comprehensively sarnpled. This revised classification is hased on a me- evaluation of all characters, including an analysis of their consistency within populations of severul species, and places primary importance on skele- tal structure and localisation of spicule types to particular regions in the skeleton. This contrasts with previous schemes which gave priority to characters such as patterns of skeletal architec- iure (e,g., Lévi, 19603), megasclere form and distribution within the skeleton (Hallmann, 1920), ectosomal structure and composition (Van Soest, 1984a), microsclere geometry and diver- sity (c.g., de Laubenfels, [936a). or combining all these characters into a a consensus matrix with u consequent proliferation of genera (e.g. Bergquist & Fromont, 1988). Re-examination of the type species of 73 nominal genera included in Microcionidae at onc time or another found that many had been mis- described, or important characters overlooked, with the consequence that previous interpreja- tions of genera are mostly incorrect. Furthermore, many of these nominal genera are not valid in à phylogenetic sense given that their supposed apomorphic characters are unstable (inconsis- tent) within otherwise closely related species. These earlier classifications had extraordinary high levels of homoplasy (Hooper, 19902], partly a consequence of relying on single morphologicul characters (which when used in isolation cur across a classification based on other single char- 570 acters). | use combinations of characters [o develop a phylogeny. Atthe species level several characters are prone to ecophenotypic variabilitv (notably growth form, effect of water depth on colour. loss of microscleres, reduced silification and variability in spicule dimensions across broad geographical ranges). whereas most other skeletal characters are more consistent within populations of single species. It is proposed that small cryptic differen- ces in skeletal characters are indicative of diver- gent, sibling species relationships, with some empirical support from biochemical and genetic data (Hooper et al., 1991). Phylogenetic analysis. Hypothesised phylu- genetic relationships based on coded, multistate (Table 47) data matrices were generated using. PAUP 3.0 (Swofford, 1991), and CONTREE was used to obtain consensus information from these analyses as outlined by Hooper & Lévi (1994). Determinations of plesiomeorphies were largely made through oulgroup comparision (Raspailiidac), bul these interpretations are com- plicated by the prevalance of analogous charac- ters throughout Porifera (or misinterpretation of homology by authors). Consequently some infor- mation from the fossil record is considered below to assist in determining character polarity. (1) Skeletal structure: Ordovician Saccospon- gta had a clathrous, fibrous reticulate skeleton, cored by axially compressed and extra-axially plumose spicule tracts, with coring spicules also protruding through fibre endings in plumose bundles (Finks, 1967). A similar structure in some living poecilosclerids, such as Esperiopsis (Hooper & Lévi, 1989), which also have a "hthistid! (desma-bearing) basal skeleton — a condition relatively widespread during the Or- dovician (Finks, 1970) — imply that the axial compressed condition may be plesiomorphic for living Clathria (Axociella) and many Raspailia ( Raspailiidae). The use of Raspailiidae as an outgroup of Microcionidae is based on their proposed sisier- group relationship (Hooper. 1991), shanng a similar range of skeletal structures (including hymedesmoid, microcionid, reniernid, reticulate and compressed architectural types), echinating acanthostyles and ectosomal specialisation. These structural similarities are considered to have a phylogenetic basis (with some empirical support from biochemical and genetic data; Hooper et aL, 1992) rather than merely repre- senting convergence of a few characters. Raspailiidae was previously included in order MEMOIRS OF THE QUEBNSLAND MUSEUM Axinellida based solely on possession of axial skeletal compression in some species (Bergquist, 1978), but this character is of dubious value when used alone occurring in many other poecilo- sclerids. (2) Ectosomal skeleton: The 'sclerosponge* Calcifibrospongia has a tangential (isodictyal) ectosomal skeleton of strongyles lying on or just below the exopinacoderm (Hartman, 1979); Vacelet (1985) and Van Soest (1984a) suggested that this feature links it with the Haplosclerida, particularly the Haliclonidae, This would infer that a tangential ectosomal skeleton is plesinmor- phic for the haplosclerids, However, similar fea- lures in some Microcionidae represent parallel development (c.g. quasi-diactinal ectosomal spicules in some Holopsamnma, quasi-diactinal structural spicules in Echinochalina). The tangen- lial layer of amphitylotes in the Myxillidae is unique to the group (e.g., Lissodendoryx), upon which Van Soest (1984h) based his classification. but the myxillid (tangential amphitylote) ec- tosomal skeleton in Acarmüus is enigmatic since its microscleres are typical of Microcionidae, Thus Hooper (1988, 19903) considered that microscleres were more easily modified, ac- quired independently or lost in the Poecilo- sclenda than was the ectosomal skeleton, which in this case defined the Myxillidae. Hajdu et al. (1994) though! that mieroselere geometry was far more conservative than that of megascleres (which might be more susceptible to phenotypic influences). Less obvious is the polarity nf the sequence Clathria-Dendrocia-Thalysias, vary- ing from a virtually aspicular ectosome to à dense, continuous palisade of erect megascleres. ] suggest that the latter condition is more plesiomorphic based on the ‘raspailiid ectosomal condition’ (i.e, smaller auxiliary spicules sur- rounding bases of protruding larger subectosomal spicules) being common to the outgroup Raspailiidae, and diminishment of ectosomal structure or loss of spicules representing a secon- dary derivation. (3) Spicule axes: It could be argued that oxcote (diactinal) megascleres represent the ancestral condition in this family because they appeared in the Cambrian (Rigby, 1986), whereas styloid (monactinal) spicules are not known earlier than the Ordovician. However, in Microcionidae oxeores (in E. (Protwophlitaspongia) for example) represent recent derivations from styloid forms (much like the derived tetractinal styloid modifications in Acarntes (Myxillidae); Van Soest et al., 1991). REVISION OF MICROCIONIDAE 571 TABLE 46. Comparison between species of Echinochalina (Protophlitaspongia) in major morphological characters.Measurements in jum. | HARACTER| E. bispiculata | | E. oxeata (Dendy) (Burton) irregularly reticulate, detritus cavernous in axis, more compact in periphery Colla m in meso| elongate or oval (180- 450) irregularly reticulate, distinct primary and d cavernous, oval (145- 550) heavily condensed axis, regularly reticulate extra-axis elongate (50- 250) E. bargibanti Hooper & Lévi regularly reticulate throughout, cavernous meshes E. tuberosa Sp.nov. E. favulosa Sp.nov. egularly plumo- reticulate, diverging near periphery, meshes even throughout irregularly reticulate, wide meshes, much detritus square or round (70- 150) elongate E. isaaci sp.nov. regularly reticulate, wide even meshes diverging near periphery elongate (370-650) E. collata sp.nov irregularly reticulate, fibres 2% cored oval or elongate (120-440) 20-50 22-55 15-90 Spicules in eletal tracts Spicule well silicified | well silicified silification Corin poorly silicified pauci- or multispicular | paucispicular multispicular uni- or aspicular uni- or aspicular px av a massive- lobate, tubular digits on surface branching, cylindrical, digitate cylindrical digitate, arborescent (4) Spicule ornamentation: Hypercalcified sponges (’sclerosponges’) were major con- tributors to reef building during the Palaeozoic and Mesozoic (Hartman & Goreau, 1975; Hartman, 1979), with extant species (in Merliida and Ceratoporellida) having entirely acanthose choanosomal megascleres (verticillate acanthos- tyles). Vacelet (1985) suggested that sclerospon- ges were polyphyletic having affinities with many demosponge groups, and if living species represent relict ancestral groups of Demospon- giae, then there is argument to suggest that acan- those megascleres are ancestral. Conversely, smooth monaxons were predominant in Palaeozoic and Mesozoic sponges such as Sac- cospongia (Rigby, 1986), in which case the smooth condition may be more primitive. This is a subject of continual speculation, both hypotheses equally supported by tenable argu- uni- or paucispicular unispicular clumped lobate- digitate poorly silicified well silicified | well silicified | well silicified | well silicified pauci- or rarely multispicular pauci- or EM multispicular multispicular multispicular uni- or paucispicular uni- or paucispicular uni- or aspicular unispicular or aspicular erect, cylindrical digit,honey- combed digitate, lobate- digitate, or encrusting thickly encrusting, ments (e.g., Van Soest, 1984a, 1987). Within Microcionidae it is considered that acanthose spicules are plesiomorphic, shared by most species, whereas smooth spicules represent a derived secondary loss of spination. This inter- pretation is consistent with similar determination for Raspailiidae (Hooper, 1991). (5) Echinating spicules: Echinating spicules are known for 7 families of Poecilosclerida and may represent a synapomorphy for the suborders Microcionina and Myxillina (Hajdu et al., 1994) (i.e. absent from the base group Mycalina). Not all genera within these families have them and their absence is interpreted as a (derived) secon- dary loss (with the corollary that echinating spicules in Microcionidae is retention of an an- cestral character). Similarly, smooth echinating or pseudo-echinating spicules, such as in Echinoclathria and Echinochalina, have probab- MEMOIRS OF THE QUEENSLAND MUSEUM 2/1— Artemisina LÀ Siri om $/2— Antho (A ) (9128/3) B Lei SÀ 2/332 5/4j9rT ee — — Antho (P.) 8/49/U- Antho (I.) 9/3/8/55/3) — Echinoclathria (9/32/1173 Holopsamma 19/4—- Echinochalina (E.) | 13/94 urs" Echinochalina (P ) Gap M rm41555— Pandaros JAM 5/4 Clathria (D.) 9 — Clathria (W.) " Clathria (C.) 2/3——— Clathria (T ) —6/5— Clathria (I.) a26 ini Clathria (M.) — 7a Clathria (A.) 11/Mo/+——. Raspailia FIG. 315, Hypothesised relationships between genera of Microcionidae based on computer-generated phylogenetic analysis using parsimony (PAUP 3.0; Swofford, 1991). Each number on the cladogram (x/y) corresponds to a character and character slate respectively (Table 47) and represents an evolutionary change from a plesiomorphic to an apomorphic state. The tree is derived from a consensus of 9 possible minimum length trees, based on an unweighted, unordered, multistate character set (consensus information = 0.609), with plesiomorphy determined by outgroup comparisons with the Raspailiidae, ly lost their spination and are a derived condition because in most species they are spined. (6) Chelae morphology: Hajdu et al. (1994a) subordinal classification of Poecilosclerida large- ly based on modifications to chelae (i.e., Microcionina with only palmate forms, Myxil- lina with modified bi- or tridentate modifications, Mycalina with sigmancistra derivatives), Pal- mate chelae were thought to be ancestral given their prevalence throughout Poecilosclerida and co-occurrence with other forms of chelae, whereas arcuate chelae were derived from ar- chetypal palmate morphology. Similarly anchorate chelae were thought to have arisen from arcuate modifications to the primitive pal- mate archaetypes. Lateral ridges on the shaft of anchorate chelae were interpreted by Hajdu et al. (1994a) as the beginning of two other (new) lateral alae (i.e. acquisition of new structures), which occasionally meet in the middle of the shaft to produce a continuous ridge along the length of the spicule. An alternative view is that these ridges may be the remnants of the point of attach- ment between the lateral alae and shaft (i.e., a reduction from the existing fused structure), If this latter interpretation is correct (and il is ener- gelically more probable than the acquisition of à new structure), then anchorate chelae may TABLE 47, List of characters and character states used to judge apomorphy in the construction of the cladogram of relationships between genera of Microcionidae based on outgroup comparisons with members of the family Raspailiidae. States marked with an asterisk indicate pleisiomorphic condition(s) and discussed in the Analysis of Morphometric Characters. Consistency Index (CI) is indicated for each character obtained from parsimony ananlysis (Swafford, 1991) REVISION OF MICROCIONIDAE 1, Growth form specialisation. (CI = 1.0) 7. encrusting or hulbous-encrusting growth forms. 2.* massive, branching, lobate, flabellate, yasifonm or tubular growth forms. 3. specialised honeycomb-reticulate growth forms composed of lightly anastomosing flattened fibre-branches (lacunae). 2. Ectosomal skeleton specialisation. (CI = 0.6) t. mem- braneous,skin-like exterior, with single category of (subec- losornal) auxiliary spicule extending from choanosomal skeleton protruding through surface forming discrete paratangential or erect brushes. 2. with single category of (subectosomal) auxiliary spicule on surface forming tangential, paratangential or plumose tracts. 3, with two categories of auxiliary spicules.smaller eclosomal spicules generally overlaying larger subectosomal spicules forming, discrete bundles or continuous palisade on surface, 4,* with apecialisedectosomal skeleton composed of smaller auxiliary spicules surrounding hases of protruding larger subectosomal spicules. 3. Subectosomal skeleton structure. (CI 0.5) 1. radial with perpendicular bundles or single spicules supporting ectosome. 2.* plumosc, plumoreticulate, or disorganised tracts of spicules in peripheral skeleton, 4. Differentiation within choanosomal skeleton. (C1 = (0.667). chosnosomal skeleton more-or-less undifferentiated, unstructured. 2. choanosomal skeleton well structured, hy- medesinoid to reticulate, but lacking any differentiated com- ponents. 3.* choanosomal skeleton well structured, predominantly reticulate, differentiated into two distinct components. 5. Fibre development and skeletal architecture. (CI = 0,714) J. choanosomal skeleton without spongin fibres (or indehinite fibres), with spicules forming more-or-less disorganised hali- chondroid, vaguely ascending, longindinal reticulate tracts. 2, choanosomul skeleton with poorly developed spongin fibres, primacy (basal or axial) renieroid component cored by acanthose principal spicules secondary plumose, subisodictyal or plumo- reticulate component cored by sipooth principal spicules. 3. choanosomal skeleton. with well developed spongin fibres, primary skeleton compressed, remeroid, cored smaller smooth principal styles, secondary skeleton vestigial or greatly reduced, consisting only of larger smooth principal styles embedded in peripheral] fibres forming sparse radial tracts. 4. ehoanosomal skeleton with well developed spongin fibres forming more-or- less evenly reticulate or plumo-reticulate meshes throughout in massive forms, or thick basal layer of spongin and hymedesmoid structure in encrusting forms. 5. choanosomal skeleton with well developed spongin fibres woven into Natlened reticulate branches or forming cuntinuous sheets. without iny regular architecture, cored by criss-cross of auxiliary styles, 6." chounosomal skeleton with well developed spongin fibres torm- ing compressed reticulate axis and well differentiated radial, plumose or plumo-reticulate extm-axis. 6. Compression of choanosomal skeleton, (CI = O08) +. choanosomal skeleton without any marked ixial compression or differentiated axial and exwa-axial regions. 2, choanosomal 573 skeleton withour marked axial compression but well difieren- tiated axial and extra-axial (radial, plumose of plumorehcu- lale) regions. 3.* chosnosomal skeleton with noticeably compressed axis and well differentiated axial and exira-axial (radial, plumose or plumoreticulate) regions. 4. choanosomal skeleton hymedesmoid or microcionid, with basal layer of spongin lying on substrate (with or without ascending, non-anustomosing fibre nodes), and bases of principal spicules standing perpendicular lo substrate, 5, choanosomal skeleton evenly renieroid or isodictyal reticulate throughout with well developed spongin fibres coned by smooth principal styles, 7. Derivation of spicules coring fibres, (CI = 0,75) J.* choanosomal fibres or skeletal tracts cored by one ec more category of principal spicules. 2. choanosomal fibres or skeletal tracts cored by auxiliary spicules but prwtially or wholly replaced by detritus, 3. choanosomal fibres or skeletal tracts cored by auxiliary spicules idenlica) 1o these iñ bo- losomal and subectosomal skeletons. 4. choanosomal fibres or skeletal tracts cored by auxiliary spicules different from those in peripheral skeleton, 8. Differentiation of primary and secondary skeletal coni- ponents (CI = 1.0) J.* primary and secondary skeletons undifferentiated, 2, primary skeleton renieroid cored by axial ly or basally compressed tracts of acanthostyles secondary skeleton cored by smooth principal styles in plumose, sub- isodictyal or plumoreticulaté tracts. 3. primary skeleton renieroid cored hy axially or basally compressed tracts of acanthostrongyles, secondary skeleton cored by smooth prin- cipal styles in. plumose, subisodicryal or plumoreticulate tracts. 4. primary skeleton axially compressed spongin fibers Cored hy renieroid tracts of sparsely spined principal styles intermingled with plumose or plumoreticulate tracts of smooth principal styles, overlaid by secondary extra-axia) plamose skeleton cored by larger smooth principal styles. 5. primary reiieroid reticolate skeleton cored by smooth principal Uyles wad echinaled by identical spicules, with secondary radial extra-axial skeleton om exterior edge of skeleton only cored by larger smooth principal styles, 9. Den vation of echinaung spicules: (CI 20.444) J, echia- img spicules absent. 2.* special category of (acantho)styles present eehinating fibres, differentiated from principal spicules. J. echinating smcules styles or acanthostyles undil- ferentiated trom principal spicules coring spongin fibres. 4. echinating spicules styles or acanthostyles representing prin» cipal spicules, but different from those coring fibres, 5. echinating spicules oxeas or anisoxeas representing principal spicules, buy different from those coring fibres, 10. Modifications to chelae microscleres. (CI = 0.5) I. chelae absent. 2.* isochelae palmate. J. 1sochelne arcuate- like 4. Isochelae aqchorate-like, 11, Ornamentanon of toxa microscleres. (C) = 06671 7 loxas absent, 2.* toxas with predominantly smooth points. 3. toxas With predominantly spined points, 574 preceed arcuate chelac in the series from palmate to bidentate- or tridentate-derived chelac. How- eve "s precor problem with dian et al. (19943) proposal is that there is no distinc- tion between these three chelae types, being one of grade (and perhaps not of clade). Frequent modifications to chelae can be seen in all three chelae morphs (curvature and thickening of the shaft, possession of unguiferous forms, reduction of alae to *tceth', fusion or detachment of alae from shaft, anchorate chelae without lateral ridges or palmate chelae with vestigial, ridge-like alae), such that the terms ‘arcuate’ and *anchorate' became a matter of degree rather than an absolute descriptor. Several Microcionidae genera have been estab- lished solely on the basis of bidentate- or triden- tate-derived isochelae, including arcuate and bidentate sigmoid chelae (Anaata, Anthoarcuata, Bipocillopsis, Damoseni, Dendrocia, Paradoryx, Qasimella, Wetmoreus) and anchorate forms (Cionanchora, Folitispa). However most do not have true arcuate or anchorate chelae (as defined by Hajdu et al., 19942), given that the lateral alae are not fully formed and still fused with the shaft for a greater proportion of their length, or the taxa do not belong in Microcionidae.. Dendrocia, Bipocillopsis and Damoseni are pivotal to the interpretation and are discussed further below. Within Microcionidae several species have dif- ferent chelae geometries but have homologous skeletal features (e.g., with megasclere geometry and skeletal stucture that indicates close affinities to each other), The Australian endemic genus Clathria (Dendrocia) has a nearly complete series of chelae extending from ‘typical’ palmate isochelae in one species (with completely fused lateral alae and straight shaft), modified palmate chelae seen in most species (i.c., with thickened curved shaft and partially detached lateral alae, verging on 'arcuale') to anchorate-like chelae seen in two species (i.e, in which there are com- pletely detached lateral alae and lateral ridges on the shaft). In other morphological characters species are very similar. The existence of this nearly complete transformation series within one genus raises the possibility that 'anchorateness' (or the detachment of lateral alae from the shaft of chelae) may have occurred more than once within the Poecilosclerida (an hypothesis dis- counted by Hajdu et al. (1994a), who suggested that anchorate and arcuate modifications to chelae were homologous within a single phylogeny comaining bidentale-derived taxa). nterpretation of this transformation continuum MEMOIRS OF THE QUEENSLAND MUSEUM in Clathria (Dendrocia) from "typical! palmate tà detached 'anchorale' chelae suggests that detach- ment of the lateral alae from the shaft of the icule, leaving the residual ‘scar’ or lateral ridge along the shaft, may be a simple reduction process pccurring more than once in the group, and that *anchorateness' may not be homologous throughout the order. Other examples, such us the strongly unguiferous sigmoid chelae of Bipacil- lopsis and Damaseni, of indeterminable arcuate or anchorate derivation, are less easily account- able and might validly be excluded from this family. However, the latter genus also has oxhorn toxas which supports its present inclusion in Microcionidae. De Laubenfels’ (19362) solution to this prob- lem (in which species with modified isochelae have skeletons and spiculation otherwise struc- turally and geometrically identical to those con- taining palmate chelae), was to assign every occurrence of a modified chela to a new genus, with the Consequence that there were nearly as many genera as species in some families, De Laubenfels’ (19362) classification had extraordi- nary high levels of homoplasy and the inferred relationships based on isochelae geometry eut across classifications based on both skeletal ar- chitecture (e.g., Hallmann, 1912, 1920) and ec- tosomal characteristics (e.g., Van Soest, 19845). This proposal was rejected by most contemporary authors. Hooper (19903) provided an alternalive proposal that considered modified isochelac to be homoplastic, and a classification based primarily on this feature was both unparsimonious and in- congruent with other structural and geometnc features within the Poecilosclerida. Hajdu et al. (1994a) correctly noted that many cases of "ercuateness' and “anchorateness’ in the literature of Poecilosclerida have not been subsequently substantiated by re-examination of original material using techniques other than routine light microscopy (e.g. many tumed out to be merely modified palmate forms), and this is also true far most recorded instances within the Microcionidae. However, from evidence presented here itis likely that ‘anchorateness” and ‘arcuateness’ may be a homoplasy for the Poecilosclerida, and consequently Hajdu et al. (19943) major reorganisation of the Poecilo- sclenda requires further refinement, as to family composition. (7) Toxa morphology: Van Soest et al. (1991) set à precedent for interpretation of loxi geometry in which oxhom-like toxas (including wing-shaped toxas) were considered to be an- REVISION OF MICROCIONIDAE cestral because they are shared by outgroups such Myxillidae, Whereas aecolada-like toxas (includ- ing raphidiform and oxeole toxas) are more deri ved. (8) Growth form: Microcionids in areas nf en- vironmental extremes, or cryptic habitats where competition for space and other resources may be high, may show a trend in reduction of skeletal characters. In these cases it is common to find fibres reduced to a simple, heavy basal layer of spongin lying on the substrate (e.g., C. (Microciona)). Although some relict (possible ancestor) groups such as 'sclerosponges' have a similar growth Form, particularly those species which lose their basal calcareous skeletons (e.g., deficient Merlia), it 1s considered here that in most cases am encnisting habit and à reduced basal skeleton is a derived condition and adaptive strategy, enabling colonization and survival in intertidal and cryptic habitats. Moreover, it is certain that this feature has arisen independently many times and in many different sponge groups. Within Microcionidae there are several specialised growth forms, such as ‘honeycomb’ reticulate characteristic of Holopsamima, with may have some systemalic value, whereas other growth forms occur throughout Ponfera and arc more difficult to interpret phylogenetically. Phylogenetic analysis (Fig. 315) shows two fundamental groups of genera (A, B) differen- tiated primarily by their skeletal structure (char- acter 6), complexity or differentiation of the skeleton (character 4), and presence or absence, respectively, of a secondary renieroid reticulate skeleton overlaying the primary (reticulate, plumo-reticulate, plumose or hymedesmoid) skeleton (character 5). One group (B) containing Artemisina, Antho and Echinoclathria has sug- gested myxillid-like features, most possessing a secondary renicroid skeleton. The other group (A) containing Clathria, Echinochalina, Holop- samma and Pandaros hàs suggested similarities to raspailiids, including skeletal structure of typi- cal genera (e.g., compare Clathria (Axociella) and Raspailia) and atypical genera (e.g. compare Echinochalina (Echinochalina) and Echinodic- tyum), It is suggested that the "raspáiloid' group (A) retains more ancestral features common te the outgroup, whereas the *myxilloid' group (B) is more denved. Within group (A) there are three taxa indicated as possibly polyphyletic (C. (Wilsenella), C (Dendrocia), Pandaros), The first two may be simply more derived than other Clatfiria at the base of the tree, or they may represent full genera. 375 The latter explanation ts rejected given that the characters inferring polyphyly (partial substito- tion of coring megascleres by detritus in C. (Wil- sonella), and loss of principal spicules completely in C. (Dendrocia)) are homoplasious representing convergences via functional ác- quisition or secondary loss of particular features rather than significant apomorphies, and their status is recognised only at the subgenus level. The third taxon, Pandaros, is incertae sedis. It is an anomolous, monotypic genus with greatly reduced fibre and spicular characteristics, nol represented in the Indo-Pacific, and probably best considered a highly modified Clathria-like species stemming from the Clathria group. The use of subgeneric taxa within this clas- sification, following the precedent adopted for the Raspailiidae (Hooper, 1991), is admittedly partially artificial given the existence of these homoplasies, but no alternative is presently avail- able that allows both the construction of a sound phylogenetic hypothesis as well as the production of a useful, working classification for this large family of sponges (given especially the relatively poor character set thal exists for sponges in general). Within Antho and Clathria the use of subgeneric taxa provides a convenient means to manage large groups of species within these genera, despite the possibility that some of the subgenene criteria are of dubious phylogenetic value (e.g., possession of acanthose strongyles in place of acanthose styles in A. (Antho) and A. (Plocamia), respectively; presence of detritus within fibres in C. {Wilsonella)y; encrusting habit and plumosc fibre nodes in C. (Microciona)). Conversely, other subgenera have a more sound phylogenetic basis and are more easily justified within the classification presented here (e.g., stylote versus oxeote structural megascleres in £ (Echinochalina) and E. (Protephlitaspongia), presence or absence of ectosomal specialisation in C. (Thalysias) and C, (Clathria)). Within Clathria there are many species-groups that could be used 1o subdivide these taxa further (e.g., ‘spicata’, ‘procera’ 'phorbasiformis" groups; see also Hooper ct al., 1991; Hooper & Lévi, 1994), Many of these groups contain species that span across several subgenera and as such are of limited usefulness in phylogenetic reconstruc- tion, but they are most useful in interpretation of sister-group relationships in a biogeographical context (Hooper & Lévi, 1994), and these species groups will he considered further in a more detailed study Indo-west Pacific microeionids, 576 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 48. List of species included in Microcionidae with their current taxonomic assignments. SPECIES CURRENT ORIGINAL GENUS abietina Lamarck abrolhosensis Bp.nov. abyssorum r acanthifolium Duchassaing & Michelotti acanthodes Hentschel acanthostyli Hoshino acanthotoxa Stephens acanthotoxa Lévi & Lévi [preocc.] aceratoobtusa Carter aculeata Ridley adioristica de benfels affinis Carter affinis Topsent [preocc.] africana Lévi alata Dendy amabilis Thiele ambigua Bowerbank amiranigiensis nom.nov. anchorata Carter anchoratum Carter angularis Sarà & Siribelli angulifera Dendy angulosa Duchassaing & Michelotti anomala Burton anomala Hallmann anonyma Burton Spongia Dictyacylindrus Pandaros Clathria Thalysias Eurypon Micraciona Micraciona Clathria Dictyociona Microciona Hymeraphia Microciona Clathria Stylotellopsis Microciona [for Colloclathria ramosa Dendy] Dictyocylindrus Echinonema Microciona Clathria Pandaros Rhaphidophlus Echinochalina Microciona ASSIGNMENT C. (Thalysias) C. (Wilsonella) Synonym of (Antho) dichotoma (Esper) Pandaros synonym of C. (Thalysias) cactiformis (Lamarck) C. (Clathria) C. (Clathria) sce C. (Microciorui) claudei nom.nov. C. (Microciona) synonym of ral (Thalysias) abietina (Lamarck) C. (Microciona) C, (Micraciona) see C. (Microciona) campécheae nom.nov. C. (Microciona) synonym of à (Dendrocia) prenda endenfeld C. (Thalysias) Plocamionida (Anchinoidae) C. (Thalysias) C. (Clathria) synonym of à (Thalysias) cactiformis (Lamarck) C. (Microciona) C. (Clathria) Mycale (Mycalidae) C. (Thalysias) E. (Echinochalina) C. (Microciona) antarctica Topsent antarcticus Koltun [preocc.] anthoides Lévi antyaja Burton & Rao aphylla sp.nov. apollinis idley & Dendy appendiculata Lamarck uraiosa Hooper & Lévi arborea Tanita arborea Lendenfeld arborescens Ridley arbuscula Row arbusculum Duchassaing & Michelotti archegona Ristau arcifera Schmidt arciger Schmidt arcuophora Whitelegge arenifera Carter armata Bowerbank arteria de Laubenfels aruensis Hentschel ascendens Cabioch asodes de Laubenfels assimilis Topsent atoxa Bergquist & Fromont ailantica Sarà atrasanguinea Bowerbank australiensis Carter australiensis Ridley Anchinoe Stylotellopsis Clathria Dendrocia Amphilectus Spongia Clathria (Thalysias) Litaspongia Plectispa Rhaphidaphlus Ophlitaspongia Pandaros Artemisina Tenacia Suberites Clathria Echinoclathria Microciona Axociella Hymeraphia Microciona Eurypon Clathria Dictyociona Echinoclathria Microciona Wilsonella Ophlitaspongia C. (Microciona) synonym of (Thalysias) koltuni Hooper C. (Clathria) C. (Clathria) C. (Thalysias) Artemisina Thalysias) C. (Thalysias) Echinoclathria Holopsamma C. (Thalysias) C. (Clathria) Ptilocaulis (Axinellidac) Artemisina Echinoclathria Artemisina C. (Clathria) synonym of olopsamma laminaefavosa Carter C. (Microciona) C. (Thalysias) C. (Thalysias) C. (Micraciona) C. (Clathria) C. (Micraciona) C. (Clarhria) Echinoclathria C. (Microciona) C. (Wilsonella) E. (Echinochalina) australis Lendenfeld australis Whitelegge | axinelloides ndy axociona Lévi barba Lamarck barbadensis Van Soest barleei Bowerbank bargibanti | Hooper & Lévi basiarenacea Boury-Esnault basifixa Topsent basilana Lévi basispinosa Burton beanii Bowerbank bergquistae sp. nov. beringensis Hentschel biclathrata nom.nov, bihamigera Waller bispiculata Dendy bispinosus Whitelegge bitoxa Burton bitoxifera Koltun brattegardi Van Soest & Stone brepha de Laubenfels brevispina Lendenfeld brondstedi nom.nov. rondsted] borealis nom.noy. REVISION OF MICROCIONIDAE RIGINAL GENUS Clathria Plumo- halichondria Ophlitaspongia Clathria barba Plocamilla Isodicrya E. (Prato- phlitaspongia) Rhaphidophius Ophlitaspongia Clathria Microciona Isodictya Phakellia |for Microciona clathrata Whitelegge] Microciona Siphonachalina Rhaphidophlus Hymantho Axociella [for Clarhria robusta Koltun] Antho Aaala Thalassodendron [for Hymedesmia ennata pr" CURRENT SPECIES — O "MEINEN synonym of rella incrustans var. arenacea (Carter) (Crellidae) synonym of rella incrustans (carier) Crellidae) Echinoclathria C. (Clathria) E. (Echinochalina) Antho (Plocamia) C: (Clathria) E. (Proto- phlitaspongía) C. (Thalysias) C. (Microciona) C. (Clathria) synonym of . (Microciona) antarctica (Topsent) synonym of A. (ndha) involvens Schmidt) Echirioclathria Echinoclathria C. (Clathria) Pronax (Anchinoidae) E. (Proto- phlitaspongia) synonym of . (Thalysias) lendenfeldi Ridley & Dendy C. (Microciona) C. (Thalysias) C. (Clathria) A.(Antho) C. (Micraciana) synonym of .( Thalysias) cactiformis (Lamarck) C. (Microciona) burtoni Lévi cactiformis Lamarck caelata Hallmann caespes Ehlers caespilosa Carter calla de Laubenfels calochela Hentschel calopora Whitelegge calypsa Boury-Esnault campecheae nom.rnov. canaliculata Whitelegge cancellaria Lamarck cantabrica Orueta carbonaria Lamarck coriocrassus Bergquist & Fromont carnosa Bowerbank carteri Topsent carteri Ridley & Dendy cercidochela Vacelet & Vasseur cervicarnis Thiele chalinoides Carter chartacea (Whitelegge) chelifera Hentschel chelifera Lévi [preocc.] circonflexa Lévi Microciona Clathria Clathria (Microciona) [for Clathria prolifera Burton] Plocamilla Spongia Clathria Scapalina Echinonema Axociella Hymeraphia Clathria Clathria [for Hymeraphia affinis Topsent] Esperiapsis Spongia Rhaphidophlus Spongia Rhaphidophlus Microciona Clathria Echinoclathria Clathriopsamma Rhaphidophlus Axinella Clathria Spanioplon Microciona Plocamilla Antho (Plocamia) 577 A. (Antha) C. (Microciona) C. (Clathria) C. (Microciona) C. (Clathria) Antho (Plòcamia) C. (Thalysias) C. (Clathria) unrecognisable Phanohalichondria (Anchinoidae) C. (Microciona) C. (Thalysias) C, (Clathria) C. (Clathria) C. (Microciona) C. (Axociella) C. (Thalysias) C. (Clathria) Haliclona (Chalinidae) C. (Thalysias) C. (Microciona) synonym of . (Clathria) foliacea Topsent synonym of olopsamma favus (Carter) C. (Wilsonella) C. (Thalysias) Echinoclathria Antho (Isopenectva) C. (Clathria) sec C. (Microciona) tunisiae nom.nov. 578 SPECIES cladoflagellata Carter MEMOIRS OF THE QUEENSLAND MUSEUM ORIGINAL GENUS Axinella clathrata Schmidt Tenacia claudei nom.nov. cleistochela Topsent |for Microciona acantholoxa Lévi & Lévi] Clathria coccinea Berguist Microciona coccinea Duchassaing & Michelotti collata sp.nov. conectens Hallmann compressa Schmidt concentrica Lendenfeld confragasa Halimann conica Lévi contexta Sara contorta Bergquist & Fromont copiosa Topsent capiosa var. curacaoensis Amdt Thalvsias Wilsonella Clathria Antherochalina Ophlitaspongia Clathria Echinoclathria Dictyociona Clathria Clathria coppingeri Ridley Clathria coppingeri var. aculeata carallitincta Dendy coralloides Olivi corallorhizoides Fristedt coriacea Bowerbank corneolia Hooper & Lévi corona Lieberkiihn Clathria Clathria Spongia Clathria Isodictya Clathria (Thalysias) Halichondria CURRENT ASSIGNMENT synonym of Y hingclathria chalinoides (Carter) synonym of d (Thalysias) virgultosa Lamarck) C. (Microciona) C. (Microciona) C. (Microciona) Spirastrella (Spirastrellidae) E. (Proto- phlitaspongia) C. (Clathria) C. (Clathria) Cymbasiela (Axinellidae) Echinoclathria C. (Clathria) Echinoclathria C. (Clathria) synonym of - (Thalysias) virgultosa (Lamarck) synonym of c (Thalysias) schoenus (dc Laubenfels) C. (Thalysias) Syn of | . (Thalysias) lendenfeldi Ridley & Dendy synonym of e ( Thalysias) vulpina (Lamarck) C. (Clathria) synonym of dendari complicata (Lundbeck), (Myxillidae) Aniho (Plocamia) C. (Thalysias) synonym of . (Clathria) coralloides (Olivi) corticata var. elegans Lendenfeld clathrata Whitelegge [preocc. claviformis Hentschel cliftoni Beet nk coralliophilus Thiele costifera Hallmann craspedia sp. nov. crassa Lendenfeld crassa Carter cratitia Esper cienichela Alander curvichela Hallmann curvichela Vacelet & Vasseur [preocc.] cullingworthi Burton curvispiculifera Carter cylindrica Kidley & Dendy &tlindrica sensu im & Byeon cylindricus Kieschnick [preocc.] darwinensis sp.nov. dayi Lévi decumbens Ridley delaubenfelsi Lévi delaubenfelsi Little delicata Lambe dendyi Berquist & Fromont densa Burton dentata Topsent depressa Sigmatella Microciona Clathria Hymeniacidon Rhaphidophlus Clathria Antherochalina Holopsamma Spongia Microciona Wilsonella Microciona Clathria Microciona Esperiopsis Axociella Rhaphidophlus Clathría Clathria Rhaphidophlus Holoplocamia Clathria Microciona Microciona Clathria Clathria synonym of e (Wilsonella) australiensis (Carter) see C. (Clathria) biclathrata nom.nov, C. (Wilsonella) synonym of ntho (Plocamia) frondifera (Lamarck) C. (Thalysias) C. (Thalysias) C. (Thalysias) C. (Clathria) Halopsamma C. (Thalysias) C. (Microciona) C. (Dendrocia) see C. (Microciona) vaceletlia nom.nov. C. (Thalysias) ? C. (Clathria) virtually | unrecognisable C. (Axociella) see C (Microciona) simae sp.nov. synonym of . (Thalysias) kieschnicki Hooper C. (Thalysias) C. (Clathria) C. (Clathria) C. (Thalysias) Antho ( Plocamia) synonym of - (Clathria) | prolifera (Ellis & olander) C. (Microciona) C. (Micraciona) sy nonym of - (Thalysias) fascicularis ‘opsent C. (Clathria) Sara & Melone ORIGINAL GENUS REVISION OF MICROCIONIDAE CURRENT ASSIGNMENT dianae Schmidt dianae Topsent dichotoma Esper dichotoma Lévi diechinata Hallman digitata Lendenfeld digitiformis Row discreta Thiele distincta Thiele ditoxa Stephens dives Topsent dubia Kirkpatrick duplex Sarà dura Whitelegge dura var. mollis Hentschel eccentrica Burton Suberites Artemisina Spongia Ophlitaspongia Clathria Thalyssodendron Ophlitaspongia Microciona Hymeraphia Eurypon Microciona Microciona Clathria Clathria Clathria Ophlitaspongia echinata Alcolado Axociella echinonematissima Wilsonella Carter egena iedenmayer elastica Lévi elastica Sara [preocc.] elegans Vosmaer elegans Lendenfeld elegans Lendenfeld elegans Ridley & Dendy elegantula Ridley & Dendy elegantula Dendy elliptichela Alander encrusta Kumar ensiae sp.nov. erecta Ferrer- Hernandez erecta Topsent Echinoclathria Clathria Clathria Clathria Antherochalina Plectispa Plocamia Clathria Artemisina Microciona Clathria Plocamia Artemisina C. (Microciona) synonym of rtemisina apollinis (Ridley Dendy) A.(Antho) Echinoclathria synonym of ras (Thalysias) lendenfeldi Ridley & Dendy Echinoclathria Echinoclathria C. (Clathria) C. (Thalysias) C. (Microciona) ? Plumo- halichondria (Anchinoidae) C. (Thalysias) C. (Microciona) C. (Dendrocia) synonym of . (Clathria) squalorum jedenmayer C. (Isociella) C. (Microciona) C. (Clathria) Echinoclathria C. (Clathria) see C. (Clathria) sarai nom.nov. ? C. (Clathria) virtually unrecognisable Raspailia (Raspailiidae) Holopsamma Antho (Plocamia) C. (Dendrocia) Artemisina C. (Microciona) C. (Thalysias) C. (Wilsonella) Antho (Plocamia) Artemisina erectus Thiele eurypa de Laut nfels fallax Bowerbank fascicihiris opsent fasciculata ilson fascispiculifera arter |fauroti Topsent favosa Whitelegge /favosa Lamarck favulosa sp.nov. favus Carter favus var. arenifera Carter felixi sp.nov. ferrea de aubenfels fetita owerbank filifer Ridley & Dendy filifer var. cantabrica Orueta filifer var. mutabilis Topsent filifer var. spiniferü Lindgren flabellata Topsent abellata iley & Dendy flabellata Burton fiabellifera i ooper & Lévi flabelliformis Carter foliacea Topsent fpliascens acelet & Vasseur foraminifera Burton & Rao Rhaphidophlus Dictyociona Microciona Clathria Clathria Microciona Axosuberites Clathria Spongia Echinoclathria Echinoclathria Fisherispongia Microciona Rhaphidophlus Rhaphidophlus Rhaphidophlus Rhaphidophlus Ophlitaspongia Phakellia Rhaphidophlus Clathria (Thalysias) Echinonema Clathria Clathria Aulenella 579 C. (Thalysias) C. (Thalysias) C. (Microciona) C. (Thalysias) C. (Thalysias) C. (Microciona) C. (Axociella) synonym of c (Thalysias) cactiformis (Lamarck) synonym of . (Echinochalina) barba (Lamarck) E. (Proto- phlitaspongia) Holopsamma synonym of olopsamma laminaefavosa Carter E. (Echinochalina) C. (Wilsonella) Phorbas (Anchinoidae) C. (Thalysias) C. (Clathria) cantabrica (Orueta) C. (Thalysias) mutabilix (Topsent) see C. (Thalysias) spinifera (Lindgren) synonym of e (Axociella) nidificata (Kirkpatrick) synonym of e (sociella) macropora Lendenfeld C. (Clathria) C. (Thalysias) synonym of . (Thalysias) cactiformis (Lamarck) C. (Clathria) C. (Clathria) C. (Wilsonella) 580 jrondifera var. dichela Hentsche frondifera var. | major Hentschel coides owerbank fusterna sp.nov. gabrieli Dendy georgiaensis nom.nov. igantea ndenfeld glabra Ridley & ndy globose ndenield Rürgonoides ndy gracea Bakus racilis idley racilis arer gradalis Topsent grisea Hentschel ymnazusa chmidt haematodes de hallmanni sp.nov son Axocielita MEMOIRS OF THE QUEENSLAND MUSEUM RIGINAL GENUS " ] CURRENT SPECIES gi ASSIGNMENT Microciona Microciona [for Axociella toxitenuis Bergquist & Fromont] Reniera Spongia Halichondria Clathria 1 Clathria Clathria Ophlitaspongia Ophlitaspongia [for Ophlita- spongta thielei urton] Halme Echinoclathria Halme Echinodictyum Anthoarcuata Echinonema Echinoclathria Clathria Leptosia Placamia Microciona Heteroclathria haplotoxa Topsent Leptoclathria C. (Microciona) C. (Microciona) C. (Axaciella) C. (Clathria) Antho (Plocamia) | synonym of . (Thalysias) vulpina (Lamarck) synonym of d (T hal 'sias) vulpina (Lamarck) synonym of a (Thal sias) vulpina (Lamarck) see C, (Thalysias) major Hentschel Terpiosella (Suberitidae) C. (Thalysias) E. (Echinochalina) C. (Axociella) synonym of olopsamma laminaefavosa Carter synonym of E. aeria Ese, rba (Lamarck) synonym of olopsamma crassa Carter C. (Clathria) A.(Antho) synonym of ral { Thalysias) procera (Ridley) synonym of chinoclathria subhispida Carter C. (Microciona) C. (Microciona) Antha (Plocamià) C. (Microciona) A.(Antho) C. (Thalysias) C. (Microciona) C.(Thalysias) | hartmeyeri | Hentschel hechteli nom.nov. | hentscheli nom.nov. | hesperia sp.nov. | heterospiculata | Brondsted Duchassaing & Michelotti hispanica Ferrer- Hernandez hirsuta Hooper & Lévi hispidula Ridley hjorti Arnesen horrida Row | hymedesmioides an Soest ignis —— uchassaing & Michelotti | illgi Bakus | imperfecta Dendy inanchorata | Ridley & Dendy | incrustans Bergquist incrustans Carter incrustans _ Svarceyskij indica Dendy. indica Thomas indistincta Bowerbank indurata Hallmann inhacensis Thomas inornata | Hallmann Clathria [for Microciona microchela Hechtel] [for Hymeraphia e ndenfeldi Hentschel] Microciona Microciona Clathria Thalysias Artemisina Clathria (Thalysias) Amphilectus Echinoclathria Ophlitaspongia Clathria (Microciona) Thalysias illawarrae sp.nov. - Plocamilla Clathria Clathria Isociella Echinonema Raspailia Clathria Oasimella Hymedesmia Clathria Clathria Ophlitispongia Clathria C. (Clathria) eyhonymi of .(Aniho) tuberosa (Hentschel) C. (Thalysias) C. (Microciona) C. (Thalysias) A (Antho) C. (Microciona) C. (Clathria) unrecognisable Artemisina C. (Thalysias) C. (Clathria) Echinoclathria C. (Clathria) C. (Microciona) Tedania (Tedaniidae) C. (Microciona) Antho (Plocamia) C. (Dendrocia) C. (Clathria) C. Usociellà) Crella (Crellidae) ? synonym of A. ntho) involvens (Schmidt) C. (Clathria) Artemisina synonym of locamionida ambigua (Bowerbank) Anchinoidae) synonym of . (Thalysias) cactiformis (Lamarck) C. (Clathria) Echinoclathria SPECIES ORIGINAL GENUS intermedia Whitelegge intermedia Burton intexta Carter involvens Schmidt irregularis Burton irregularis Lendenfeld isaaci sp.nov. isochelifera Uriz isodictyoides Van Soest ixauda Lévi cesar ndy jecusculum owerbank johnsoni de ubenfels jolicoeuri Topsent jovis Dendy ugosa ilson juncea Burton juniperina (var. A) Lamarck Jieupering (var. ) Lamarck kasumiensis Tanita kentii Bowerbank kieschnicki Hooper kilauea de Laubenfels koltuni Hooper lista ooper & Lévi laboutei Hooper & Lévi labyrinthica Schmidt laciniosa Bowerbank & Norman laevigata Lambe Echinoclathria Paresperia Microciona Myxilla Marleyia Halme Echinoclathria Rhaphidophlus Microciona Phakellia Hymeniacidon Cornulum Rhaphidophlus Artemisina Clathria Clathria Spongia Spongia Raspailia Microciona Clathria (Thalysias) Axocielita Clathria (Thalysias) Clathria E. (Proto- phlitaspongia) Reniera Isodictya Clathria REVISION OF MICROCIONIDAE CURRENT ASSIGNMENT E. (Echinochalina) Esperiopsis (Desmacididae Rhabderemia (Rhabderemiidae) A.(Antho) C. (Clathria) synonym of olopsamma laminaefavosa Carter E. (Proto- phlitaspongia) E. (Echinochalina) C. (Thalysias) C. (Microciona) synonym of C (Isociella) macropora Lendenfeld C. (Microciona) Cornulum (Coelosphaeridae) C. (Thalysias) Artemisina synonym of a (Thalysias) virgultosa (Lamarck) C. (Clathria) C. (Thalysias) synonym of ral (Thalysias) coppingeri Ridley ? Pandaros uncertain placement C. (Microciona) C. (Thalysias) C. (Thalysias) C. (Thalysias) C. (Clathria) E. (Proto- phlitaspongia) C. (Clathria) synonym of C. (anri barleei Bowerbank) C. (Clathria) laevis Bowerbank Microciona C. (Microciona) laevissima Dendy Hymedesmia C. (Microciona) lore: ced Anaata C. (Microciona) lambda Lévi Leptoclathria C. (Thalysias) lambei Koltun Microciona C. (Axociella) lambei Burton Heteroclathria Antho (Plocamia) laminaefi AVOA, Holopsamma Holopsamma i s pym of axa olopsamma Lendenfeld Halme laminaefavosa Carter leighensis [for Microciona , . nom.nov. rubens Bergquist] C. (Microciona) for Microciona lematolae [ ; lacenta sensu de C. (Thalysias) Sp;nov. Laubenfels] leporina Lamarck Spongia Echinoclathria tevi & Siribelli Microciona C. (Microciona) levii sp. nov. - Echinoclathria levis Echi s mila ym of , chinonema rella incrustans Lendenfeld (Carter) (Crellidae) Kidiey & Dendy Clathria C. (Thalysias) lendenfeldi see C. (Microciona) Hentschel Hymeraphia hentscheli [preocc.] nom.nov. inane : Axocielita C. (Thalysias) lindgreni [for Clathria A nom.nov. ramosa Lindgren] C. (Wilsonella) lipochela Burton Clathria C. (Clathria) lissocladus Burton Rhaphidophlus C. (Thalysias) lissosclera Bergquist & Clathria C. (Clathria) Fromont lithophoenix de ; Laubenfels Plocamia A.(Antho) litos Clathria ; Hooper & Lévi — (Clathriopsamma) ©: (Wilsonella) lizardensis sp.nov. - C. (Microciona) lobata Vosmaer Clathria C. (Clathria) lob e (QV ise f obosa ] . (Wilsonella) Lendenfeld Clathriopsamma australiensis (Carter) . f longichela A s "Cla a Clathria . (Clathria Topsent anchorata (Carter) fougisp iculum Microciona C. (Microciona) longistyla Burton Microciona C. (Microciona) lorgiroxá, Hymeraphia C. (Thalysias) loveni . Mycale Fristedt Clathria (Mycalidae) 581 582 SPECIES Duchassaing & Michelotti macrochela Lévi macroisochela Lévi macropora Lendenfeld macropora Lendenfeld macratoxa Bergquist & Fromont madrepora Dendy maeandrina Ridley major Hentschel manaarensis Carter marissupéri Pulitzer-Finali massalis Carter maunaloa de Laubenfels mediterranea Babic melana Van Soest & Stentoft membranacea Thiele menoui Hooper & Lévi meyeri Bowerbank michaelseni Hentschel microchela Stephens microchela Hechtel [preocc.] microcionides Carter microjoanna de Laubenfels micronesia de Laubenfels micropora Lendenfeld micropunctata Burton & Rao MEMOIRS OF THE QUEENSLAND MUSEUM ORIGINAL GENUS Pandaros Microciona Clathria Claihria Plectispa Axociella Clathria Clathria Clathria Dictyocylindrus Clathria Thalysias Microciona Artemisina Echinachalina melana Van Soest Artemisina Ophlitaspongia Clathria Ophlitaspongia Hymeraphia Eurypon Microciona Plumo- halichondria Microciona Microciona Halme Tenacia Clathria CURRENT ASSIGNMENT Echinodictyum (Raspailiidae) C. (Microciona) C. (Clathria) C. (Isociella) Holopsamma C. (Axociella) synonym of . (Clathria) spongodes Dendy C. (Clathria) C. (Thalysias) Antho (Plocamia) C. (Claihria) Reniera (Chalinidae) C. (Thalysias) synon mof .(Antho) involvens (Schmidt) E. (Echinochalina) Artemisina C. (Thalysias) C. (Clathria) C. (Clathria) C. (Thalysias) C. (Clathria) see C. (Thalysias) hechteli nom,nov. Plocamionida (Anchinoidae) C. (Microciona) C. (Microciona) synonym of olopsamma crassa Carter C. (Thalysias) C. (Clathria) microxea Vacelet & Vasseur mima de Laubenfels minor Burton minuta Van Soest minutula Carter mixta Hentschel mollis Kirkpatrick monticularis Ridley & Dendy morisca Schmidt morlensii Brondsted mosulpia Sim & Byeon mulabilis Topsent multiformis Whitelegge multipes Hallmann multipora Whitelegge multitoxaformis Bergquist & Fromont murphyi sp. nov. mutans Sara mutula Bowerbank myxilloides Dendy naikaiensis Hoshino namibiensis Uriz. nervosa Lévi nexus Koltun nidificala Kirkpatrick nidus-ves, Lendenfeld rum noarlungae sp.nov. nodosa Carter normani Burton notialis sp. nov. noto Tanita novaezealandiae Brondsted Parátenaciella Ophlitaspongia Ophlitaspongia Rhaphidophlus Microciona Clathria Clathria Axinella Clathria Clathria Clathria Rhaphidophlus Ceraochalina Clathria (Plectispa) Clathria Axociella Ophlitaspongia Halichondria Clathria Eurypon Microciona Axociella Bipocillopsis Ophlitaspongia Halme Echinoclathria Hymantho Ophlitaspongia Microciona C. (Microciona) C. (Microciona) Echinoclathria C. (Thalysias) Rhabderemia (Rhabderemiidae) C. (Wilsonella) Pranax (Anchinoidae) Aulospongus (Raspailiidae) synonym of A. (Antha) involvens (Schmidt) C. (Clathria) C. (Clathria) C. (Thalysias) synonym of | Echinoclathria subhispida Caner C. (Clathria) synonym of C. (Thalysias) rubra (Lendenfeld) C. (Axociella) C. (Clathria) Echinoclathria synonym of C. (Clathria) barleei (Bowerbank) C. (Dendrocia) C. (Thalysias) C. (Microciona) C. (Thalysias) C. (Clathria) C. (Axociella) synonym of Holopsamma laminaefavosa Carter C. (Clathria) Echinoclathria C. (Microciona) Echinoclathria Echinoctathria C. (Microciona) CURRENT SPECIES ORIGINAL GENUS ASSIGNMENT Raley ica Dirrhopalum Antho (Plocamia) nuda Hentschel Clathria C. (Thalysias) obli i pohregil Esperiopsis C. (Clathria) oculata Burton Clathria C. (Clathria) resi Axociella C. (Thalysias) QUUM ad Alcyonium A. (Antho) orientalis Rhaphidophlus C. (Thalysias) originalis de Esperiopsis C. (Thalysias) ornata Dendy Bubaris Antho (Plocamia) oroides Schmidt Clathria Agelas (Agelasidae) osismica Cabioch Microciona C. (Microciona) s no m of ) ostacina ; . (Clathria Rafinesque Spongia rolifera (Ellis & Solander] aene Bernat Ophlitaspongia Echinoclathria oxeata Proto- E. (Proto- Burton phlitaspongia phlitaspongia) gxeifera Ferrer- Clathria A.(Antho) oxeotus Van Soest Rhaphidophlus C. (Thalysias) oxitoxa Lévi Clathria C. (Thalysias) oxneri Topsent Hymedesmia C. (Clathria) d diced Wilsonella C. (Clathria) pachyaxia Lévi Axociella C. (Thalysias) pachystylaLévi Clathria C. (Clathria) A f apilla Dunn . synonym of phlitaspongia . (Microciona) Bowerbank ien (Grant) papillosa Thiele Clathria C. (Clathria) synonym of PUE Phakellia chinoclathria leporina ? synonym of A. Va Artemisina (Artho) involvens (Schmidt) parkeri sp. nov. — - Echinoclathria arinena de Microciona C. (Microciona) partita Hallmann Clathria C. (Clathria) parva Lévi Clathria C. (Axociella) patula sp.nov. - C. (Axociella) påuorspicula Rhapidophlus C. (Clathria) UA synonym of Dendehfeld Thalassodendron c (Thalysias) REVISION OF MICROCIONIDAE rubra (Lendenfeld) paucispina Sarà & Siribelli pauper Brondsted pectiniformis arter pellicula itelegge elligera Kchmidt pennata Lambe ennata rondsted [preocc.] ennata — uchassaing & Michelotti penneyi de ubenfels erforata forat g erforata ìn part ndenfeld y phorbasiformis sp.nov. ilosus idley & Dendy placenta Lamarck lacenta de aubenfels [preocc.] planum Carter plena Sollas linthina de aubenfels lumosa ontagu lumosa entschel jin litzer-Finali lurityla ulitzer-Finali piniformis Carter plana Carter poecilosclera Sarà & Siribelli polita Ridley prima Brondsted primitiva Koltun primitiva Burton procera Ridley Antho Clathria Echinonema Clathria Clathria Desmacella Hymedesmia Pandaros Holoplocamia Antherochalina Antherochalina Amphilectus Spongia Microciona Microciona Plocamia Microciona Spongia Artemisina Echinoclathria Clathria Dictyocylindrus Microciona Microciona Hymedesmia Lissoplocamia Microciona Clathriella Rhaphidophlus 583 A.(Antho) C. (Clathria) synonym of g (Thalysias) cactiformis (Lamarck) C. (Clathria) Dictyonella (Dictyonellidae) C. (Microciona) see C. (Microciona) brondstedi nom.nov. Echinodictyum (Raspailiidae) Antho (Plocamia) synonym of Antho (/sopenectya) chartacea (Whitelegge) C. (Clathria) C. (Thalysias) Megaciella (lophonidae) C. (Thalysias) see C. (Thalysias) lematolae sp.nov. C. (Microciona) Antho (Plocamia) C. (Microciona) Pronax (Anchinoidae) Artemisina Holopsamma C. (Clathria) C. (Clathria) synonym of c (Thalysias) virgultosa (Lamarck) C. (Microciona) C. (Microciona) Antho (Plocamia) C. (Microciona) Antho (Isopenectya) C. (Thalysias) 584 procumbens sensu Brondsted raductitoxa oshino grolifera Ellis & olander prolifera Burton roxima | Lundbeck roxima | uchassaing & Michelotti pecans ade aubenfels pugio Lundbeck punicea sp.nov. pustulosa Carter yramida ndenfeld prs rondsted uadriradiata | Carter yyerolfplia ller uinqueradiata arter rameus Koltun ramosus. Kieschnick ramosa Lindgren [preocc.] ramosa Dendy [preocc.] | ramosa Hallmann ramsayit Lendenfeld raphanus Lamarck raphida sensu Hechtel raraechelae Van Soest MEMOIRS OF THE QUEENSLAND MUSEUM | ; CURRENT SPECIES ORIGINALGENUS “oC Clathria Thalysias Spongia Clathria Hymedesmia Thalysias Clathriopsamma Hymedesmia Microciona Halichondria Clathria Microciona Microciona Antherachalina Microciona Axociella Rhaphidophlus Clathria Colloclathria Echinoclathria Thorecta Spongia Clathria Rhaphidophlus _venosa (Alcolado) synonym of Ascetta proce ndenfeld (Calcarea) C. (Clathria) C. (Clathria) see C. ( Clathria) burtoni nom.nov. C. (Microciona) Xestospongia (Petrosiidae) C. (Wilsonella) C. (Microciona) Antho (Isopenectya) synonym of habderemia minutula (Carter) (Rhabderemiidae) C. (Microciona) C. (Dendrocia) C. (Clathria) C. (Microciona) Phakellia (Axinellidae) Cyamon (Raspailiidae) synonym of i (Aiociella) nidificata (Kirkpatrick) C. (Thalysias) see C. (Wilsanella) lindgreni nom.nov. see C, (Thalysias) amiranteiensis nom.nov. Holopsamma synonym of - (Wilsonella) australiensis (Carter) C. (Clathria) synonym of liona rhaphida Boury-Esnault (Clionidae) synonym of . (Thalysias) rarispinosa Hechtel rectangulosa Schmidt reinwardti Vosmaer reinwardti var. nn idley reinwardli var. subcylindrica Ridley renieroides Lendenfeld repens Duchassaing & Michelotti reticulata Lendenfeld reficulaja Bergquist & Fromont reticulata Whitelegge rhaphidotoxa Stephens rhopalophora Hentschel riddlei sp. nov. ridleyi Lindgren ridleyi Hentschel ridleyi Dendy robusta Dendy rabusta Koltun [preocc.] rotunda Hallmann rubens Lendenfeld rubens Bergquist [preocc.] rubens var. dura Lendenfeld rubens var. lamella Lendenfeld rubra Lendenfeld rugosa Hooper & Lévi rugosa Duchassaing & Michelotti sainrvincenr sp.nov. Microciona Clathria Clathria Clathria Clathria Antherochalina Thalysias Clathriopsamma Ophlitaspangia Echinochalina Clathria Hymeraphia Rhaphidophlus Plocamia Echinodictyum Microciona Clathria Echinaclathria Thalassodendron Microciona Thalassodendron Thalassodendron Echinonema Clathria (Clathriopsamma) Thalysias synonym of . (Microciona) calla (de Laubenfels) C. (Clathria) C. (Thalysias) synonym of . (Thalysias) vulpina (Lamarck) Claas reinwürdti Vosmaer An mof hakellia flabellata (Carter) (Axinellidae) synonym of estospongia subtriangularis (Duchassaing) (Petrosiidae) C, (Wilsonella) Echinoclathria E. (Echinochalina) C. (Clathria) C. (Micraciona) Echinoclathria C. (Thalystas) Antho (Plocamia) E. (Echinochalina) C. (Thalysias) see C. (Clathria) borealis nom.nov. Holopsamma C. (Clathria) see C. (Microctona) leighensis nom.nov. pony of C. (Thalysias) rubra (Lendenfeld) synonym of C. f alysias) rubra ‘Lendenfeld) C. (Thalysias) C. (Wilsonella) synonym of. Xestospongia | subtriangularis (Duchassaing) (Petrosiidae) Antho (Isopenectya) ORIGINAL GENUS REVISION OF MICROCIONIDAE CURRENT ASSIGNMENT sarai nom.nov. saraspinifera nom.nov. sartaginula Lamarck saxicava Duchassaing & Michelotti scabida Canter schoenus de Laubenfels scotti Dendy selachia sp.nov. seriata Grant seriatus Thiele sessilis Carter shirahama Tanita sigmoidea Cuartas signata Topsent simae sp.nov, similis Thiele similis Stephens [preocc, | similis sensu Uriz simplex Lendenfeld simplicissima Norman simpsoni Van Soest skia sp.nov. spicata allmann spiculosus Dendy spiculosa var. macilenta Hentschel spiculosus Var. ramosa Hentschel spinarous arter & Hope spinatoxa oshino spinifera Lindgren [for Clathria elastica Sarà] [for Clathria spinifera Sarà] Spongia Thalysias Halichondria Clathria Clathria Spongia Rhaphidaphlus Dictyocylindrus Clathria Microciona Plocamiopsis [for Axociella cylindrica sensu Sim & Byeon] Hymeraphia Microciona Eurypon Halme Microciona C. (Microciona) Clathria Rhaphidophlus Clathria Clathria Micraciona Microciona Rhaphidophlus C. (Clathria) C. (Clathria) C. (Clathria) unrecognisable C. (Dendrocia) C. (Thalysias) C. (Microciona) C. (Isociella) C. (Microciona) synonym of e (Thalysias) vulpina (Lamarck) unrecognisable C. (Clathria) C. (Microciona) Antho (Plocamia) C. (Microciona) C. (Mieraciona) see C. (Microciorui) stephensae nom.nov. see C. (Microciona) urizae nom.nov. Holopsamma Bubaris (Axinellidae) synonym of . (Microciona) echinata (Alcolado) C. (Isociella) synonym of `, (Thalysias) lendenfeldi Ridley & Dendy C. (Thalysias) synonym of r (Thalysias) reinwardtii Vosmaer synonym of . (Thalysias) procera (Ridley) C. (Microciona) C. (Microcionu) C. (Thalysias) spinifera Sarà [preocc.] gpiniypicula anita spinosa Wilson spinulenta owerbank spongigartina del Laubenfels spongiosa urton spongiosa Dendy spongodes Dendy squalorum iedeamayer stipitata Koltun striata Whitelegge strongyla Hentschel stephensae nom.nov. sirepsitoxa Hope strepsitoxa var. robusta Dendy styloprothesis sp.nov. suberitoides Vosmaer subhispida Carter subtriangularis Duchassaing surculosa Esper svarchevskyi de Laubenfels tenebratus Whitelegye tener Caner tenuifibra Whilelegge tenuis Stephens tenuis Carter lenuispina Lendenfeld Clathria Clathria Microciena Microciona Agata Clathria Echinodictyum Clathria Clathria Artemisina Clathria Artemisina [for Microciona similis Stephens] Microciona Microciona Artemisina Echinoclathria Thalysias Spongia Microciona Rhaphidophlus Thalysias Clathria Microciona Echinaclathria Antherochalina 585 see C. (Clathria) saraspinifera nom.nov. C. (Clathria) C. (Microciona) Iophon (lophonidae) C. (Microciona) synonym of . (Clathria) spongodes Dendy E. (Echinochalina) C. (Clathria) C. (Clathria) Artemisina C. (Clathria) synonym of Artemisina numasa entschel C. (Microciona) C. (Microciana) C. (Thalystas) robusta (Dendy) C. (Thalysias) synonym of rtemisind — arciger (Schmidt) Evhinoclathria Xestospongia (Petrosiidae) ? C. (Clathria) virtually unrecognisable synonym of . (Microciona) armata (Bowerbank) synonym of . (Clathria) striata Whitelegge "7 C. (Thalysias) imperfectfy known Anonym of C. (Clathria) rubens (Lendenteld) C. (Microciona) synonym of Echinaclathria leporina (Lamarck) synonym of Echinoclathria leporina (Lamarck) 586 MEMOIRS OF THE QUEENSLAND MUSEUM a mmea ania SPECIES ORIGINAL GENUS ASSIGNMENT tenuissima Stephens terranavae Dendy tetrastyla Hentschel textile Carter thetidis Hallmann thielei Hentschel thielei Burton [preocc.] lingens sp.nov, topsenti Thiele toriuosa Uriz toxifera Hentschel toxiferuim Topsent [preocc, ] taxtmujor Topsent toxipraedita Topsent loxirecta Sarà & Siribelli Toxistricta Topsent toxistyla Sarà Ioxitenuis Bergquist & loxivaria Sarà toxotes Schmidt iransiens Hallmann iranslara . Pulitzer-Finali iricurvatifera arter tuberculata Burton tuberosa Bowerbank tuberosa Hentschel tuberosa sp.nov. tuberosacapitaja Topsent tubulatum Bowerbank tubulosa Hallmann tumulosa Bowerbank Toxitenuis Topsent Fromont [preocc.] transiens Topsent Eurypon Clathria Hymeraphia Cornulum Ophlitaspongia Hymeraphia Ophlitaspongia Rhaphidophlus Clathria Hymeraphia Stylostichon Clathria Clathria Microciona Clathria Microciona Clathria Axociella Microciona Scopalina Clathria Arlemisina Ophlitaspongia Thalysias Stylastichan Microciona Lissodendaryx Hymeraphia Haliphysema Ophlitaspongia Microciona Jromontae nom.nov. C. (Micraciona) C. (Clathria) C. (Micraciona) C. (Clathria) C. (Axociella) C. (Microciona) see C. (Axociella) georgliaensis nom.nov. C. (Thalysias) C. (Thalysias) C. (Clathria) C. (Thalysias) see C. (Microciona) antarctica (Topsent) C. (Microciona) C. (Clathria) C. (Microciona) C. (Clathria) C. (Claihría) C. (Microciona) see C. (Axociella) C. (Clathria) C. (Clathria) C. (Clathria) Artemisina Echinoclathria ? C. (Thalysias) imperfectly known synonym of - (Microciona) _ antarctica (Topsent) C. (Wilsonella) A.(Antho) E.(Protao- —— phlitaspongia) Discorhabdella (Hymedesmiidae) ? Aulospongus (Raspailiidae) E. (Echinochalina) C. (Microciona) tubulosa Koltun tunixiae nom.nov. tylota Boury-Esnault typica Kirkpatrick icum Carter typica Var. orrecta entschel ulmus Vosmaer unica Cuartas urcealata Desor urizae nom.nov. vacelettia nom.nov. varians | Duchassaing & Michelotti vasiformis de Laubenfels vasiplicata Carter venosa Alcolado ventilabrum var. australiensis vespuri ium Lamarck viminalis Lendenfeld virgula Sarà & Siribelli virgultasa Lamarck vulcani Lévi vulpina Lamarck waldoschmitii de Laubenfels walpersii Duchassaing & Michelotti wesselensis sp.nov. Artemisina [for Microciona chelifera Lévi] Artemisina Clathria Echinonema Claihria Clathria Clathria Spongia [for Eurypon —— similis sensu Unz] |for Microciona curvichela Vacelet & Vasseur] Thalysias Thalyseurypon Echinonema Microciona Phakellia Alcyanium Thalassodendron Microciona Spongia Artemisina Spongia Echinoclathria Pandaras Artemisina C. (Microciona) Cornulum (Iophonidae) C, (Clathria) synonym of i (Thalysias) cactiformis (Lamarek) synonym of i (Thalysias) reinwardti Vosmaer C. (Clathria) C. (Claihria) synonym of C. (Clathria), prolifera (Ellis & olander) C. (Microctona) C. (Microciona) Anthosigmella (Spirastrellidae) C. (Clathria) synonym of Echinodictyum mesenterinum (Lamarck) (Raspailiidae) C. (Thalysius) synonym of C. (Thalysias) cactiformis (Lamarck) Spheciospongia (Spirastrell ae) synonym of Echinachohita subhispida Caner synonym of A. (n) involvens Schmidt) C. (Thalysias) Artemisina C. (Thalysias) Echinocluthria synonym of Patou spiculifera (Lamarck) (Axinellidae) C. (Thalysias) REVISION OF MICROCIONIDAE 587 incorporating the In- donesian and Micronesian faunas (in prep.). My phylogeny, based primarily on structural fea- tures of the skeleton, con- flicts with that of Hajdu et al., 1994, Their hypothesis is based on the premise that microgeometry is less like- ly to be influenced by modification during the course of evolution than structural features. I em- phasise skeletal structure and skeletal differentiation as primary characters in the evolution of Micro- } mas | cionidae. My classification x " allows inclusion of arcuate [ J à EX u^ LL *S =| and anchorate modified i ug chelae (which would other- , 2s) Wise be included in Myxil- ‘Ss lina). Identical structural p71 features, primarily, and megasclere geometry, secondarily, in species with arcuate or anchorate chelae may indicate evolutionary similarity, whereas it is B debatable whether or not fi observed modifications to 4] chelae are homologous or ( merely convergent. SE. Pacific im. — ESSAI es « SW. Africa 1 5-SE. Africa afic NW. central A New E Pacific Biogeography. Many early New Zilan m 7 attempts to analyse SW, central biogeographic patterns of Pacific marine sponges were un- successful because authors 2Üspecies —.. £ attempted too broad a taxonomic coverage without detailed taxonomic revisions and the belated recognition thal many so- called ‘widely distributed’ species actually consist of allopatric, cryptic sibling species, thus masking potentially informative pat- terns on distribution and FIG. 316. Distribution of microcionid species and levels of endemism in biasing proportions of biogeographic provinces. Division of provincial faunas based on traditional regional endemism. By biogeographic regions (Wiedenmayer, 1989). Bar length = total number of comparison, several con- provincial species; grey hatch = % of species endemic to each province. temporary bi ogeographic 30% endemic Tropical Australian 588 Air o Yu Be y DNE 3 i ' n gm | PERONIAN mee) = wo h lj | m |. FLINDERSIAN FIG. 317. A,B, Biogeographic relationships between Australian microcionid species within Indo-west Pacific marine provinces, showing numbers of shared species between provinces (provincial endemic species circled). analyses have had greater success based on rela- tively well-revised, restricted laxa (genera, families), (e.g., Van Soest et al., 1991; Hooper, 1991; Van Soest & Hooper, 1993; Hooper & Lévi, 1994; Bergquist & Kelly-Borges, 1995). Yet these studies too have not reached any consensus concerning general area statements for shallow water marine sponges, and they fail to distinguish between vicariant events or subsequent dispersals to explain species' distributions (Hooper & Lévi, 1994). Only one (Hooper & Lévi, 1994) included species of Microcionidae, providing a prelimi- nary analysis of biogeographic patterns amongst Indo-west Pacific species, comparing levels of MEMOIRS OF THE QUEENSLAND MUSEUM species endemism throughout world marine provinces, comparing regional faunas within the i Indo-west Pacific region (targeting the New Caledonian fauna in par- | ticular), and contrasting Q these broad distribution zi patterns for Microcionidae with those of the families l Raspailiidae and Axinel- n lidae. These analyses are bs eS . taken further in this study, -P focussing in particular on f the Australian fauna, and incorporating additional data derived from the present revision. Hooper & Lévi (1994) also provided an areacladistic analysis of one species group (Clathria ‘procera’ group), compar- ing sibling species' dis- tributions and postulating y historical biogeographic patterns and relationships throughout the world's 6 0 seas. Although it is clear from this study and others aa | that historical biogeog- ee di kd raphy can provide many 157 "--aW more facets to the questions posed by biogeographers, who attempt to understand relationships between both provincial species and the provinces themselves, it is also clear that analysis of many species groups are re- quired to gain a meaningful interpretation of data and to resolve a general area Statement for the marine biome (Van Soest et al., 1991). It is inappropriate to undertake such a detailed areacladistic analyses in this present work restricted to the Australian fauna, whereas it will be much more useful to include a revised Indonesian and Micronesian microcionid fauna into analyses (in prep.). The present analysis fol- lows the format used for Raspailiidae (Hooper, 1991). From present data and the earlier analysis (Hooper & Lévi, 1994) there is little evidence for cosmopolitan microcionids. A possible exception is Clathria atrasanguinea which has contiguous REVISION OF MICROCIONIDAE 0 | MAUGEA DAMPIERIAN Bog 589 populations (i.e. substan- tiated by comparison of voucher specimens) within the W and E Indian Ocean system, Mediterranean-E Atlantic system, and W At- lantic system (though not yet recorded from the Pacific). All other reported cases of cosmopolitan species consist of 2 or more sibling species. In contrast, 9 species of microcionids are widely distributed, usually with disjunct dis- tributions (possibly ex- plained by local extinctions within intermediate provin- ces; e.g., C. cactiformis). lt is possible that some of these species also comprise more than one allopatric sibling species but it was not possible to detect any consistent differences be- tween populations based solely on skeletal charac- ters. Similarly, most species of Microcionidae are restricted to single ocean systems, and only about 5% of species range widely within any particular sys- tem (whereas most other species are much more restricted in their ranges). Worldwide 7246 of species are restricted to a single biogeographic province al- though levels of regional endemism range from 30- 80% of species (Fig. 316). Microcionids are rela- tively diverse and prevalent, with most species found predominant- ly in shallow-waters but some ranging down to 2500m depth. They com- prise between 7-16% of all species of demosponges within the various marine FIG. 317. C-E, Biogeographic relationships between Australian microcionid biogeographic regions of species within Indo-west Pacific marine provinces, showing numbers of shared the world (8% average), species between provinces (provincial endemic species circled). with two major peaks of 590 diversity indicated: Indo-Australia region and NE Atlantic (Hooper & Lévi 1994: fig.3). Within the Indo-west Pacific, extending from ihe Andaman Sea to islands of the W Pacific rim, there are 196 species of which 115 (oc about 60%) are endemic to the region. This level of endemism ts closely comparable to that in NE Atlantic species. The Indo-west Pacific species represent about 7% of the region's demosponge fauna. Australian continental marine provinces con- tain 148 species of microcionids, comprising about 11% af the entire demosponge fauna for the region, with 111 (or 75%) being endemic. Although it is not possible to postulate detailed faunistic relationships using these crude analyses mücrocionid distribution data do support the con- cept of a differentiated soulhern Gondwanan fauna and northern Tethys fauna within coastal Australia (Hooper & Lévi, 1994) based on a different data set, Temperate Australian marine provinces contain a higher diversity of mücrocionids (90 species) and greater endemism (8196) than tropical provinces (the latter with 74 species, 59% endemic) (Fig. 316). These levels of endemism are very similar to those observed for Raspailiidae (Hooper, 1991), although the temperate raspailixd fauna was less diverse but had greater endemism than did the tropical fauna. This observation is further supported consider- ing the Australian fauna in more detail (Fig. 317). Peronian (Fig, 317A), OF all Australian continen- tal provinces the SE Australian temperate (Perontan) province contains the greatest diver- sity of species (60) and also the highest propor- tion of regional endemism (28 species or 48%). Not surprisingly greatest similarities in species composition (1.e., numbers of shared species) are with the adjacent temperate provinces (Flinder- sian, Maugean) and adjacent tropical Solanderian province, with only few ‘widespread’ species common to all Indo-west Pacific provinces. Flindersian (Fig. 317B). The southern and SW Australian temperate (Flindersian) province con- tains 46 species of which 22 (47%) are endemic, showing greatest similarities to the other two southern Australian temperate provinces (Peronian, Maugean), whereas few species are shared with the adjacent NW tropical (Dam- pierian) province (no doubt reflecting the relative MEMOIRS OF THE QUEENSLAND MUSEUM importance of the Leeuwin current lo the marme biogeography ofthe west coast (Pearce & Walker, 1991; Hooper, 1994)). Maugean (Fig. 31 7C). The coal temperate Bass Strait-Tasmanian (Maugean) province contains 26 species of microcionids with only 5 (or 19%} endemic, showing greatest similarities to the other southern temperate provinces. Selanderian (Fig. 317D). The NE Australian tropical (Solanderian) province contains 45 species (18 or 40% endemic), with similarities to the adjacent temperate Peronian, tropical Dim- pienan and SE Indonesian faunas. Dampierian (Fig. 317E). A similar relationship is indicated for the tropical northwest Australian (Dampierian) province containing 42 species (17 species or 4076 endemic). From these comparisons between adjacent provincial faunas it 1s clear that the two tropical provinces (Dampierian and Solandenan) share the highest number of sympatric species and also share a large number of species with both In- donesia and the western Pacific rim islands, Nevertheless, nearly half the number of species in each of these provinces are unique. Similarly, the three southern Australian (continental, lemperate) provinces share a large proportion of their species, especially Peronian and Maugean faunas, whereas there is very little mixing (less than 15% of species) between temperate anil tropical Australian mierocionid species. Only two species of microcionids have confirmed trans-Tusman Sea distributions (Bergquist & Fromont, 1988), and only four are found in both the Solanderian and New Caledonian provinces (Hooper & Lévi, 1993a), with similar observa- tions reported for Raspailiidae (Hooper, 1991) in which both New Zealand and New Caledonian faunas have exceptionally high numbers of en- demic microcionid species (nearly 70% and 67% endemism, respectively). The Antarctic and austral islands faunas have a low diversity of microcionids (18) and few en- demic species (2896), sharing many species with adjacent provinces such as New Zealand, SW Atlantic, Subantarctic islands and SE Pacific. There are no austral temperate species found in the Australian continental faunas, as also ob- served for the Raspailiidae (Hooper, 1991). REVISION OF MICROCIONIDAE LITERATURE CITED ACKERS. R.G., MOSS, D. & PICTON, B.E. 1992. Sponges of the British Isles. A colour guide and working documenL Sth Edn. 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Esponjas del Caribe Colombiano, Pp. 1-236. (Editorial Catalogo Cientifico: Bogota D.E. Colombia), ZIMMERMANN, H. 1907. Tierwelt am Strande der Blauen Adria. Zeitschrift fiir Naturwis- senschaftlich — Medizinische Grundlagen forschung 78: 293-322. REVISION OF MICROCIONIDAE 615 PLATE 1. A, Clathria (C.) angulifera (QMG303230, N. Stradbroke I., Qld., 30m, photo author). B, Clathria (C.) conectens (QMG305135, Direction Is, GBR, 15m,, photo author). C, Clarhria (C.) kylista (QMG300035, Mooloolabah, Qld., 10m, photo author). D, Clathria (C.) murphyi (QMG300656, Houtman Abrolhos, WA, 9m, photo NCJ). E, Clathria (C.) noarlungae (NTMZ3566, Kingston S.E., SA, 5m, photo NCH). F, Clathria (C.) striata (QMG303755, North Head, NSW, 50m, photo D. Roberts). 616 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 2. A, Clathria (C.) transiens (QMG300268, Furneaux Is, Tas, 18m, photo NCI). B, Clathria (W.) australiensis (QMG301458, Cook I, NSW, 15m, photo author). C, Clathria (W.) abrolhosensis (NTMZ3218, Houtman Abrolhos, WA, 22m, photo author). D, Clathria (W.) claviformis (QMG300576, Parry Shoals, NT, 30m, photo NCI). E, Clathria (W.) ensiae (NTMZ3561, Marion Reef, SA, 6m, photo NCI). F, Clathria (W.) tuberosa (QMG303428, Mandorah, NT, im, photo author). REVISION OF MICROCIONIDAE 617 PLATE 3. A, Clathria (D.) myxilloides (QMG300613, Kangaroo I, SA, 18m, photo NCI). B, Clathria (D.) pyramida (QMG300238, Kangaroo L, SA, 6m, photo NCI). C, Clathria (M.) aceratoobtusa with Rostanga arbutus feeding (QMG300543, Long Reef, NSW, Om, photo W. Rudman). D, Clathria (M.) illawarrae (QMG304572, Shell Harbour, NSW, 2m, photo L. Miller). E, Clathria (M.) lizardensis (QMG304121, Lizard I., GBR, 9m, photo author). F, Clathria (l) selachia (NTMZ2946, Shark Bay, WA, 7m, photo author). 618 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 4. A, Clathria (I) eccentrica (NTMZ288, Darwin Harbour, NT, 1m, photo author). B, Clathria (A.) thetidis (QMG303752, North Head, NSW, 50m, photo D. Roberts). C, Clathria (T.) abietina (NTMZ2163, Darwin Harbour, NT, 10m, photo author). D, Clathria (T.) abietina (NTMZ2611, Darwin Harbour, NT, 12m, photo author). E, Clathria (T.) cactiformis (NTMZ3831, St. Patrick's Head, Tas., 6m, photo NCI). F, Clathria (T.) cactiformis (QMG300736, Houtman Abrolhos, WA, 8m, photo NCI). REVISION OF MICROCIONIDAE 619 PLATE 5. A, Clathria (T.) cancellaria (QMG300536, Houtman Abrolhos, WA, 8m, photo NCI). B, Clathria (T.) cervicornis (QMG300707, Howick Is, GBR, 20m, photo author). C, Clathria (T.) coralliophila (QMG300377, Motupore I., PNG, 8m, photo NCI). D, Clathria (T.) aphylla (QMG300477, Houtman Abrolhos, WA, 20m, photo NCI). E, Clathria (T.) craspedia (QMG301452, Cook 1., NSW, 15m, photo author). F, Clathria (T.) craspedia (QMG301452, Cook I, NSW, 15m, photo author). 620 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 6. A, Clathria (T.) coppingeri (NTMZ1152, Northwest Shelf, WA, 55m depth, photo author). B, Clathria (T.) darwinensis (QMG303375, Darwin Harbour, NT, 17m, photo author). C, Clathria (T.) hallmanni (NTMZ2218, Darwin Harbour, NT, 0m, photo author). D, Clathria (T.) hesperia (NTMZ3327, Exmouth Gulf, WA, 16m, photo NCI). E, Clathria (T.) hesperia (QMG300213, Amphinome Shoals, NW Shelf, WA, 50m, photo author). F, Clathria (T.) hirsuta (QMG300603, Whitsunday Is, GBR, 20m, photo NCI). REVISION OF MICROCIONIDAE 621 PLATET7. A, Clathria (T.) hirsuta (QMG300771, Solitary Is, NSW, 7m, photo NCI). B, Clathria (T.) lendenfeldi (unregistered, Darwin Harbour, NT, 12m, photo N.Smit). C, Clathria (T.) lendenfeldi (NTMZ3489, Monte Bello Is, WA, 4m, photo NCI). D, Clathria (T.) lendenfeldi (NTMZ64, Port Essington, NT, 6m, photo author). E, Clathria (T.) lendenfeldi (QMG305 138, Direction Is, GBR, 20m, photo author). F, Clathria (T.) phorbasifor- mis (NTMZ2138, Darwin Harbour, NT, Om, photo author). G, Clathria (T.) phorbasiformis (NTMZ2203, Darwin Harbour, NT, Om, photo author). 622 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 8. A, Clathria (T.) reinwardti (NTMZ77, Port Essington, NT, 15m, photo author). B, Clathria (T.) reinwardti (NTMZ2206, Darwin Harbour, NT, Om, photo author). C, Clathria (T.) tingens (NTMZ2231, Darwin Harbour, NT, Om, photo author). D, Clathria (T.) tingens (QMG303826, Hook Reef, GBR, 31m, photo author). E, Clathria (T.) toxifera (NTMZ2222, Darwin Harbour, NT, 0m, photo author). F, Clathria (T.) vulpina (QMG301376, Byron Bay, NSW, 18m, photo author). REVISION OF MICROCIONIDAE 623 PLATE 9. A, Clathria (T.) wesselensis (QMG300361, Wessel Is, NT, 13m depth, photo author). B, Antho (A.) tuberosa (QMG300203, Shark Bay, WA, 8m, photo author). C, Antho (A.) tuberosa (QMG300678, Houtman Abrolhos, WA, 20m, photo NCI). D, Antho (P.) ridleyi (NTMZ2142, Darwin Harbour, NT, 0m, photo author). E, Antho (P.) ridleyi (NTMZ299, Darwin Harbour, 3m, photo author). F, Antho (1.) chartacea with Rostanga feeding (NTMZ2831, Botany Bay, NSW, 2m, photo W, Rudman). 624 MEMOIRS OF THE QUEENSLAND MUSEUM PLATE 10. A, Antho (L) punicea (QMG304399, Lizard L, GBR, 18m, photo author). B, Echinoclathria axinelloides (QMG300269, Furneaux Is, Tas, 20m, photo NCI). C, Echinoclathria bergquistae (QMG303827, Hook Reef, GBR, 24m, photo author). D, Echinoclathria digitata (QMG304763, Howick Is, GBR, 21m, photo J. Kennedy). E, Echinoclathria egena (QMG300669, St. Patrick's Head, Tas., 6m, photo NCI). F, Echinoclathria levii (QMG300675, Bicheno, Tas., 30m, photo NCI). REVISION OF MICROCIONIDAE 625 PLATE 11.A, Echinoclathrianotialis (QMG300614, Kangaroo I., SA, 6m, photo NCI). B, Echinoclathria riddlei (QMG300271, Bicheno, Tas., 30m, photo NCI). C, Holopsamma arborea (QMG300239, Kangaroo I., SA, 17m, photo NCI). D, Holopsamma crassa (QMG303235, Moreton Bay, Qld., 30m, photo author). E, Holop- samma laminaefavosa(QMG301399, Byron Bay, NSW, 20m, photo author). F, Holopsamma sp. indeterminate (QMG300620, Edithburgh, SA, 6m, photo NCI). 626 MEMOIRS OF THE QUEENSLAND MUSEUM ie ia ad PLATE 12. A, Echinochalina (E.) barba (NTMZ3853, Kent Is, Tas, 20m, photo NCI). B, Echinochalina (E.) intermedia (QMG300025, Mooloolabah, Qld., 30m, photo author). C, Echinochalina (E.) tubulosa (QMG300265, Kent Is, Tas, 20m, photo NCD. D, Echinochalina (P.) collata (QMG304120, Lizard I., GBR, 9m, photo author). E, Echinochalina (P.) isaaci (QMG305464, Swain Reefs, GBR, 22m, photo author). F, Echinochalina (P.) tuberosa (QMG300039, Mooloolabah, Qid., 15m, photo author). 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 CONTENTS HOOPER, J.N:A Revision of Microcionidae (Porifera: Poecilosclerida: Demospongiae), with description of Australian species ........ llle nnn