(Part 2 of 3) Revision of Microcionidae (Porifera: Demospongiae), with description of Australian Species.

John N . A . Hooper

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

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A Guide to Authors is displayed at the Queensland Museum web site A Queensland Government Project Typeset at the Queensland Museum REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 301 generally have an aspicular secondary fibre system, but otherwise spiculation and architecture is similar between these morphs and they are obvious synonyms. The species is included in the juniperina complex (see discussion of C. (T) hirsuta) all of which have reduced skeletal and fibre characteristics. This group includes 12 species (with about 20 synonyms): C. (T) arborescens, C. (T) cactiformis, C. (T) cervicornis, C. (T) clathrata, C. (T) corneolia, C. (T) craspedia sp. nov., C. (C.) decumbens, C. (T.) fusterna sp. nov., C. (T) 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 characteristics, nor on the basis of preserved material. Clathria (Thalysias) cancellaria (Lamarck, 1814) (Figs 149-150, Plate 5A) Spongia cancel/aria Lamarck, 1814: 382, 361. Rhaphidophlus cancel/anus; Topsent, 1930: 43, p1.2, fig.6. Clathria cancel/aria; Hooper & Wiedenmayer, 1994: 270. MATERIAL. HOLOTYPE: MNHNDT528: locality unknown, Peron & Lesueur collection. OTHER MATERIAL: WA - NTMZ1249 (fragment QMG300430), QMG300594 (NCIQ66C-4272-K), QMG300536 (NCIQ66C-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. 149H). DESCRIPTION. Shape. Lamellate, massive, mostly planar growth form, with or without basal attachment, consisting of more-or-less fused erect thin digits forming a continuous 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 5RP 8/2) to orange-red alive (5YR 6/10), light grey-brown in ethanol. Oscules. Small, 2-3mm diameter, mainly between ridges; pores minute, 0.5-1mm diameter, scattered over entire surface. Texture and surface characteristics. Firm, compressible, difficult to tear. Lamellae prominently striated with longitudinal ridges, grooves and discontinuous conules; margins of lamellae digitate, microconulose. Ectosome and subectosome. Relatively thin, dense ectosomal crust, 60-95 p.m thick, composed of erect ectosomal auxiliary subtylostyles forming continuous erect dermal palisade; immediately below ectosome, protruding through palisade, are mostly paratangential subectosomal auxiliary subtylostyles; in thicker, more elongated sections larger auxiliary subtylostyles form plumose ascending tracts supporting ectosomal skeleton, but in most sections they lie paratangential to surface, arising from ends of peripheral choanosomal fibres; subectosomal region relatively 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 subtylostyles. Choanosome. Skeletal architecture irregularly reticulate, without axial or extra-axial differentiation; spongin fibres relatively heavy, imperfectly separated into primary and secondary elements, forming ovoid or elongate, relatively cavernous meshes, 140-680p,m diameter; primary fibres, 92-165pm 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 fibres 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, 42-68p,m diameter. Megascleres. Choanosomal principal styles short, thick, slightly curved towards basal end, with rounded or very slightly subtylote smooth bases, fusiform points. Length 1664185.5)218p,m, width 7.5412.6)-16pm (holotype 161(198.1)-224 x 6-(12.1)-16pm). Subectosomal auxiliary subtylostyles long, thick, straight or very slightly curved, with slightly subtylote or rounded bases, usually microspined, fusiform points. Length 134.6(218.8)-286pm, width 3 .546 .5)-8 .6p,m (holotype 148-(229.7)-265 x 3-(5.9)-8p,m). Ectosomal auxiliary subtylostyles short, thin, straight, with prominent subtylote bases, invariably microspined, fusiform points. Length 86-(97.6)-114.5p,m, width 2.543.3)-4.5pm (holotype 85-(93.4)-102 x 2-(3.7)-5p.m). MEMOIRS OF THE QUEENSLAND MUSEUM 302zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 303 FIG. 150. Clathria (Thalysias) cancel/aria (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. 1, Accolada toxas. 304zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Acanthostyles long, thick, with slightly subtylote rounded bases, large spines dispersed over apical and basal extremities, usually with aspinose neck, occasionally evenly spinose; spines conical, barely recurved. Length 67(79.8)-92.5tim, width 3.8 -(8 .6)-11.6p,m (holotype 62-(80.2)-88 x 3-(8.1)-10p,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 I: 9-(10.9)16p„m (holotype 9-(12.3)-16p.m), length II: 3(4.4)-81,im (holotype 3-(4.8)-6p,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)-166p,m, width 0.4-(0.8)-1.2p„m (holotype 17-(102.3)-186 x 0.5-(1.1)-1.5p,m). REMARKS. Clathria (Thalysias) cancellaria is most closely related to the New Caledonian C. (T.) flabellifera, both with similar skeletal architecture, spicule geometry and spicule size, and also (vaguely) growth form, surface features and live colouration (Hooper &Levi, 1993a). The two populations differ in several cryptic characters, which Hooper & Levi (1993a) suggested were indicative of sibling species relationships rather than population variability. Clathria (T.) cancellaria has only a single category of toxa microsclere, resembling neither of the two found in C. (T.)flabellifera; it 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 are substantially thicker in the WA population. Clathria (Thalysias) cancel/aria also superficially resembles C. (T.) raphana and irregular growth morphs of C. (T.) coppingeri, with lamellate, planar growth form, although fibre characteristics, 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 paucispicular dermal skeleton of C. (T.) coppingeri, although both species conform to the Thalysias condition in having differentiated ectosomal and subectosomal megascleres. Clathria (Thalysias) cervicornis (Thiele, 1903) (Figs 151-152, Table 32, Plate 5B) Rhaphidophlus cervicomis Thiele,1903a: 959, 968, p1.28, fig.24a-e; Hallmann, 1912: 177; Brondsted, 1934: 22 23, fig.23; Simpson, 1968a: 70, p1.15. - Not Rhaphidophlus cervicomis; Vacelet & Vasseur, 1971: 96 97, text fig.50, p1.4, fig.3. - - Thalysias cervicomis; de Laubenfels, 1954: 135-137, text-fig.86. Clathriacervicornis; Bergquist, 1965: 165-167, fig.14. MATERIAL. HOLOTYPE: SMF679 (fragments ZMB3141, MNHNDCL2312) Ternate, Moluccas, Indonesia, 0°48'N, 127°23'E, 1894, coll. W. Kfikenthal (dredge). OTHER MATERIAL: QLD - QMG300707 (fragment NTMZ4045). CAROLINE IS., CENTRAL W PACIFIC - USNM22892, U5NM22905, QMG304828 (NCI OCDN-0488-0). HABITAT DISTRIBUTION. Usually at base of coral reef slope, on coral rubble, rock or seagrass beds, forming tangled thickets; intertidal - 45m depth; Howick Is (FNQ), (Fig. 151G). Aru Is, Arafura Sea, Moluccas Sea, Marshall Is, Palau Is, Truk Atoll (Chuuk), Ponape. DESCRIPTION. Shape. Stoloniferous, thin, cylindrical branches, 10-25mm diameter forming single digits, erect or creeping over substrate, to dense tangled thickets with numerous, thin, stoloniferous, anastomosing, cylindrical branches, and either small, central, single basal stalk, or multiple points of attachment to substrate via branches. Colour. Pale orange or orange-red exterior (Munsell 7.5YR 7/10-2.5YR 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, unornamented when preserved. Ectosome and subectosome. Ectosomal skeleton a dense palisade of discrete, erect spicule brushes composed of smaller ectosomal auxiliary subtylostyles, supported by underlying, ascending, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 305zyxwvutsrqp A FIG. 151. Clathria (Thalysias) cervicomis (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. MEMOIRS OF THE QUEENSLAND MUSEUM 306zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 152. Clathria (Thalysias) cervicomis (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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 307 TABLE 32. Comparison between spicule dimensions of Clathria (Thalysias) cervicornis, from present material and published records. All measurements given in p.m, as range, or range (and mean), of lengths x widths. Holotype (SMF679) (Indonesia) (N=1) (GBR) (N=3) (Micronesia) (N=1 ) (Brondsted,I 934) (Am I.,Indonesia) (N=1) (de Laubenfels,1954) (Marshall Is.) (N=I ) (Bergquist, 1965; Simpson, 1968a) (Palau Is.) Absent Absent Absent Absent Absent Absent 180-(258.9)-304 x 4-(5.8)-9 112-(128.3)-155 x 2-(3.3)-5 48-(62.0)-74 232-(274.9)-325 x 4-(4.7)-7 205-(236.7)-262 x 3-(4.3)-6 105-(118.3)-135 x 3-(4.1)-5 165-315 x 6-10 265 x 4 135-325 x 2.4-7.6 160-180 194 x 7 95-132 x 3-4.5 x 3-(5.4)-7 x 4-(6.3)-9 x 3-(4.2)-5 60-82 x 2.3-5.9 63-66 x 6 50-73 x 4-6.5 Chelae I 10-(12.4)-15 10-(11.6)-14 absent 12 10 8-13.6 Chelae II 3-(6.3)-9 3-(4.4)-6 absent Not recorded Not recorded Not recorded 1 2-(35.3)-55 x 0.5-(0.9)-1.5 x I .0-(1.2)-2.0 absent 40-90 40-50 34-56 SPICULE Principal megascleres Subectosomal subtylostyles Ectosomal subtylostyles Acanthostyles Tox as 86-(117.4)-151 x 2-(3.2)-4 52-(55.6)-61 I6-(54.5)-112 47456.2)-63 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% of branch diameter. Choanosome. Skeleton regularly reticulate, with even, rectangular or ovoid, relatively cavernous meshes, 180-270p,m diameter; spongin fibres very heavy, 90-125p.m diameter, thicker at nodes, up to 200p.m diameter, cored by multispicular tracts of subectosomal auxiliary subtylostyles which occupy only 60% of fibre diameter; echinating acanthostyles moderate to lightly dispersed over fibres, predominantly clustered around fibre nodes, sometimes rare or absent completely in some specimens; mesohyl matrix moderately light, with numerous subectosomal auxiliary megascleres scattered between fibres; choanocyte chambers small, oval, 35-55p.m diameter. Megascleres (Table 32). Choanosomal principal megascleres absent or completely undifferentiated from subectosomal spicules. Subectosomal auxiliary subtylostyles long, slender or robust, straight or slightly curved near point, with prominently subtylote, polytylote, 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 and 'neck' below basal swelling; spines moderately large, recurved. Microscleres (Table 32). Palmate isochelae occur in two size classes, with 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 or thinly branching thickets. In the western Pacific and Indonesian archipelago this species is a prominent member of the fringing coral reef community, commonly found at the base of reef slopes in the rubble and sediment, whereas on the Great Barrier Reef it has been found only in the far northern sector where it is 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, although there is some variability in spicule geometries. Specimens from Chuuk lack 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 is 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 exclusively in the ectosomal skeleton or scattered between fibres. Similarly, acanthostyles in MEMOIRS OF THE QUEENSLAND MUSEUM 308zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 (1903a) and subsequent authors. Consequently, 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 cervicomis belong to C. (T) abietina. Clathria (Thalysias) cervicornis is closely related to C. (T) corneolia Hooper & Levi (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 diversity 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 differentiated principal and auxiliary megascleres), and spicule dimensions. These are discussed further below for the respective species (and see also Hooper & Levi, 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, p1.40, figs f-f', p1.42, figs i-i'; Fristedt, 1887: 459; Hallmann, 1912: 215; Hentschel, 1912: 298, 358, 361-362; Topsent, 1932: 99, p1.5, fig.1; Hooper, 1984a: 55; Hooper & Wiedenmayer, 1994: 270. Thalysias coppingeri; de Laubenfels, 1936a: 105. Clathria lendenfeldi; Brondsted, 1934: 19-20, textfig.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). PARALECTOTYPE: BMNH1881.10. 21.330 (spirit): same locality. SYNTYPES of var. thuyaeformis: MNHNDT571, 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, NTMZ1152, 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 Am 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 planardendritic reticulate, with ascending radial primary branches and interconnecting secondary branches; branches form tight meshes (4-9mm diameter); primary branch diameter 12-1 6mm, 6.5-9.5mm midway, 1.5-4mm near apex of fan; connecting branches 1.5-3mm diameter. Colour Consistent, unpigmented, light beigebrown 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; microscopically hispid, irregular, consisting of tight reticulation 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 principal styles usually protrude through ectosomal skeleton with bases embedded in peripheral spongin fibres, especially at fibre junctions and on fibre endings; detritus also often present on sur- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 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 310zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 154. Clathria (Thalysias) coppingeri Ridley (QMG300106). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of principal and auxiliary subtylostyles. G, Palmate isochelae. H, Accolada toxas. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 311 TABLE 33. Comparison between present and published records of Clathria (Thalysias) coppingeri. All measurements given in p.m, as range, or range (and mean), of lengths x widths (N=25).zyxwvutsrqponmlkjihgfedcbaZY SPI CULE Holotype (Ridley, 1884a) Choanosomal principal styles 200-340x 15 Subectosomal auxiliary styles 150-250 x 5-6.3 Specimens (N= 2) Hol S.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Specimens (N= 16) zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFED (I ndonesia) zyxwvutsrqponml kjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Specimen (N= 1) (I ndian Ocean, . ^. .otyPe j um penna var. beta (MNHNDT571) 179-(254.1)-332 x 9-(16.2)-21 134-(196.5)-279 x 1.6-(4.6)-8 Ectosomal auxiliary styles 75-(108.8)-I 33 x Echinating acanthostyles 66-(99.5)-118 x Chelae Tox as (Hentsche1,1912; Brondsted,I 934) 164-340x 10-20 144-256 x4-6 (Pacific Ocean) Arafura Sea) 150-(228.4)-318 x 3.5-(13.6)-26 136-(202.2)-323 x 1-(4.9)-12 17 i .146.3) - 10 10-(13.8)-17 1-(3.7)-7 50-110 10-17 61497.3)-150 x 1.2 - (6.5) - 14 6-(13.1)-25 72-149 x 2-10 7-19 zyxwvutsrqponmlkjihgfedcba 39-126 58-(111.6)-191 x 28-(140.6)-245 x 0.5-(1.4)-2.5 0.5-(1.7)-4 face, sometimes replacing dermal skeleton entirely; subectosomal peripheral skeleton not sharply delineated from deeper choanosomal regions with choanosomal fibres usually immediately subdermal; meshes formed by fibre anastomoses often wider in peripheral skeleton. Choanosome. Skeletal architecture irregularly reticulate, without differentiation between axial or extra-axial regions; spongin fibres heavy, 40110Rm diameter, without any clear division between primary or secondary elements; fibres form oval or rectangular meshes, 100-220m diameter in axis, 280-405Rm 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 scattered 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, occasionally smooth, fusiform points. 164-362 x 1-9 69-(110.8)-I 42 x 1.1-(3.1)-5 110 x 6.3 201-520 x 4-24 Ectosomal auxiliary subtylostyles smaller, thinner than subectosomals, invariably with microspined bases. Acanthostyles variable in size, characteristically 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 contort, 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; typical morph (64% of specimens, including type material) with thicker ascending almost dendritic (primary) branches radiating outwards to produce an arborescent appearance; second morph (36% of specimens, including type material of S. juniperina) have even branching, even branch sizes and mesh sizes closely resemble Echinodictyum cancellatum (Raspailiidae). Foreign detritus in skeleton: abundant in choanosomal mesohyl of deeper water specimens (70 m depth), 312zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM rare in samples from shallower habitats (38-46 m depth). Ectosomal 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) (7%), 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 renieroidreticulate 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 (81% of specimens), with distinct primary (ascending) and secondary (transverse) fibre elements (12%), or noticeably heavier fibres in the axis than in the peripheral skeleton (7%). Echinating spicules: moderately heavy acanthostyles and choanosomal 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 at random throughout skeleton (20%). Mesohyl matrix: light with collagen found mainly around fibre nodes (93% of specimens), or heavy, granular (7%). Megasclere geometry: Principal spicules predominantly basally spined, but with variable proportion of entirely smooth spicules in specimens, ranging from 0-10% of spicules sampled (67% of specimens), 16-30% (27%), up to 56% of spicules (6% of specimens). Larger auxiliary subtylostyles 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), common (33%), rare (21%), or absent entirely (20%). Toxas: abundant (26% of specimens), common (20%), 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 southern 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 latitudes) for the west coast material (P>0.05), for all spicule types, although the inclusion of Queensland material (27°S) in analyses did reveal significant differences (commonly 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 or widths between specimens collected from 3846m or 70-90m depth ranges (P>0.05) for all spicule categories except acanthostyles. REMARKS. This species has a characteristic planar, flabellate growth form similar to C. (C.) loveni Fristedt (1887: 459) (from North America) and C. (C.) ultnus 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 possession of two size categories of auxiliary spicules places the species in the Clathria (Thalysias) group although it is atypical of most other species in having only rudimentary ectosomal structure, with sparse spicule brushes. This species belongs to 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) arrangement of choanosomal megascleres protruding through fibres, and dense echination of peripheral fibres). Included in this group are: C. (T.) lenden- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 313 TABLE 34. Summary of results of one-way ANOV's (Model I), testing for tate points (whereas in variability in spicule lengths and widths between locality and bathymetric distribu- typical material they are tion of Clathria (Thalysias) coppingeri. fusiform); ectosomal spicules occur in light paratangenti al bundles near the surface (versus Choanosomal (350) 12.56 P<0.0005 3.21 P>0.05 styles L not forming brushes); fibres are lightly cored W (350) 2.89 P<0.05 2.33 P>0.05 by paucispicular tracts Subectosomal (350) 8.18 P<0.0005 2.75 P>0.05 of choanosomal styles, styles L which produce an ir(350) 0.29 P>0.05 0.03 P>0.05 W regular renieroid Ectosomal (350) 8.87 P<0.0005 2.76 P>0.05 styles L reticulation (whereas most specimens have (350) 0.52 P>0.05 0.06 P>0.05 W aspicular fibres); acanAcanthostyles 9.95 P<0.0005 6.39 P<0.025 (350) L thostyles are only sparsely spined, and P>0.05 W (350) 0.37 P>0.05 0.82 together with (275) 0.69 P>0.05 0.02 P>0.05 Chelae L choanosomal styles, al(325) 11.48 P<0.0005 0.03 P>0.05 Toxas L ways form ascending P>0.05 0.02 P>0.05 W (325) 0.78 extra-fibre tracts within Number of groups: the skeleton. This 1.4 locality groups (27, 19, 16, 12°S latitude) specimen was initially 2. 2 depth groups (38-46m, 70-90m depth) assigned to C. (T.) coppingeri with some feldi, C. (T) major, C. (C.) caelata, C. (C.) in- hesitation, but there is no doubt that it belongs to anchorata, C. (T) clathrata, C. (T) costifera (in- this species and probably represents the southerneluding the nominate species Clathria spicata, C. most extent of its geographical range. bispinosa, C. whiteleggei and Ophlitaspongia ^There is a nomenclatural complexity that remembranacea, which have since been shown to quires brief comment. Wiedenmayer (1989) sugbe conspecific with other species of the spicata gested that the senior 'variety' name of S. group; Hooper et al., 1990)). This assemblage of juniperina (viz. thuyaefonnis) had priority over species does not appear to constitute a natural taxon because it cuts across a classification based Ridley's (1884a) subsequent species name, but on other (possibly more important) characters this is not accepted here. Lamarck (1814 (viz. Clathria and Thalysias). Nevertheless, all described several distinct 'varieties' of S. species are very close in fibre construction and juniperina, for which he used Greek symbols but spicule geometry, and in fact C. (T) coppingeri not a trinomen. It was Topsent (1932) who suband C. (T) lendenfeldi can only be easily differen- sequently elevated Lamarck's 'variety B' to a tiated on the basis of their respective growth subspecific rank by using it in a trinomen (ICZN forms, which is very characteristic for the former, Article 45fii), using the name thuyaefonnis as a but relatively variable in the latter species (see noun whereas it was previously used as an adjecHooper et al., 1990). tive by Lamarck. This subsequent designation is The single known specimen from southern therefore considered to be infrasubspecific, not Queensland differs in many respects from other subspecific (ICZN, Article 450, and does not take populations, accounting for most of the precedence over Ridley's (1884a) species desigvariability documented above. In shape it is close nation. Further support of this opinion is that S. to Lamarck's (1814) variety thuyaefonnis (Topjuniperina thuyaeformis is a composite taxon as sent, 1932: p1.5, fig. 1), whereas most samples examined resemble type material (Fig. 15314). some of Lamarck's syntypes belong to different This specimen also has a well developed ec- species. Thus the choice of the name coppingeri tosomal skeleton (consisting mainly of larger over thuyaefonnis is also supported by ICZN subectosomal megascleres which surround the Article 57g, and the pragmatic argument that bases of protruding choanosomal styles); true Ridley's (1884a) name has now become well (smaller) ectosomal auxiliary spicules have has- known for this species. LOCALITY' SPICULE (N) F DEPTH2 Prob. F Prob. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 314zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Thalysias) coralliophila (Thiele, 1903) (Figs 155-156, Table 35, Plate 5C) Rhaphidophlus coralliophilus Thiele, 1903a: 959,968, p1.28, fig.25a-d. Rhaphidophlus coralliophila; Hallmann, 1912: 177. Tenacia coralliophda; Burton, 1934a: 560. & Wiedenmayer, 1994: Clathriacoralliophila; Hooper, 270. cf. Microciona prolifera; Vosmaer, 1935a: 611, 645. MATERIAL. LECTOTYPE: SMF1784 (fragment ZMB3145): Ternate, Moluccas, Indonesia, 0°48'N, 127°23'E, 1894, coll. W. Kiikenthal (dredge). PARALECTOTYPE: SMF7 87 (fragment MNHNDCL2309): same locality. OTHER MATERIAL: QLD - BMNH1930.8.13.107. PNG QMG300377 (NCIQ66C-4518-A). HABITAT DISTRIBUTION. Growing on on Seriotopora coral, bivalves and dead coral substrates; shallow intertidal to 8m depth; Low Is (NEQ) (Fig. 155H); also Moluccas, Indonesia; Motupore I., S. PNG. DESCRIPTION. Shape. Thin, thick, or massively encrusting, covering up to 80mm` surface. Colour Red, dark orange or pale pinkish alive (Munsell 5R 8/2 - 1OR 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, compressible; 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 supported by well developed series of discrete subectosomal plumose brushes, the latter not protruding 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 (hymedesmoid), 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 (inicrocionid) fibre nodes cored by erect multispicular tracts of choanosomal principal spicules, congregated especially on ends of fibre nodes, forming plumose brushes and producing a series of ascending plumose or occasionally anastomosing tracts; subectosomal tracts mostly per- pendicular, less commonly longitudinal in thinner sections, always plumose; echinating acanthostyles 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, 120-250Rm diameter. Megascleres (Table 35). Choanosomal principal styles long or short, slightly curved, with rounded or only slightly subtylote, smooth bases, tapering to sharp fusiform points; barely differentiated from subectosomal auxiliary spicules, being only marginally thicker and more curved and with predominantly smooth bases. Subectosomal auxiliary subtylostyles relatively long, straight, thin, sharply pointed, with mostly microspined subtylote bases. Ectosomal auxiliary subtylostyles short, relatively thick, slightly curved, usually with subtylote microspined bases. Acanthostyles relatively long, thin, subtylote with aspinose necks, spines slender, long, slightly recurved. Microscleres (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 completely fused to shaft, and front ala widely separated from lateral alae. Toxas very abundant, short, thin, mostly wingshaped 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, up to 425 p.m 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 (1903a) and Burton's (1934a) 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 315 AB 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 316zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 156. Clathria(Thalysias)coralliophila (Thiele) (BMNH1930.8.13.107). A, Choanosomal skeleton. B, Fibre characteristics (x219). C, Echinating acanthostylc. D, Acanthostyle spines. E-F, Bases of subectosomal and ectosomal auxiliary subtylostyles. G-1-1, Palmate isochelae. I, Wing-shaped and u-shaped toxas. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 317 Were it not for the TABLE 35. Comparison between present and published records of Clathria presence of a (Thalysias) coralliophila (Thiele). All measurements are given in vim, denoted as specialised ectosomal range (and mean) of spicule length x spicule width (N=25). skeleton this specimen Le,ctotype Paralectotype GBR (BMNH PNG would be included in SPICULE (5MFI784) (SMF787) 1930.8.13.107) (NC1Q66C4518A) the Microciona group, 145-(252.8)406 x 164-(279.8)-349 x 285-(347.5)-408 x 224-(345.2)-452 x Choanosomal illustrating the dif- principal styles 7-(10.5)-13 10-(11.5)-14 12-(13.8)-15 10-(12.4)-15 ficulty in recognising 321-(355.4)-390 x 233-(302.7)-350x 312-(390.4)-488 x 223-(347.3)-430 x Subectosomal generic concepts sole- auxiliary styles 6-(7.2)-9 547.3)-11 547.9)-10 5-(7.2)-9 ly on the basis of skeleI04-(150.1)-208x IO2-(115.8)-134 x 1184140.9)-174 x I09-(121.5)-138 x Ectosomal tal architecture: auxiliary styles 2-(3.1)-4 2-(3.2)-4 2-(3.1)-4.5 2-(3.5)-5 leptoclathriid, hym48-(56.3)-7I x 52-(66.8)-78 x 84-(95.0)-105 x 76-(85.8)-95 x Echinating edesmoid or micro- acanthostyles 4-(5.2)-6 3-(5.2)-8 4-(6.3)-8 5-(6.6)-8 cionid (e.g., the Chelae I 2-(4.6)-8 2-(4.4)-8 3-(5.3)-8 3-(5.1)-8 concepts of Lep11-(13.1)-16 11-(12.8)-15 11-(13.5)-17 10-(12.1)-15 toclathria, Microciona, Chelae II 18-(76.6)-118 x 6-(35.2)-84 x 68-(140.3)-244 x 32-(125.5)-211 x Clathria). Toxas 0.5-(0.7)-1.0 0.5-(0.9)-1.2 0.5-(1.4)-2 0.5-(1.2)-2 Specimens from the Great Barrier Reef and Papua New Guinea also differ from the In- Colour. Dark red alive (Munsell 2.5R4/10); greydonesian population in having some larger brown in ethanol. spicules (notably toxas, principal styles and acan- Oscules. Small, up to 2.5mm diameter, scattered thostyles; Table 35), but generally spicule size over entire surface, not apparently confined to and spicule geometry are comparable between all any particular region. specimens, and there is no doubt that these Texture and surface characteristics. Harsh, firm, populations are conspecific. This species is related to the New Caledonian compressible, flexible; surface rugose with C. (T.) araiosa, differing in the distribution of prominent striations, raised ridges and subecspines on acanthostyles and auxiliary spicules, tosomal grooves running longitudinally and and specific dimensions of megascleres (which radially. are generally smaller in C. (T.) araiosa), and Ectosome and subectosome. Dense ectosomal Hooper & Levi (1993a) suggested that these skeleton consisting of erect plumose brushes of similarities in skeletal architecture and spicule smaller ectosomal auxiliary subtylostyles formgeometry may be indicative of a sibling species ing a continuous palisade on surface; subecrelationship. tosomal skeleton plumose with erect brushes of larger auxiliary subtylostyles arising and divergClathria (Thalysias) costifera Hallmann, 1912 ing from ends of choanosomal spicules in (Figs 157-158) peripheral fibre skeleton; subectosomal megascleres protrude into and partially interClathria costifera Hal!mann, 1912: 215-218, p1.31, mixed with smaller ectosomal spicules although fig.2, text-fig. 44; Vosmaer, 1935a: 648; Guiler, both spicule categories with distinctly localised 1950: 6; Hooper & Wiedenmayer, 1994: 270. distribution. Pseudanchinoe costifera; de Laubenfels, 1936a: 109. MATERIAL. HOLOTYPE: AME650: E. coast of Hinders I., Bass Strait, Tas, 40°01'S, 148°02'E, coll. FIV 'Endeavour' (dredge). OTHER MATERIAL: S. AUST - AME1035 (dry). VIC - QMG300666 (NCIQ66C-3633-P) (fragment NTMZ3798). HABITAT DISTRIBUTION. Rock reef; 15-60m depth; Bass Strait (Tas), Fumeaux Is (Vic), Kingston (SA) (Fig. 1570). DESCRIPTION. Shape. Large, thinly flabellate, plannar, 170-220mm high, 120-270mm wide, with small cylindrical basal stalk, 24-55mm long, 18-38mm diameter, corrugated apical margins. Choanosome. Choanosomal skeletal architecture irregularly reticulate, with heavy fibres and ascending primary and transverse secondary fibre components; primary fibres have paucispicular core of both choanosomal principal and subectosomal auxiliary megascleres, and choanosomal styles also protrude through fibres to form ascending, plumose brushes near periphery; secondary fibres entirely aspicular; echinating acanthostyles relatively sparse at core, slightly more numerous towards peripheral skeleton; mesohyl matrix moderately light, with few megascleres scattered between fibres. 318zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Megascleres. Choanosomal 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)-305p,m, 4 -(5.8)-7Rm width (holotype 129-315 x Subectosomal auxiliary subtylostyles slender, straight or very slightly curved near basal end, with smooth well developed subtylote bases, fusiform points. Length 296-(321.3)-342p,m, width 3-(3.8)-5p,m (holotype 198-336 x 3-7p,m). Ectosomal auxiliary subtylostyles slender, straight or slightly curved at centre, with prominently subtylote smooth bases, fusiform points. Length 117-(152.2)-175pm, width 2(2.6)-4p,m (holotype 112-158 x 2-3.5p,m). Echinating acanthostyles slender, usually slightly curved at centre, with subtylote bases, fusiform points, rudimentary spination, small spines, aspinose `neck'region proximal to base. Length 76-(85.4)-95p,m, width 4-(4.3)-5Rm (holotype 56-92 x 4-7 pm). Microscleres. Isochelae absent. Toxas accolada, abundant, long, thin or rhaphidiform, with slightly angular central curvature, straight arms, unreflexed points, sometimes completely straight. Length 176-(215.2)-264p,m, width 0.5-(0.8)-1.0p,m (holotype 110-315 x 0.51.2p,m). REMARKS. Vosmaer's (1935a) 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 a single size category of auxiliary spicule (whereas C. (T)costtfera obviously belongs to C. (Thalysias) in having two categories). Although apparently initially identified only by superficial comparison with the holotype (according 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 Islands 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 scattered 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). Clathria (Thalysias) craspedia sp. nov. (Figs 159-160, Plate 5E F) - MATERIAL. HOLOTYPE: QMG301436: Wommin Reef, S. of Cook Island, Tweed Heads, NSW, 28°12.1'S, 153°34.8'E, 22m depth, 04.ii.1993, coll. J.N.A. Hooper & 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.ii.1993, coll. J.N.A. Hooper & S.D. Cook (SCUBA). HABITAT DISTRIBUTION. In sand, coral rubble coral substrata at base of granite boulders; 15-22m depth; Tweed River region (NSW) (Fig. 159G). DESCRIPTION. Shape. Growth form erect, lamellate, digitate or bulbous-lobate, 120230mm long, 8-50mm diameter, partially burrowed 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 contiguous lamellae, occasionally isolated, 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 (10R 6/10), grey-brown in ethanol. Oscules. Large, 2-5nun diameter, slightly raised above surface, with membraneous lip, scattered over lateral margins of digits or on apex of digits. Texture and surface characteristics. Firm, compressible, flexible; surface fleshy, mostly smooth, relatively even in cylindrical specimens, or with crenellated margins in erect bulbous specimens. Ectosome 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 subtylostyles arising from terminal choanosomal spongin fibres; mesohyl matrix light in choanosome but more darkly pigmented in peripheral skeleton. Choanosome. Regularly reticulate, widemeshed, with heavy spongin fibres differentiated into primary and secondary elements, but no axial compression or differentiation between 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 160pm diameter; primary ascending fibres, 45-90p,m diameter, cored by 4-8 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 319 FIG. 157. Clathria (Thalysias) cost ((era 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. MEMOIRS OF THE QUEENSLAND MUSEUM 320zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 321zyxwvutsrqp ABC 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, 180-360iim connecting fibres, 40-6511,m diameter, with 1-3 diameter; echinating acanthostyles absent; spicules abreast; fibre reticulation forms cavern- mesohyl matrix heavy but only lightly pigmented 322zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 323 thoughout choanosome, with numerous auxiliary spicules scattered between fibres; choanocyte chambers oval, 30-45Rm diameter. Megascleres. Choanosomal principal styles long or short, slender, straight, with evenly rounded smooth bases, telescoped points. Length 103(221.4)-305p,m, width 3-(4.1)-6Km. Subectosomal auxiliary subtylostyles long, slender, straight, with smooth subtylote bases, fusiform points. Length 2554316.6)-3611.1.m, width 2.5-(3.8)-4.5p..m. Ectosomal auxiliary styles short, slender, straight, with smooth subtylote or evenly rounded bases, fusiform points. Length 1074135.4)17411m, width 1.5-(2.3)-3Rm. Echinating megascleres absent. Microscleres. Palmate isochelae in single size class but variable length, with front ala slightly longer than lateral alae, lateral alae fused completely to shaft, front ala entire, and apex of spicule characteristically constricted, pointed. Length 5-(10.7)-141Lm. Toxas moderately short, thick, wing-shaped, with rounded or slightly angular central curvature, non-reflexed arms. Length 16-(42.7)-761i.m, width 0.8-(1.9)-2.5p..m. ETYMOLOGY. Greek kraspedon, edge or border; occuring in the transition zone between the Solanderian and Peronian biogeographic provinces. REMARKS. This is a sibling species of C. (T) cervicornis, initially referred to that species based on skeletal structure and spicule types; it is separate by its different growth forms, surface 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 terminations such as telescoped points, smooth subtylote swellings, principal and auxiliary megascleres are differentiated within fibres, and isochelae have a terminal tooth-like constriction). These differences in skeletal characters might be intraspecific variability, and the 4 populations (i.e., including C. (T) fusterna sp. nov. and C. (T) corneolia) may represent a single, widely dispersed species. I consider major differences in growth forms, surface features and live colouration are consistently correlated to skeletal differences, supporting distinct taxa for the Tweed River and Gulf of Carpentaria populations (C. (T) fusterna below and Hooper & Levi, 1993a). Whereas C. (T) cervicornis is habitually long, thin, cylindrical, digitate, attached to the substrate 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'S, 130°47.0'E, 19m depth, 24.ix .1993, coll. J.N.A. Hooper, L.J. Hobbs & B. Alvarez (SCUBA). HABITAT DISTRIBUTION. Coral pinnacle near mouth of estuary, high sediment, turbid water; 19m depth; Darwin Harbour (NT) (Fig. 161H). DESCRIPTION. Shape. Arborescent, very thinly branching, reminiscent of an Axinella (Axinellidae), 290mm 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, convoluted, 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 slightly raised membraneous lip. Texture and surface characteristics. Soft, compressible, flexible branches, more harsh in ethanol; surface optically hispid, fleshy alive, even, bulbous, turgid, non-porous, but contracting 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, 400-500Rm apart, extending 300-450[Lm from surface, surrounded at base by paratangential tracts of both larger and smaller auxiliary subtylostyles, sometimes in plumose brushes surrounding base of principal spicule, more often in tangential or paratangential tracts; echinating acanthostyles also erect peripheral fibres, protruding through surface; subectosomal skeleton usually reduced with peripheral 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 (ectosomal) or larger (subectosomal) auxiliary spicules, both appearing to be intermingled in MEMOIRS OF THE QUEENSLAND MUSEUM 324^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 (110-160pm diameter) than at periphery (60-90pm 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 bifurcation and relatively few transverse connecting 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, cored by 1-3 principal spicules abreast, interconnecting primary fibres mainly in axial region of skeleton, producing oval or elongate fibre meshes, generally smaller at core (120-190Rm diameter) than periphery (170-240pm diameter); fibres moderately heavily echinated by acanthostyles, evenly distributed over fibres although possibly more abundant on exterior surface of fibres, especially in peripheral skeleton; mesohyl matrix moderately light, including some auxiliary spicules scattered between fibres; choanocyte chambers small, oval, 12-24pm diameter. Megascleres. Choanosomal principal styles long, robust, slightly curved near base, entirely smooth, evenly rounded bases without any tylote swelling, long, tapering, fusiform points. Length 188(301.8)-492Rm, width 4-(12.8)-21Rm. Subectosomal auxiliary subtylostyles long, slender, straight, slightly subtylote microspined bases, fusiform points. Length 210-(282.2)365 p.m, width 3-(4.3)-6Rm. Ectosomal auxiliary subtylostyles short, slender, straight, slightly subtylote microspined bases, fusiform points. Length 1154135.3)153Rm, width 1.5-(2.1)-2.5Rm. Echinating acanthostyles long, slender, straight or slightly curved at centre, slightly subtylote bases, evenly spined except for aspinose 'neck' proximal to base; spines large, recurved, sharp; points sharp or slightly rounded, spinose. Length 96-(104.8)-116Rm, width 3-(5.6)-11Rm. Microscleres. Palmate isochelae abundant, single size class, unmodified, with lateral and front alae approximately same length, long, lateral alae entirely fused to shaft, front ala detached along lateral margin. Length 15416.8)-181km. Toxas abundant, wing-shaped, thick, variable in length, with wide central curvature, slightly reflexed arms. Length I: 73-(111.2)-124p,m, width 2-(3.8)-8p,m; length II: 17-(26.2)-36p..m, width 0.5-(1.7)-2.0p..m. ETYMOLOGY. For the type locality. REMARKS. Clathria (T.) darwinensis is similar to C. (T.) lendenfeldi, C. (C.) inanchorata and C. (T.) coppingeri of the spicata group having choanosomal principal spicules protruding through peripheral spongin fibres forming a hispid surface. It differs from these species, and to some extent the concept of the spicata group, having smooth principal styles enclosed within spongin fibres, only protruding through fibres at 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-renieroid skeletal architecture, and toxa morphology, 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 vestigal spines), possession of 2 size classes of auxiliary styles (versus one size class), thinly branching gross morphology (versus bulbous branches), and spicule dimensions. Clathria (Thalysias) dubia (Kirkpatrick, 1900) (Figs 163-164) Microciona dubia Kirkpatrick, 1900a: 128, 136, 141, p1.12, fig.3,3a, p1.13, fig.2a-f. Cionanchora dubia; de Laubenfels, 1936a: 108. Clathria dubia; Hooper & Wiedenmayer, 1994: 270. cf.Microciona prolifera; Vosmaer, 1935a: 608, 643. MATERIAL. HOLOTYPE: BMNH1898.12. 20.37: Flying Fish Cove, Christmas I., Indian Ocean, 100 25.5S, 105°40'E, coll. Mr Andrews (dredge). HABITAT DISTRIBUTION. Coral rubble; probably intertidal; Christmas I. (Indian Ocean) (Fig. 163H). DESCRIPTION. Shape. Thickly encrusting lamella, 12mm diameter, on eroded bivalve shell. Colour. Yellow preserved. Oscules. Not seen. Texture and surface characteristics. Compressible; optically smooth surface. Ectosome and subectosome. Ectosome microscopically hispid, with bundles of ectosomal auxiliary megascleres protruding through surface, forming a relatively thick dermal palisade, arising from subdermal brushes of subectosomal REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 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 acanthostyle. 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 subtylostyles arising from ends of choanosomal megascleres. Choanosome. Choanosomal skeletal hymedesmoid, with a thin layer of spongin lying on substrate, 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 subtylostyles long or short, slightly curved, usually with prominently microspined bases, occasionally smooth, bases subtylote, points fusiform. Length 132-(195.6)-292p,m, width 7-(10.6)-16[Lm. Subectosomal auxiliary subtylostyles long, straight, with prominent subtylote, microspined MEMOIRS OF THE QUEENSLAND MUSEUM 326zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 subtylostyle. F, Palmate isochela. G, Wing-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 327 bases, fusiform points. Length 2184280.8)314Rm, width 4.5-(5.9)-7Rm. Ectosomal auxiliary subtylostyles straight, with well formed tylote, microspined bases, fusiform points. Length 86-(110.2)-15311m, width 3-(3.9)-5Rm. Acanthostyles short, thick, tapering clubshaped, with large recurved spines on basal portion of shaft, aspinose point; spines on basal swelling often bifurcate, greatly recurved. Length 35-(46.4)-54Rm, width 5-(8.2)-11Rm. Microscleres. Isochelae modified palmate, very small, often anisochelate, with lateral alae completely fused to shaft, front ala often bifurcate or trifurcate, producing multiple, partially fused teeth. Length 2-(4.8)-811m. Toxas divided into two morphs - I: very short, ? oxhorn, relatively thick, slightly curved at centre, with slightly reflexed points. Length 4(6.6)-9Rm, width 0.5-(1.1)-1.5Rm. II: Accolada, long, thin, with gently rounded or angular curvature, with straight points. Length 1124195.2)295Rm, width 0.5-(1.4)-2Rm. REMARKS. De Laubenfels (1936a) assigned this species to Cionanchora because it supposedly had anchorate (rather than palmate) isochelae, although differing in no other respect from typical species of Clathria (Tlialysias). 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, p1.2, fig2; Thiele, 1903a: 957; Hallmann, 1912: 177; Levi, 1961a: 136-137, text-fig.10, pl.]. 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 (fragment BMNH1930.7.1.7): same data. OTHER MATERIAL: NT - NTMZ3113 (fragment QMG300579), NTMZ3146 (fragment QMG300219). INDONESIA - SMF1788. HABITAT DISTRIBUTION. Coral reef and coral rubble; 16-20m depth; known Australian distribution: Parry Shoals, Timor Sea (Fig. 165H); also Moluccas and Sulawesi, Indonesia (Thiele, 1899, 1903a), Vietnam (Levi, 1961a). 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 surface. Choanosome. Very irregularly reticulate, cavernous, with very large primary fibres running longitudinally through branches, up to 140Rm diameter, interconnected by smaller tangled secondary fibres, up to 70Rm diameter, producing vaguely triangular skeletal meshes, up to 450Rm 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 rnicrospined 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. 328zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM in the diversity and geometry of its spicules, but it differs again in growth form (being bushy, flattened branching), skeletal structure (more irregularly reticulate) and toxa morphology (includes asymmetrical sinuous forms). It is possible Specimens (2) Holotype (Levi, 1961) SPICULE that all three species are extreme morphological (Timor Sea) (NME119) (Vietnam) variants of a single species, in which case reinChoanosomal 1744224.0_292 1424197.7)-259 190-300 principal wardti would have priority, but all these morx10-22 x 6-(10.9)-14 x 9-(13.7)-19 styles phological differences are consistent within each Subectosomal 1944241.2)-285 165-(226.9)-262 growth form type (morphospecies) and probably 255-310 x 6-7 auxiliary x 4-(6.6)-9 x 4-(6.3)-8 represent fixed genetic differences. styles Clathria (T.) erecta is also vaguely similar to C. Ectosomal 85-(102.7)-134 92-(138.5)-198 100-200 x 3-5 auxiliary (T.) vulpina in the overall structure of spongin x 3-(4.2)-6 x 344.2)-5 styles fibre skeleton and spicule skeleton, both species 61-(68.5)-75 64472.6)-78 Echinating having a characteristic, more-or-less triangular 80-95 x 9-10 acanthostyles x 5-(6.7)-8 x 6-(6.8)-8 skeletal network of fibre meshes fully cored by 4-(6.1)-8 7 4-(6.3)-9 Chelae I principal styles, although this fibre reticulation is 10-17 10-(12.2)-14 12-(13.4)-15 Chelae II much more regular in the latter species. This I 2-(197.0)-265 25-(138.5)-204 structural feature is prominent and their inferred 120-160 x zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Toxas x 0.5-(1.2)- I .5 x 0.8-(1.1)-1.5 similarities are immediately obvious upon casual observation, but the two species differ from each Toxas accolada, relatively long, thin, with other in most other respects. The presence of two size classes of isochelae, small, angular central curvature, or rounded centrally, straight arms and reflexed points; including contort forms, has not been recorded previously for C. (T.) erecta but are consistently juvenile forms resemble oxhorn toxas. Larvae. Incubated parenchymella larvae small, present in all specimens including the type spherical, 140-190iLm diameter, with light material. Thiele's (1899, 1903a) Indonesian mesohyl matrix and larval toxas dispersed within specimens are identical to the Timor Sea specimens in most respects (see Table 36), axis. whereas Levi's (1961a) material from Vietnam differs slightly in growth form (compare Levi's REMARKS. This species is a sibling of C. (T.) (1961a) Plate 1 with Fig. 16514 of the present reinwardti based on similarities in skeletal strucstudy), and spicule dimensions are relatively ture (even-meshed, cavernous primary and larger. secondary tracts), geometry of some spicules TABLE 36. Comparison between present and published records of Clathria (Thalysias) erecta (Thiele). Measurements in pim, denoted as range (and mean) of spicule length x spicule width (N=25). ? (smooth, curved principal styles; robust subectosomal auxiliary subtylostyles; slender accolada toxas with slightly angular central curvature; 2 sizes of isochelae), and live colouration (orangebrown). They differ significantly in growth form (C. erecta is arborescent, thickly branching; C. reinwardti has only thin or slightly thick cylindrical branches), surface features (prominent surface conules versus completely smooth or irregularly rugose surface), and acanthostyle geometry (tapering, sharply pointed and subtylote 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 (Wilson, 1925; de Laubenfels, 1954), is also very similar to both C. (T.)erecta and C. (T.)reinwardti 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 Carpentaria, Qld, 11°12.7'S, 142°05.9'E, 21m depth, 30.xi.1991 (dredge). QMG301008: NW. of Port Musgrave, Gulf of Carpentaria, Qld, 11°18.9'S, 140°55.8'E, 41m depth, 27.xi.1991 (dredge). OTHER MATERIAL: GULF OF CARPENTARIA, - QMG301013, QMG303462. RED SEA - PIB0004-17 (fragment QMG300064). HABITAT DISTRIBUTION Soft sediments, mud, shell grit; 21-58m depth; Shelburne Bay, Torres Strait 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 acanthostyles. 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 embedding in soft sediments); rhizomous roots encrusted with sand and shell fragments; apex of club usually large, 50-100mm diameter, 60110mm long, composed of fused, tightly anastomosing digits; adjacent digits fused with dense fleshy surface; apex of digits tapering slightly pointed. Colour Grey-brown on-deck (Munsell 2.5Y 8/2), grey in ethanol. Oscules. Few small pores, 0.5mm diameter, possibly oscules, scattered near apex of digits (seen in preserved material only). Texture and surface characteristics. Stalk tough, wiry, flexible, apex of club softer, more compressible but with firm axis; slightly convoluted surface 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 ectosomal skeleton; ectosomal and subectosomal spicule skeletons very dense but together comprise only 10-20% of branch diameter. 330zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 331 A E a D o zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFE 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, 90-2001im diameter, heavily collagenous, forming tight oval or rectangular meshes, 150-40011m diameter; fibres not obviously differentiated into primary or secondary elements, but meshes slightly more cavernous in peripheral skeleton than in axis; fibres virtually fully cored by multispicular tracts of both subectosomal auxiliary subtylostyles and choanosomal principal styles, together occupying 80-90% fibre diameter, interconnected by very large, bulbous fibre nodes, 160-400p.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 MEMOIRS OF THE QUEENSLAND MUSEUM 332zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 333 8 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. I, Holotype QMG303240. MEMOIRS OF THE QUEENSLAND MUSEUM 334zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 subtylostyle. F, Wing-shaped toxas. G, Palmate isochelae. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 335 martrix heavy, with few auxiliary spicules scattered between fibres but abundant microscleres lining small oval choanocyte chambers, 20451.tm 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)-355tan, width 5413.9)25p.m. Subectosomal auxiliary subtylostyles long, thick or thin, mostly straight, with subtylote microspined bases, only occasionally smooth bases, and fusiform points. Length 2114369.2)385t.Lm, width 4-(9.5)-121J.m. Ectosomal auxiliary subtylostyles short, straight or slightly curved near basal end, subtylote microspined bases, fusiform points. Length 99-(127.7)-163Rm, width 2-(3.2)-4Rm. Echinating acanthostyles thick, robust, relatively short, heavily spined but with bare neck and point; spines moderately large, conical erect (not recurved). Length 73-(82.8)-96vm, width 3-(7.4)-12vm. 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 5411.8)Toxas wing-shaped, relatively thick, with slightly angular central curve, arms at wide angles from centre, straight or slightly reflexed points. Length 18-(41.4)-63iLm, width 1.541.9)2.5 p.m. ETYMOLOGY. Latin fusterna , club or knotty part of a tree. REMARKS. This species is possibly a very atypical, highly specialised population of C. (T) cervicomis 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) cervicomis. However, there are subtle differences in skeletal characteristics that consistently differentiate the two populations: possession of differentiated principal and auxiliary spicules (whereas cervicomis has undifferentiated structural megascleres), acanthostyle spines are erect, conical (not recurved), a single size class of palmate isochelae (not two), and slightly subtylote bases on principal and auxiliary spicules (not prominently subtylote as in most C. cervicomis). 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 complex of four: the cylindrical C. (T) cervicomis 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 & Levi, 1993a). This species is discussed 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.i.1985, coll. J.N.A. Hooper. HABITAT DISTRIBUTION. Encrusting under beach rock and coral rubble; intertidal pools; NT (Fig. 169H). DESCRIPTION. Shape. Thinly encrusting, up to 1.5mm 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 1 mm diameter, scattered 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 smooth, even, without conules, ridges or canals, and encrustation conforms exactly with contours of substrate. Ectosome and subectosome. Opaque in life, 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 subtylostyles; moderate quantities of detritus in ectosomal skeleton; subectosomal region extensive, occupying 70% of sponge thickness, composed of mostly paratangential tracts of larger auxiliary subtylostyles gradually ascending and diverge at surface. Choanosome. Skeletal architecture hymedesmoid in choanosomal (basal) region, but distinctly plumose towards peripheral skeleton; spongin fibres consist of a basal layer of spongin lying against substrate, 18-35Rm thick, with bases of choanosomal principal subtylostyles and acan- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 336zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA thostyles embedded in spongin and standing perpendicular to substrate; choanosomal principal subtylostyles morphologically close to subectosomal 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 35-48Rm diameter; mesohyl matrix heavy, granular, with small amounts of detritus Megascleres. Choanosomal principal subtylostyles 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)-419.5Rm, width 849.3)11Rm. Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, almost indistinguishable from choanosomal megascleres but with prominent spined subtylote bases. Length 284.5(362.2)-450Rm, width 2-(3.2)-4.5Rm. Ectosomal auxiliary subtylostyles short, thin, fusiform, with microspined subtylote bases. Length 944121.2)-15 ltim, width 0.8-(1.4)2.5 Rm. Acanthostyles subtylote, fusiform, relatively evenly spined although spines less heavily concentrated in 'neck' region, proximal to base, heavier on apical and distal extremities; spines relatively small, weakly formed. Length 52(59.5)-72Rm, width 3-(4.5)-6.5Rm. Microscleres. Palmate isochelae relatively common, 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 5410.8)17Rm. Toxas accolada, moderately common, long, thin, almost straight, with only slight angular central curvature, straight arms, straight (unreflexed) points. Chord length 174-(208.0)481p.m, width 0.4-(0.8)-2.0Rm. Associations. Single known specimen growing next to encrusting sponges (Renieria, Haliclona, Mycale), polychaete worm tubes (Pomatoleios kraussii) and simple ascidians. ETYMOLOGY. For E.R. Hal!mann in recognition of his contributions to Australasian microcionids. REMARKS. It is difficult to define C. (T.) hallmanni in any single unique character apart from the close resemblance between choanosomal principal and subectosomal auxiliary subtylostyles. As far as can be ascertained 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, especially those from the Indo-Malay archipelago, rarely include details on live colouration or surface details. But none of these species match the present one in spicule geometry either. Consequently, C. (T.) halhnanni can be differentiated from other encrusting (hymedesmoid) Clathria (Thalysias) species in: grey-brown live colour; even (unornamented) surface, i.e., lacking subectosomal drainage canals commonly found in thinly encrusting species; plumose ectosomal and subectosomal skeletal structure as well as extensive paratangential tracts composed of both sorts of auxiliary spicules in the periphery; entirely smooth, relatively short and thin choanosomal subtylostyles, barely different from the subectosomal auxiliary subtylostyles except for pattern of spination; evenly spinous acanthostyles, unmodified palmate isoehelae, and thin, nearly straight toxas with unreflexed arms. None of these features are unique or particularly distinctive by themselves but their combination is unique 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, fragment QMG304991): 1.8km N. of Bessieres I, Anchor Is, Exmouth Gulf, WA, 21 0 30.6S, 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 substrates; 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, resembling species of Cymbastela (Axinellidae); margins 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 337zyxwvutsrqp AgBzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC FIG. 169. Clathria (Thalysias) hallmanni sp.nov. (holotype NTMZ2218). A, Choanosomal principal subtylostyle. 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. 338zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 339 voluted, concentric ridges 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 smooth, even, upper surface more conulose (with terminal oscules), longitudinal folds, ridges or convoluted folds and small digits. Ectosome and subectosome. Discrete surface brushes produce specialised ectosomal skeleton; brushes composed of ectosomal auxiliary subtylostyles on outer surface forming thick, erect bundles but not continuous palisade, with 1 or several choanosomal principal styles also protruding through surface associated with ectosomal brushes; subectosomal auxiliary subtylostyles intermingled with ectosomal spicules but originating slightly lower in peripheral skeleton; subectosomal region greatly reduced with peripheral choanosomal fibres lying immediately below ectosome; mesohyl matrix moderately heavy in peripheral region. Choanosome. Choanosomal skeleton almost regularly renieroid reticulate although renieroid pattern severely disrupted by heavy concentrations of echinating spicules (both principal styles and acanthostyles); spongin fibres very well developed, dark brown, imperfectly divided into primary (90-140iun diameter) and secondary (25-45 diameter), and very large fibre nodes (up to 220Rm 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; secondary 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 abundant, particularly at fibre nodes, also contributing to ectosomal spicule brushes, with only small 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 (45-1151im diameter) than at core (170-25011m diameter); choanocyte chambers oval, 35-551m diameter, often lined by isochelae; mesohyl matrix moderately heavy but only lightly pigmented. Megascleres. Choanosomal principal styles straight or slightly curved near centre, with rounded or very slightly subtylote bases, bases usually smooth, occasionally microspined, long tapering fusiform points. Length 1624187.3)21311m, width 8-(11.1)-14p.m. Subectosomal auxiliary subtylostyles straight or very slightly curved near basal end, slightly subtylote bases lightly microspined, fusiform points. Length 121-(138.1)-168Rm, width 4(4.6)-5.5p.m. Ectosomal auxiliary subtylostyles only slightly shorter than subectosomal spicules but consistently thinner, with smooth subtylote bases, fusiform points. Length 97-(121.1)-147Rm, width 2-(2.9)-4.5p.m. 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)-112p,m, width 4Microscleres. Palmate isochelae very abundant, poorly silicified, sigmoid, with short, sharp, vestigial unguiferous alae. Length 11-(13.7)-151.t.m. Toxas absent. ETYMOLOGY. Latin hesperius, western; from WA. REMARKS. This species is borderline between Clathria 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 subectosomal spicules can be consistently differentiated by their thickness as well as the absence or presence of microspi nes on their base, respectively, even though there is no marked difference in length between the two categories. Clathria (T) hesperia has a distinctive lamellate growth form with differentiated osculiferous and porous faces. Its skeleton is a mixture of plumose tracts (reminiscent of C. (M.) coccinea, particularly its plumose fibre nodes, or the MEMOIRS OF THE QUEENSLAND MUSEUM 340zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 subtylostyle. 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. Palmate isochelae are vestigial, unguiferous, sigmoid reminiscent of C. (T.) michaelseni (which is an encrusting, hymedesmoid species, has toxas, and differs from this species in virtually every other respect). Clathria (Thalysias) hirsuta Hooper & Levi, 1993 (Figs 173-175, Table 37, Plate 6F, 7A) Clathria (Thalysias) hirsuta Hooper & Levi, 1993a: 1259-1264, figs 19-20, table 10; Hooper & Wiedenmayer, 1994: 270. MATERIAL. HOLOTYPE: QMGL2746 (fragment NTMZ1551): Cairns region, Qld, 16°56'S, 146°00'E, 1982, coll. A. Kay (trawl). PARATYPES: QMGL2750 ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 341 FIG. 172. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Thalysias) hesperia sp.nov. (holotype QMG300213). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of subectosomal auxiliary subtylostyle. F, Reduced sigmoid palmate isochelae. 342zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM (fragment NTMZ1555), QMGL2754 (fragment NTMZ1560): Cairns region, Qld, 16°56'S, 146°00'E, coll. A. Kay (trawl). OTHER MATERIAL: QLD QMG300328 (NCIQ66C-1893-X; fragment NTMZ3513), NTMZ3494, QMG303040, QMG304767, QMG300081, QMG303971. NSW QMG300771 (NCIQ66C-1185-F). NEW CALEDONIA - QMG301274, QMG301325, QMG301340. for whole of length, longer than front ala; toxas accolada and wing-shaped morphs, very thin, sometimes slightly sinuous, rarely raphidiform, only slightly curved at centre, with straight nonreflexed arms or only slightly reflexed points (or exclusively raphidiform with small angular central curve and straight arms). HABITAT DISTRIBUTION. Rock, dead coral and coral rubble substrates, usually on broken substrates, sides of bornmies, or in gullies; 7-30m depth; Shelburne Bay, Howick Is (FNQ), Whitsunday Is (NEQ), Noosa Heads, Stradbroke I. (SEQ), Solitary Is (N. NSW) (Fig. 1730). Also New Caledonia lagoon (Hooper & Levi, 1993a). REMARKS. Clathria (T.) hirsuta was originally described and illustrated from both New Caledonian and Queensland populations (Hooper & Levi, 1993a), in which it was reported that the New Caledonian population possessed ectosomal auxiliary subtylostyles (i.e., belonging to Thalysias) whereas Queensland specimens did not (i.e., belonged to Clathria). Since this publication 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 differences between these two populations are discussed in Hooper & Levi (1993a). Two 'atypical' specimens from SE. Australia (QMG300328, G300771) (which are included in the diagnosis above, in brackets) further illustrate the considerable variability of this species (toxas are only raphidiform, auxiliary spicules lack tylote bases, and a structured ectosomal skeleton is present (Figs 174-175)). These specimens are reminiscent of the 'reduced' New Caledonian population. At first glance this species lacks any remarkable or unique feature that stands it apart from other Clathria (Thalysias), but it possesses an unusual combination of characters not seen in any other species. It is superficially similar to C. (T) vulpina in growth form, but spiculation and fibre characteristics are quite different between the two species. Its skeletal architecture and spiculation is also very similar to C. (T) schoena (i.e., USNM22404; which may be different again from Rhaphidophlus schoenus of authors; e.g., Simpson, 1968a; Alcolado, 1980; Van Soest, 1984b), but these species differ significantly in their growth form, spongin fibre architecture and ectosomal characteristics. Clathria (T) hirsuta has very lightly spined acanthostyles, comparable with those of C. (T) transiens, and it is also closely related to that species in its fibre characteristics and spiculation, although they DESCRIPTION. (See Hooper & Levi, 1993a). DIAGNOSIS (Table 37). Tubular, lobo-digitate, reticulate-honeycombed, excavated growth forms superficially resembling Phakellia cavernosa; bright red (or orange-red) conules, paler pink or white between conules, prominent subdermal drainage canals; large oscules scattered between surface projections; texture firm, compressible, slightly arenaceous; surface prominently conulose, conules pointed (or rounded, fleshy); ectosome with irregular, tangential or paratangential layer of intermixed ectosomal and subectosomal subtylostyles (or with light palisade of smaller auxiliary styles forming 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, running longitudinally through branches, ascending to surface, interconnected by 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 and fusiform points; subectosomal auxiliary subtylostyles 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 microspined bases (or with simply rounded, smooth bases); 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 ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 343 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, Subectosomal auxiliary style. J, Ectosomal auxiliary style. K, Echinating acanthostyles. L, Raphidiform toxa. M, Palmate isochelae. N, Section through reduced specimen. 0, Known Australian distribution. P. Holotype QMGL2746. Q, Atypical QMMG300328. 344zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 345 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. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 346zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clathria (Thalysias) juniperina TABLE 37. Comparison between Australian and New (Lamarck, 1814) Caledonian populations of Clathria (Thalysias) hirsuta Hooper & Levi. measurements in i.tm, denoted (Figs 176-177) as range (and mean) of spicule length x spicule width Spongia juniperina Lamarck, 1814: 444; Lamarck, (N=25).zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 1816: 373. Specimens Specimens Holotype Clathria juniperina; Hooper & Wiedenmayer, 1994: (N= 6) (N= 3) (New SPI CULE (QMGL2746) 270. (Queensland) Caledonia) (Cairns region) Not Pandaros juniperina; Duchassaing & Michelotti, 1864: 90, p1.19, fig.3. Clioaposomal 1634178.3)-194 132-(155.2)-I95 96-(141.4)-I68 pnncipal Not Thalysias juniperina; de Laubenfels, 1936a: 105x 2-(4.8)-8 x 4-(7.8)-12 x 3-(5.8)-I2 styles 107 (see synonymy for T. virgultosa below). Sub.ectosomal 179-(241.8)-284 1414216.9)-293 163-(204.3)-248 Rhaphidophlus clathratus; Hallmann, 1912: 209; Topauxiliary sent, 1920b: 17-18; Topsent, 1932: 97, p1.5, fig.6, x I.5-(3.4)-5 x 1.5-(3.1)-5 x I.8-(3.1)-4.0 styles text-fig.3. Ectosomal 74-(100.1)-115 72-(92.0)-111 Not Tenacia clathrata Schmidt, 1870: 56, 80. auxiliary styles Echinating acanthostyles absent 37-(57.8)-73 x 2-(4.4)-7 x 1.5-(2.1)-3 x 0.8-( 1.9)-2.5 4I-(60.4)-79 x 2.5-(4.4)-8 34-(51.9)-72 x 2-(3.5)-5 Chelae I 3-(4.8)-6 3-(4.3)-6 6-9, rare Chelae II 9-(10.8)-I2 9411.5)-14 10-12, rare 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 differentiated from the subectosomal auxiliary spicules, an 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 by Hooper & Levi (1993a). 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 consequently 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. MATERIAL. LECTOTYPE: MNHNDT570: SW. coast 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. 176J). 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. Oscules. Not observed Texture and surface characteristics. Harsh, firm in dry state; surface characteristics range from relatively smooth, even, with white subderrnal canals in encrusting forms, to irregularly microconulose or clathrous in more massive forms. Ectosotne and subectosome. Ectosomal skeleton crust-like, easily detachable, relatively thin but dense palisade of erect or paratangential brushes supported by paratangential tracts of larger subectosomal auxiliary megascleres immediately below surface; peripheral fibres immediately subectosomal with vaguely ascending multispicular subectosomal tracts arising to surface. Choanosome. Choanosomal skeleton irregularly reticulate, with very heavy spongin fibres forming 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 typically heavily echinated, some enveloping echinating megascleres entirely, some fibres without echinating megascleres; mesohyl matrix light, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 347 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, L,ectotype. I, Paralectotype MNHNDT3354. J, Australian distribution. with numerous choanosomal styles dispersed be- (sinuous), with smooth subtylote bases. Length tween fibres.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 169-(253.5)-310p,m, width 4-(5.4)-6.51.1..m. Megascler es. Choanosomal principal styles Ectosomal auxiliary subtylostyles with straight or slightly curved near basal end, with prominent subtylote, smooth bases. Length 93smooth, rounded or very slightly subtylote bases. Length 1704244.4)-28011m, width 9-(10.1)- (102.3)-110ii.m, width 2-(3.7)-4.5Rm. Acanthostyles small, stubby, with rounded or 12Known only from Australia: m. only slightly subtylote bases, with few spines and Subectosomal auxiliary subtylostyles straight or curved, sometimes with multiple curves extensive aspinose regions on necks and points; MEMOIRS OF THE QUEENSLAND MUSEUM 348zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 177. Clathria (Thalysias) juniperina (Lamarck) (lectotype MNHNDT570). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostylc. D, Acanthostyle spines. E, Palmate isochclae. F, Accolada, sinuous and U-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 349 spines large, bulbous, erect. Length 45457.0)65p.m, width 5-(6.1)-81.1.m. Microscleres. Palmate isochelae unmodified, incompletely differentiated into two size classes; lateral alae entirely fused to shaft, approximately same length as front ala and completely detached from front ala. Length I: 647.1)-9p,m, length II: 12413.8)-16.5pm. Toxas very variable in length, mostly thin, ranging from accolada forms with large central curvature and slightly reflexed points, large curvature and simply u-shaped, to asymmetrical sinuous forms. Length 554122.4)-180p,m, width 0.841.1)-1.5p.m. REMARKS. C. (T) juniperina differs from that of Hartman (1955), Simpson (1968a), Wiedenmayer (1977) and Van Soest (1984b), who included Spongia juniperina, Spongia virgultosa, Clathria copiosa, Microciona 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.e., nominotypical population of Spongiajuniperina) inferring a widely disjunct geographical distribution. Three explanations are possible: 1) W. Indian 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 distributed '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) virgultosa 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 conspecific. 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 depauperate skeleton (fibres shed some or all of their coring spicules) and principal and auxiliary spicules are similar in geometry. Clathria (Thalysias) kieschnicki Hooper, in Hooper & Wiedenmayer, 1994 Rhaphidophlus cylindricus Kieschnick, 1900: 569, p1.44, fig.10. Not Esperiopsis cylindrica Ridley & Dendy, 1886: 340. Clathria (Thalysias) kieschnicki Hooper, in Hooper & Wiedenmayer, 1994: 271. MATERIAL. None. Holotype PMJ missing; (F. Wiedenmayer, pers. comm.). HABITAT DISTRIBUTION. Ecology unknown; Thursday I., Torres Strait (FNQ). DESCRIPTION. Shape. Bifurcate digitate, with short cylindrical 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 conules. Texture and surface characteristics. Fragile; surface with prominent, irregularly distributed conules, and detachable skin-like crust. Ectosome and subectosome. Ectosome relatively thick, with discrete plumose bundles of ectosomal auxiliary subtylostyles, forming a continuous palisade, below which plumose tracts of subectosomal auxiliary subtylostyles protrude through ectosome. Choanosome. Choanosomal skeletal architecture irregularly reticulate, with heavy, lamellated spongin fibres, 60-160p,m diameter, not obviously divisible into primary or secondary elements, forming ovoid meshes, 90-150pm diameter; fibres cored by irregular multispicular tracts of choanosomal principal styles and also fewer subectosomal auxiliary subtylostyles; echinating acanthostyles abundant, evenly distributed. Megascleres. Choanosomal principal styles straight or slightly curved, with smooth bases. Length 90-180p,m, width 9-15p.m. Subectosomal auxiliary subtylostyles straight, with microspined bases. Length up to 270p,m, width up to 18 p.m. Ectosomal auxiliary subtylostyles are fusiform, straight or slightly curved, with microspined bases. Length 135-230p,m, width 4-8 Rm. Acanthostyles subtylote, with bare necks. Length up to 135p.m, width up to 181.m. Microscleres. Palmate isochelae. Length 15 p.m. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 350zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Toxas thin (but of unknown geometry), occurring in trichodragmata or singly. Length 70100p.m. REMARKS. This species is poorly characterised because Kieschnick's (1900) description is brief and does not differentiate it from other arborescent, 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 the 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, p1.4, fig.5, p1.6, fig5. Clathria (Thalysias) koltuni Hooper, in Hooper & Wiedenmayer, 1994: 271. MATERIAL. None: `Syntypes' (ZIL 10637, 11437) (not seen). HABITAT DISTRIBUTION. Substrate unknown; 610-860m depth; Budd Coast, Wilkes Land, Antarctica. DESCRIPTION. Shape. Thinly encrusting, up to only 1mm thick. Colour. Red alive. Oscules. Not seen. Texture and surface characteristics. Even surface. Ectosome and subectosome. Erect choanosomal principal subtylostyles protruding through surface with bundles of smaller auxiliary subtylostyles dispersed around principal spicules. Choanosome. Hymedesmoid, with choanosomal principal and subectosomal auxiliary subtylostyles erect on basal spongin; bundles of echinating acanthostyles clumped around erect structural megascleres. Megascleres. Choanosomal principal subtylostyles long, straight, fusiform, with subtylote bases and evenly microspined in basal third of spicule. Length 400-750[Lm, width 26-36p.m. Ectosomal and subectosomal auxiliary styles very long, slender, straight, fusiform, with microspined subtylote bases. Length 4306301im, width 8-12p,m. Echinating acanthostyles short, club-shaped, fusiform, with prominent subtylote base and evenly spined over entire length of spicule. Length 100-260p,m, width 10-14p.m. 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 previously referred to Pseudanchinoe), in spicule dimensions and spicule ornamentation, but has few other noteworthy features. antarcticus is 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, p1.28, fig.5, p1.29, fig.6, p1.47, 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, textfig.19. Thalysias lendenfeldi; de Laubenfels, 1936a: 105. Microciona lendenfeldi; Dawson, 1993: 37. Spongia abietina, in part, Lamarck, 1814: 450. Echinonema anchoratum var. lamellosa; Whitelegge, 1901: 82. Not Echinonema anchoratum var. lamellosa Lendenfeld, 1888: 219. Clathria spicata Hallmann, 1912: 210; Dendy, 1922: 65-66, p1.5, fig.2, p1.13, fig.4a-f; Burton, 1959a: 244. Clathria diechinata Hallmann, 1912: 211; 1914a: 268 [nomen nudurn]. Thalysias spicata; de Laubenfels, 1936a: 105. ClathriawhiteleggiiDendy, 1922: 67, p1.7, fig.1, p1.13, figs 5a-f; Burton, 1931a: 344-345; Burton, 1959a: 245; Levi, 1963: 66. Thalysias whiteleggei; de Laubenfels, 1936a: 105. Clathria coppingeri var. aculeata Hentschel, 1912: 363. Rhaphidophlus bispinosus Whitelegge, 1907: 503504. Clathria bispinosa; Hallmann, 1912: 177,211. cf. Microciona prohfera; Vosmaer, 1935a: 610, 636, 669. MATERIAL. HOLOTYPE: BMNH1887.5.2. 107: off Port Jackson, NSW, 33°40'S, 151°40'E, HMS ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 351 FIG. 178. Clathria (Thalysias) lendenfeldi Ridley & Dendy (NTMZ2095). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyles. 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, QMG300830, QMG303507, QMG303523, QMG304946. NT - NTMZ2821. WA - NTMZ3060, NTMZ3384, QMG310535 (NCIQ66C-1518-Q) (fragment NTMZ3489), QMG310423 (NCIQ66C-1291-T) (fragment NTMZ3463), QMG310423 (NCIQ66C-1318-X) (fragment NTMZ3468). TAS - QMG311436 (NCIQ66C-3745-M) (fragment NTMZ3822). ANDAMAN SEA, THAILAND - NTMZ3657, NTMZ3659. RED SEA - PIB0004-11 (fragment QMG300057). SOMALIA, EAST AFRICA - PIBOCB12-367 (fragment QMG300062). HABITAT DISTRIBUTION. Rock reefs and dead coral heads; intertidal to 108m depth; widespread IndoPacific; 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), 352zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 353 Red Sea and Somalia (present study), Saya de Malha (Dendy, 1922) to the Natal coast (Burton, 1931a). DESCRIPTION. (See Hooper et al., 1990). REMARKS. Variation has been comprehensively 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 C.(T.) major, differing only substantially by its hair-like toxas, sharply pointed auxiliary megascleres, statistical differences in spicule dimensions and various biochemical features (Hooper et al., 1990), and both are members of Hallmann's (1912) spicata' group. 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 microClathria (Thalysias) major Hentschel, 1912 conulose; ectosome ranging from sparsely scat(Figs 180-181) tered smaller auxiliary spicules to dense, erect, continuous crust; choanosomal principal sub- Clathria frondifera var. major Hentschel, 1912: 361. tylostyles also protrude through surface; subec- Clathria (Thalysias) major; Hooper et al., 1990: 133135, figs 1, 3, 5, 6; Hooper & Wiedenmayer, 1994: tosomal skeleton poorly developed, 272. paratangential, composed of larger auxiliary subtylostyles; choanosomal skeleton irregularly MATERIAL. HOLOTYPE: 5MF977 (fragment reticulate, heavy spongin fibres divided into MNHNDCL2303): Straits of Dobo, Aru I., Indonesia, primary (ascending) and secondary (transverse) 6°S, 134°50'E, 40m depth, 20.iii.1908, coll. H. Merton components, producing regular or irregular (dredge). OTHER MATERIAL: (Hooper et al., 1990) meshes; fibre-meshes heavier in axis; fibres WA - NTMZ3338, NTMZ3360, CSIROEMG001. generally uncored, some with uni- or pauci- HABITAT DISTRIBUTION. Rock reefs and dead spicular tracts of principal spicules, and abun- coral heads; intertidal to 82m depth; Bedout I., Port dantly echinated by both acanthostyles and Hedland, Mary Anne I., Direction Is, Exmouth Gulf principal subtylostyles especially at fibre junc- (WA); Bynoe Harbour, Darwin Harbour, Port Estions ('spicate'); choanosomal principal sub- sington (NT) (Fig. 180H); also Aru Is, Indonesia tylostyles long, curved or straight, sharply (Hentschel, 1912). pointed, usually with heavily spined bases (119- DESCRIPTION. (See Hooper et al., 1990). (229.6)-492pm x 1.8-(12.9)-35p,m); subectosomal auxiliary subtylostyles long, straight, DIAGNOSIS. Variable growth forms ranging fusiform, slightly subtylote, microspined bases from low, foliose, bushy, subspherical, clathrous (136-(241.5)-404Rm x 2-(4.6)-15Rm); ec- digitate, to flabellate or digitate fans, usually with tosomal auxiliary subtylostyles short, thin, long basal stalk and flattened or irregularly straight, fusiform, with microspined subtylote cylindrical branches; bright red to orange-red bases (62-(123.4)-194,m x 2-(3.0)-10Rm); alive; oscules small, congregated into special acanthostyles slender, long or short, fusiform, pore areas on lateral sides of branches or between slightly subtylote, evenly and lightly spined, surface conules; flabellate specimens may have spines small, recurved (49488.1)-151pm x 2- Phakellia-like pores grouped into stellate pore(6.4)-141,m); palmate isochelae small, rarely areas; surface irregularly microconulose with close-set subdermal ridges and striations; ecmodified, narrow lateral alae completely fused to tosomal skeleton ranges from very few tangenshaft, approximately same length as front ala; tially placed ectosomal auxiliary subtylostyles to lateral alae completely and widely separated from dense, erect or paratangential brushes of ecfront ala (6-(12.5)-25p.m long); toxas accolada to tosomal spicules; subectosomal skeleton wing-shaped, very thin (hair like); larger toxas plumose, paratangential tracts of larger auxiliary usually accolada, straight, with slight but sharp subtylostyles; both larger auxiliary and principal angular central curvature and unreflexed arms; spicules protrude through surface singly or in smaller toxas usually wing-shaped, with large plumose bundles; choanosomal skeleton ircentral curvature and slightly reflexed arms; regularly reticulate; fibre characteristics, skeletal toxas found singly and in bundles (dragmata) structure and distribution of megascleres and microscleres identical to C. (T) lendenfeldi; X 0.4-(1.3)-3.611M). (7-(136.4)-3611IM zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 354zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA CD 0 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)-38iim x 5-(15.1)-36p,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)-43911m x 2-(5.7)-141J.m); ectosomal auxiliary subtylos- tyles short, straight, subtylote microspined bases, usually with rounded apex (quasi-tornotes) and terminal spines (84-(136.8)-193p,m x 243.8)9p,m); acanthostyles relatively slender, long or short, fusiform, subtylote, with large spines on base and apex but nearly aspinose 'neck' (77 (112.7) 144Rm x 3-(7.8)-15p.m); palmate isochelae small, unmodified, wide lateral alae completely fused to shaft, approximately same - - ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 355 FIG. 181. Clathria (Thalysias) major Hentschel (QMG300153). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E-F, Bases and apex of principal and auxiliary subtylostyles. G, Palmate isochelae. H-I, Accolada toxas. 356zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 0 0 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, Oxhom - 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)-16Rm long); toxas accolada or wing-shaped, the former long, very thick, with large rounded central curvature, straight or reflexed arms, the latter short, thin, widely curved at centre with reflexed arms (27-(108.9)-390vm x O.6-(2.3)-5m). REMARKS. The species is a cryptic sibling of C. (T) lendenfeldi with a sympatric but more restricted distribution. In gross morphology, sur- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 357 face features, live colouration and in many of its skeletal characters C. (T.) major is indistinguishable from its sibling. However, they can be reliably 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 significant quantities of the chemical 2,6dibromophenol of potential commercial importance as an lodoform' 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: UM (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 protruding 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 subtylostyles long, fusiform, slightly curved, subtylote, with microspined bases. Length 1884381.6)646p,m, width 10-(14.6)-19p.m. Subectosomal auxiliary subtylostyles long, thin, fusiform, prominently subtylote, lightly microspined, occasionally with smooth bases. Length 3074403.6)-48211m, width 3-(4.4)-611m. Ectosomal auxiliary subtylostyles, short, thin, straight, prominently subtylote, smooth bases, usually polytylote shafts. Length 1414162.4)197p,m, width 2-(2.9)-41J.m. Acanthostyles long, thin, slightly subtylote, with lightly microspined base and central portions, aspinose points and neck regions. Length 96-(108.8)-125p,m, width 3-(6.6)-41..m. Microscleres. Isochelae sigmoid (bidentate) anchorate, with small alae attached only at their bases. Length 15-(17.2)-19Rm 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 384122.6)-23911m, width 1-(3.5)-6p.m. REMARKS. Hentschel (1911) initially overlooked the presence of toxas in this species, although 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 published 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 stellidenna Carter). Clathria (Thalysias) phorbasiformis sp. nov. (Figs 183-184, Plate 7F-G) MATERIAL. HOLOTYPE: NTMZ2138: Dudley Point, East Point Aquatic Life Reserve, Darwin Harbour, NT, 12°25.0'S, 130°49.1'E, intertidal, 27.ix.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 MEMOIRS OF THE QUEENSLAND MUSEUM 358zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ OF1DzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 0 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.i.1985, QMG300149 (fragment NTMZ2223), QMG300150 (fragment NTMZ2237). coral boulders or in crevises and pools; intertidal; Darwin Harbour (NT) (Fig. 183H). HABITAT DISTRIBUTION. Encrusting on laterite rock, dead coral, exposed at ELWS tides, under dead Shape. Thickly encrusting, 0.7DESCRIPTION. zyxwvutsrqponmlkjihgfedcbaZYXWVU 1.3cm thick, producing thin cylindrical ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 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. 360zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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, 0.10.25mm 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 ectosomal 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 protruding, 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; subectosomal region variable in thickness dependent on proximity of peripheral fibres to surface, containing plumose columns of larger auxiliary subectosomal subtylostyles, not associated with fibres, but often bound together by collagen; subectosomal 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 architecture 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 principal subtylostyles, in plumose tufts or singly, particularly abundant at fibre nodes; fibre anastomoses form oval or elongate meshes, 250-800Rm diameter; fibres thicker in deeper areas of choansome (70-100gm diameter) than in periphery (55-801.1,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 ectosomal 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 randomly throughout mesohyl. Thinly encrusting growth forms — hymedesmoid skeletal construction with basal layer spongin lying on substrate, uncored but very heavily echinated by both acanthostyles and choanosomal subtylostyles standing perpendicular 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. Me gascleres. Choanosomal principal subtylostyles fusiform, tapering to long points, slightly curved near the basal end or occasionally straight, with subtylote, mostly smooth bases, occasionally roughened subapically or slightly tubercular. Length 245.2-(425.8)-583.1p,m, width 10.2(19.5)-33.811m. Subectosomal auxiliary subtylostyles long, fusiform, mostly straight, with subtylote microspined or occasionally smooth bases. Length 275-(386.2)-485.3p,m, width 4.049.6)Ectosomal auxiliary subtylostyles relatively short, straight or whispy, slender, with subtylote, relatively heavy basal microspination. Length 70.04140.5)-261 .2pLm, width 1.2-(4.0)-6.9p,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)-132.4p,m, width 4.6-(8.4)-12.711m. Microscleres. Palmate isochelae abundant, subdivided into 2 size categories, smaller ones sometimes contort; lateral alae completely attached to shaft, approximately same length as front ala but completely detached from it. Length!: 10-(14.9)21.9p.,m, length II: 2.5-(5.6)-9.2Rm. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 361 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(95.1)-222.9p,m, width 0.5-(1.4)-2.5 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 subtylostyles 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 Hallmann's spicata group (see C. (T) coppingeri). Incorporation of echinating acanthostyles secondarily into fibres has been observed in some specimens of C. (Dendrocia)dura, C. (D.) imperfecta, and to a lesser degree C. (D.) myxilloides, but these instances are infrequent, inconsistent (i.e., seen in some sections of the skeleton but not in others), and probably aberrant. A similar phenomenon has been described for C. (T.)orientalis by Brondsted (1934) but this too is atypical for the species (whereby the larger auxiliary subtylostyles usually core fibres). Analogous structures 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) phorbasifonnis differs from other species in the phorbasiformis' complex in gross morphology and spicule geometry. In live surface features and colouration it some resemblance to thickly encrusting C. (T) reinwardti, although spicule geometry, spicule size, skeletal architecture and fibre characterisitics are clearly different between them. Clathria (Thalysias) placenta (Lamarck, 1814) (Figs 185-186) Spongia placenta Lamarck, 1814:374; 1815:356. Wilsonella placenta; Topsent, 1930:24, p1.3, fig.8. Clathria placenta; Hooper & Wiedenmayer, 1994: 273. Not Microciona placenta; de Laubenfels, 1954:146147, text-fig.94. MATERIAL. HOLOTYPE: MNHNDT552: King I., Bass Strait, Tas, 39°50'S, 144°00'E, Peron & Lesueur collection. HABITAT DISTRIBUTION. Ecology unknown; Bass Strait, Tasmania (Fig. 185G). DESCRIPTION. Shape. Thick, flabellate growth form, 180mm high, 170mm wide, up to 1 Omm thick, with even margin; probably originally 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 pattern. Ectosome 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 subectosomal spicule tracts, the latter embedded in peripheral skeleton; choanosomal fibres immediately subectosomal. Choanosotne. Choanosomal skeleton irregularly reticulate with primary (ascending) and secondary (transverse) fibres; primary fibres (105175pm diameter) cored by multispicular tracts of subectosomal auxiliary styles, occupying up to 60% fibre diameter, tracts becoming plumose peripherally; secondary fibres (35-88p.m diameter) without coring spicules; all fibres heavily echinated by small acanthostyles sometimes nearly enveloped in spongin; fibre anastomoses form irregular oval and rectangular meshes (1455101.1m diameter); mesohyl matrix light, with few loose megascleres dispersed between fibres. Megascleres. Choanosomal principal megascleres absent or completely undifferentiated 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)-285pm, width 5-(6.6)-8pm. 362zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM A Cs/ FIG. 185. Clathria (Thalysias) placenta (Lamarck) (holotype MNHNDT552). 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 52454.4)(138.1)-156p.m, width 2-(2.6)-4‘m. 5811,m, width 3.5-(4.2)-6p.m. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 363 Clat hria ( Thalysias) placent a (Lamarck) (holotype MNHNDT552). A, Skeleton, ectosome detached. FIG. 186. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Palmate isochelae. F, Accolada toxas. 364zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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)-1411m. Toxas accolada, moderately long, thick, with only very slight central curvature and straight points. Length 105-(119.5)-148Rm, width 1.0(1.4)-2.0ikm. REMARKS. Lamarck's (1814) holotype from Bass Strait and de Laubenfels'(1954) sample USNM22908 from Truk, Caroline Is are not conspecific; the latter becomes C. (T.) lematolae sp. nov. The Truk sample is only similar to C. placenta in having fibres cored by subectosomal auxiliary spicules instead of principal spicules (i.e., the principal and larger auxiliary spicules are undifferentiated in their geometry). In most other details the two species can be readily distinguished (the Chuuk sample has an encrusting growth form, an extremely smooth surface, skeletal structure is hymedesmoid including possession of a very extensive subectosomal skeleton, occupying almost half of the sponge diameter, acanthostyles are about twice the size of those in C. placenta with much more robust spination, toxas are slightly accolada but moreso wingshaped, megascleres are mostly subtylote, and dimensions of most spicules differ). Topsent (1930) implied that C. (T.) placenta) was similar to C . (Wilsonella) australiensis (Carter), 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)placenta is a member of the juniperina' complex having a 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.) cactifonnis). Clathria (Thalysias) procera (Ridley, 1884) (Figs 187-188, Table 38) Rhaphidophlus procerus Ridley, 1884a:451-452, p1.39, fig.k, p1.42, fig.o; Burton, 1931a:343, p1.23, fig.2. Clathria procera; Dendy, 1922:64, p1.2, figs 6-7; Burton, 1938a:28-29; Burton, 1959a:243; Levi, 1963:66; Bergquist, 1967:164-165, text-fig.3; Thomas, 1973:34-35, p1.2, fig.5, p1.7, fig.3; Bergquist, 1977:65; & Wiedenmayer, Hooper, 1994: 273. Tenacia procera; Burton & Rao, 1932:340; Burton, 1934a:559; Burton, 1934b:28. Rhaphidophlus spiculosus Dendy, 1889b:75, 86, 87, 99, p1.4, fig.4 [Gulf of Manaar, Ceylon]; Dendy, 1922:64. Clathria spiculosa; Dendy, 1905:171-173, p1.8, fig.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, 1912:363364. Not Clathria spiculosa var. macilenta; Hentschel, 1912:364 [Aru I., Arafura Seat Echinonema gracilis Ridley, 1884a:617, p1.54, fig.1; Dendy, 1922:64. Rhaphidophlus gracilis; Ridley & Dendy, 1887:152, 242, 252; Topsent, 1892b:24. Clathria gracilis; Dendy, 1905:171; Vosmaer, 1935a:634. Not Rhaphidophlus arborescens Ridley, 1884a:450451, p1.40, fig.L, p1.42, fig.n; Burton & Rao, 1932:340. cf. Microciona prolifera; Vosmaer, 1935a:610, 634, 669. MATERIAL. HOLOTYPE: BMNH1882.2.23. 313: Off East Point, Port Darwin, NT, 12°24.5'S, 130°48.0'E, 14-22m depth, coll. HMS `Alert' (dredge). PARATYPE: BMNH1882.2.23.311: same locality. HOLOTYPE of R. spiculosus: BMNH1889.1.21.5 (fragment BMNH1954. 2.23.101): Gulf 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, 9°14S, 51°02'E, 48m depth (dredge). HOLOTYPE of C. spiculosa var. ramosa: SMF1698 (fragment MNHNDCL 2304): Straits of Dobo, Am I., Arafura Sea, Indonesia, 6°S, 134°50'E, 20.iii.1908, 40m depth coll. H. Merton (dredge). OTHER MATERIAL: QLD- NTMZ3983, QMG 301032, QMG303514QMG304392, QMG304771. NT- NTMZ2604, QMG303582. WA- NTMZ1308. INDIAN OCEAN B MN H1907 .2.1.63, BMNH 1954.2.23.113, BMNH1954.2.23.114. HABITAT DISTRIBUTION. On loose, soft substrates (sand, mud, gravel, shell grit) associated with shallowwater or deeper offshore reefs; 11-78m depth; widespread throughout the Indian Ocean and Indowest Pacific; Gulf of Carpentaria, Low Is, Direction Is, Snake Reef, Turtle Is (FNQ); Bynoe Harbour, Darwin Harbour, Cape Wessel, Arafura Sea (NT); Port Hedland (WA)(Fig. 187H); also Scottburgh, Natal (Burton, 1931a; Levi, 1963), Cargados Carajos, Seychelles, Amirante, Red Sea and Arabian Sea (Ridley, 1884a; Ridley & Dendy, 1887; Dendy, 1922; Burton & Rao, 1932; Burton, 1959a; Thomas, 1973b); Tuticorin, Cape Comorin, Palk Straits, and Madras Straits, Gulf of Manaar (Burton & Rao, 1932; Burton, 1938a), Aru Is, Indonesia (Hentschel, 1912), Hawaii (Bergquist, 1967; 1977). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 365 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 flattened 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 attachment to substrate expanded, rhizomous; gross morphology superficially resembles Junceela gorgonian whip-coral. MEMOIRS OF THE QUEENSLAND MUSEUM 366zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 38. Comparison between present and published records of spicule diverging) towards dimensions for Clathria (Thalysias) procera (Ridley). Measurements in 11,m periphery; fibres imperfect(N=25). ly divided into ascending primary fibres (40-751i,m Specimen Specimen Clarhria diameter) and transverse HoIotype (N=1) (N=1) spiculosa var. Specimens , (BMNH1882.2., secondary components SPICULEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA (Thomas, ram osa (Burton, (N=8)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 3.313) (130-210vm diameter); 1973b) (SMFI 698) 1938) Choanosomal styles 322-(334.2)348 x 11(13.2)-16 (common) 2484283.2)309 x 16(18.2)-22 (uncommon) 263-288 x 1518 (rare) Subectosomal styles 318-(334.2)358 x 8-(9.2)11 342-(367.1)393 x 8-(11.3)14 276-345.5 x 9.2-12.5 Ectosomal styles I72-(258.4)295 x 3-(5.6)-7 124-(178.2)290 x 2-(3.8)-5 112.3-235.6x 2.2-5.8 Acanthostyles 91-(99.2)-110 x8-(11.0)-13 89-(104.1)-114 x7-(11.6)-16 72-98.9 x 912.1 Chelae I 12-(15.3)-18 13-(15.6)-19 14.5-19.2 Chelae II 4-(6.6)-10 6-(8.3)-10 6-10.5 18-(60.5)-122 x 0.8-(1.3)- I .5 31-(101.2)-145 x 0.9-(1.2)-1.8 62-142x 1.2-2 Toxas 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 obvious stiff axis; basal region woody, more rigid than central or apical regions; surface optically smooth, without conules or other surface processes, microscopically hispid with minute subdermal canals and grooves. Ectosome and subectosome. Well developed series of erect spicule brushes forming a continuous palisade, composed of ectosomal auxiliary subtylostyles; ectosomal brushes embedded on ultimate fibres, with echinating acanthostyles and choanosomal principal styles protruding through bases of each spicule brush; subectosomal auxiliary subtylostyles form tangential or paratangential tracts below ectosomal skeleton; choanosomal principal styles embedded 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, yellow spongin fibres forming tight anastomoses near core, becoming more plumose (or merely primary fibres multispicular, cored by subec201-310 x 4-12 tosomal auxiliary subtylostyles occupying up to 90% of fibre diameter; 210-294 x 4-8 280 x 8 secondary fibres less heavily cored, occasionally 180-200 unispicular; fibre reticula100 x6 tion producing irregularly oval or eliptical meshes at 58-75 x 4-9 present core (110-275Rm 12-16 16 diameter), becoming wider, 9 more rectangular at periphery (230-425p.m <147 45-56 diameter); echinating acanthostyles more heavily concentrated 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 supporting ectosomal skeleton; mesohyl matrix relatively light in axial region with many loose subectosomal auxiliary megascleres scattered between fibres. 220-360 x 13 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 367 FIG. 188. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hr ia ( Thalysias) pr ocer a (Ridley) (QMG300166). A, Choanosomal skeleton. B, Fibre characteristics (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. 368zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Microscleres (Table 38). Palmate isochelae incompletely divided into 2 size categories, both abundant, larger unmodified, smaller often contort (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, variable in length, with pronounced central curvature, slightly reflexed or straight points. REMARKS. In live colour, surface characteristics, texture, gross morphology, spiculation and skeletal architecture this species is quite distinctive. 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. (T) cactifonnis although the 2 species are not conspecific as supposed by Burton & Rao (1932), where C. (T) cactifonnis has an aspicular secondary fibre skeleton and lacks any axial compression. The principal megascleres echinating fibres and absence of principal spicules from within the fibre core indicates it belongs to Hallmann's (1912) spicata' group. Records of C. procera subsequent to Ridley (1884a) make no mention of choanosomal principal spicules echinating peripheral fibres, although this feature is characteristic for the species. Conversely, authors following Dendy (1922) note that there are two classes of auxiliary megascleres, both of similar length but different thickness, the thicker ones coring fibres and the thinner ones scattered in the mesohyl, but this distinction was not corroborated from re-examination of any material. Hallmann (1912), Dendy (1922), Burton & Rao (1932), Burton (1938a) and subsequent authors included a number of other species as synonyms of C. (T) procera, but most of these synonymies are not supported here. Clathria spiculosa var. macilenta is certainly different from C. (T) procera and is clearly a synonym of C. (T) reinwardti. In contrast, Clathria spiculosa var. ramosa Hentschel (SMF1698) is conspecific with C. procera, having 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, thin, evenly cylindrical branches bifurcating but not anastomosing, and bifurcate branch tips superficially 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 (1931a, 1938a) 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 (1935a) C. (T) arborescens is a distinct species from C. (T) procera, both species differing substantially in their spicule geometry, spicule sizes and skeletal architecture. Clathria reinwardti var. palmata 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 to C. (T.) procera, but shows virtually no axial compression of the choanosomal skeleton, spicule geometry is clearly different, and the two species are not considered to be synonyms. Clathria (Thalysias) ramosa (Kieschnick, 1896) (Figs 189) Rhaphidophlus ramosus Kieschnick, 1896:533; Kieschnick, 1900:569-570, p1.45, figs 47-50. Clathria ramosa; Hooper & Wiedenmayer, 1994: 273. Not Clathria ramosa Lindgren, 1897:482 483; Lindgren, 1898:308-309, p1.17, fig.9, p1.18, fig.15, p1.19, fig. 16; Hentschel, 1912:367. Not Thalysias ramosa; de Laubenfels, I936a:105. Not Colloclathria ramosa Dendy, 1922:74-76. - MATERIAL. HOLOTYPE: PMJ Porif.92: Thursday I., Torres Strait, Qld, 10°35'S, 142°13'E, no other details 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 369 A CD curved, with basal spination. Length 150-480p,m, width 13-2411m. Subectosomal auxiliary subtylostyles long, thin, prominently subtylote with microspined bases. Dimensions unknown. Ectosomal auxiliary subtylostyles identical in geometry to larger auxiliary spicules. Dimensions unknown. Acanthostyles cylindrical, club-shaped, evenly spined, subtylote. Length up to 150p,m, width 9-13p.m. Microscleres. Palmate isochelae in 2 size classes. Length up to 13p.m. 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 inclusion of the species in Rhaphidophlus. Until the presently missing holotype is re-examined, the affinities of this species remain uncertain. Clathria (Thalysias) reinwardti Vosmaer, 1880 (Figs 190-192, Table 39, Plate 8A-B) Clathria reinwardti Vosmaer, 1880:152; Vosmaer, FIG. 189. Clathria (Thalysias) ramosa (Kieschnick) (redrawn from Kieschnick, 1900). A, Choanosomal 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 significantly in diameter; fibres only lightly cored by choanosomal principal subtylostyles within axis of skeleton, occasionally absent; fibres usually more heavily cored towards periphery; echinating acanthostyles abundant; character of soft parts unknown. Megascleres. Choanosomal principal subtylostyles prominently subtylote, straight or slightly 1935a:610, 632, 639; Bergquist & Tizard, 1967:184-186, p1.4, fig.2; Bergquist et al., 1971:102-106; Van Soest, 1989:223, fig.34; Hooper & Wiedenmayer, 1994: 273. Clathria reinwardti var. subcylindrica Ridley, 1884a:446-448. Rhaphidophlus reinwardti; Kelly Borges & Bergquist, 1988:141-143, figs 3-4, p1.3f. Clathria typica var. porrecta Hentschel, 1912:298, 359-360. Tenacia typica var. porrecta Hallmann, 1920:771. Clathria spiculosa var. 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. macilenta: SMF1514 (fragments MNHNDCL2242, 2250): Straits of Dobo, Am I., Arafura Sea, Indonesia, 6°S, 134°30'E, 40m depth, 20.iii.1908, coll. H. Merton (dredge). LECTOTYPE of C. reinwardti var. subcylindrica BMNH1881.10.21.260: 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 MEMOIRS OF THE QUEENSLAND MUSEUM 370zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ C. typica var. porrecta: SMF1653 (fragment MNHNDCL2228): Straits of Dobo, Aru I., Arafura Sea, Indonesia, 6°S, 134°50'E, 22.iii.1908, 12m depth, coll. H. Merton (dredge). OTHER MATERIAL: NT AMZ3099, AMZ4311 (RRIMP-0917); NTMZ176, NTMZ177, NTMZ270; NTMZ1094, NTMZ2080, NTMZ2121, NTMZ2206, NTMZ 2211, NTMZ2227, NTMZ2232, NTMZ2264, NTMZ2389, NTMZ2423, NTMZ2543, NTMZ 2545, NTMZ2554, QMG300179 (NTMZ2717), NTMZ472, NTMZ228, NTMZ435, NTMZ2174, NTMZ2197, QMG303260, NTMZ3150, NTMZ55, NTMZ77, NTMZ348, NTMZ350, NTMZ359, NTMZ361, NTMZ362, NTMZ363, NTMZ364, NTMZ441, NTMZ1364, NTMZ 1371, NTMZ1378, NTMZ2493, NTMZ2514, NTMZ3299, NTMZ3308, NTMZ570, NTMZ 586, NTMZ574, NTMZ1327, NTMZ2502, NTMZ3242, NTMZ3247, NTMZ3251, NTMZ 3254, NTMZ3256, NTMZ3264, NTMZ3271, NTMZ3275, NTMZ3279, NTMZ3288, NTMZ 3296, NTMZ3310, NTMZ324, NTMZ333, NTMZ602, NTMZ38, NTMZ40, NTMZ50, QMG300753 (NCIQ66C-4677-Y, fragment NTMZ3906). WA - QMG301121, QMG301135, QMG301169, NTMZ3336 (NCIQ66C-1450-C). QLD- QMG300824, QMG304085, NTMZ4018, NTMZ4021, NTMZ4022, QMG303014, NTMZ4043. INDONESIA - QMG303687 (NCIOCDN-1285-H), SMF1589. PNG- NTMZ2561, NTMZ2562, NTMZ2563, NTMZ2564, QMG300371 (NCIQ66C4495-A), QMG300375 (NCIQ66C-4516-Y), QMG 300383 (NC1Q66C-4547-.1), QMG303104. PHILIPPINES - QMG300344, QMG300304 (NCIQ66C5727-Q). 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 0-10m depth, occasionally deeper. Speculated that association with dead coral substrates indicates some role in reef bioerosion; widely distributed throughout Indo-west Pacific; Darwin Harbour, Parry Shoals, Timor Sea, Port Essington, Orontes Reef, Trepang Bay, Cobourg Peninsula, Wessel Is (NT); Hibernia Reef, Cartier I., Sahul Shelf, Direction I. (WA); Gulf of Carpentaria, Cockburn Is, Cape York, Shelburne Bay, Blanchard Reef, Adolphus I. (FNQ) (Fig. 190H); also Cebu, Negros Orientale, Philippines (present study), Chuuk, Caroline Islands (present study), Hon Rai I., Vietnam (present study), Solomon Is (Bergquist et al., 1971), Motupore I., PNG (Kelly Borges & Bergquist, 1988; present study), Am Is, Sulawesi, Lesser Sumba Is, Indonesia (Vosmaer, 1935a; Van Soest, 1989, present study). DESCRIPTION. Shape. Typically simple digitate, stoloniferous, cylindrical or occasionally laterally compressed branches (7-25mm diameter), forming meandering digits with mul- tiple points of attachment to substrate; no differentiation between branches and stalk, with branches attaching directly to substrate; branches frequently anastomose with adjacent branches sometimes forming complex intertwined digits; free branches mostly simple, rarely bifurcate; several thickly encrusting, bulbous specimens also collected, presumably immature growth stages. Colour Very light orange (Munsell 5YR 8/4), orange-brown (7.5YR 8/2-4), orange-red-brown (2.5YR 7/8), light brown (10R 7/4), to grey-white (2.5Y 8/2) pigmentation alive, orange-brown (5YR 8/4) to grey-white (2.5Y 8/2) in ethanol; ectosomal membrane varies from colourless (opaque), to grey (2.5Y 8/2); subectosomal and choanosomal regions generally darker than periphery, usually clearly visible through ectosomal membrane when alive. Oscules. Abundant, relatively large (5602760t.un diameter), predominant on lateral sides of branches; oscules slightly raised with prominent membraneous lip (often orange pigmented) surrounding aperture; generally thicker specimens have larger oscules raised further above surface and more prominent subectosomal sculpturing; oscules collapse in air. Texture and surface characteristics. Soft, compressible, flexible, moderately easy to tear; surface smooth, pellucid, semi-translucent in life, with prominent stellate subectosomal channels radiating towards oscules, particularly in thicker specimens, more even surface ornamentation in thinner specimens; ectosomal membrane collapses upon dessication and preservation, becoming roughened and pocked with ridges and cavities. Ectosome and subectosome. Thin layer of smaller ectosomal auxiliary subtylostyles form discrete brushes erect on surface, in a continuous palisade, supported by long or short subectosomal plumose tracts from peripheral fibre skeleton; ectosomal region generally poorly collagenous but variable in thicker specimens; subectosomal region cavernous with lacunae (120-660p.m diameter) supported paucispicular plumose tracts 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 ectosomal skeleton. Choanosotne. Skeletal architecture irregularly or semi-regularly reticulate, with anastomosing spongin fibres (40-120p.m diameter) forming curved oval, straight triangular or rectangular meshes (50-47011m diameter) in choanosome; REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 371 A FIG. 190. Clathria (Thalysias) reinwardti Vosmaer (NTMZ2174). A, Choanosomal principal styles. B, Echinating 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. 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; abundant auxiliary spicules scattered between fibres; mesohyl matrix light, poorly pigmented, surrounding ovoid to eliptical choanocyte chambers (130-250p,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 strongylote points. MEMOIRS OF THE QUEENSLAND MUSEUM 372zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 39. Comparison between spicule dimensions (in p.m) between type specimens and other material of Clathria (Thalysias) reinwardti Vosmaer from different localities (N=25). SPICULE Choano- 1 2 4 3 5 6 7 111-(195.1)- 131-(201.6)- 1464192.9)- 110-(190.3)135-215 x 236 x 7259 x 7273 x 6280 x 56-10 (11.4)-21 (11.1)-16 (12.4)-18 (11.8)-21 I13-(235.5)- 141-(246.1)- 141-(219.7)- 113-(236.1)170-211 x 337 x 4301 x 3337 x 3326 x 33-9 (6.7)-14 (7.6)-11 (5.9)-11 (6.8)-16 66-(102.3)- 85-(106.5)- 82-(104.1)- 66-(100.7)84-102 x 3166 x 2162 x 2157 x 2170 x 26 (4.1)-7 (3.6)-7 (3.9)-7 (3.9)-8 42-(60.1)39-(57.9)3I-(59.1)45-(64.7)59-74 x 479 x 3-(7.4)- 78 x 3-(7.4)- 72 x 3-(6.6)- 79 x 3-(7.5)11 13 12 10 13 8 9 116-(195.6)- 1484200.5)264 x 9279 x 8(13.1)-21 (15.9)-21 155-345 (stout) 223-305 x 5-16 135_305 155_323 x (spinet') 3-12 120-130 (slender) 97-181 x 37 Acanthostyles 50-70 x 6.3 54-82 x 310 Chelae I 13-19 10-19 12-15 4-8 4-8 2-(6.I)-9 2-(5.4)-9 2-(6.0)-9 2-(5.8)-9 4-(6.6)-9 4-(6.4)-9 52-246 x 0.5-2 8-(121.2)237 x 0.5(1.2)-3.1 12-(119.3)186 x 0.5(1.2)-2.5 9-(102.5)173 x 0.5(1.2)-2.5 16-(125.8)236 x 0.5(1.3)-3.0 I2-(124.1)228 x 0.5(1.2)-2.5 24-(115.6)186 x 0.5(1.1)-2.0 somal styles Subectosomal styles Ectosomal styles Chelae II Toxas 22-174 x 0.5-2 146-(233.0)- 137(238.6)317 x 5317 x 5(7.2)-13 (7.6)-13 69-(102.7)- 73492.8)139 x 4170 x 4(5.1)-8 (3.9)-8 35460.3)31-(58.2)77 x 5-0.9)- 76 x 648.7)12 13 10-(14.4)-21 10-(14.0)-19 10-(14.3)-18 10-(14.2)-19 10-(14.9)-21 11-(13.9)-19 Source: subcylindr ica (BMNH1881.10.21.260). 3. Lectotype var. m acilent a 1. Holotype (Vosmaer, 1880). 2. Lectotype of var. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFE (SMF 1514). 4. Northern Territory specimens. 5. Papua New Guinea specimens. 6. Indonesian specimens. 7. Philippines specimens. 8. Micronesian specimen. 9. Vietnam specimen - Subectosomal auxiliary styles straight or sometimes slightly curved near basal end, relatively thick, with fusiform, sharp points, sometimes 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 abundant, scattered throughout mesohyl and lining choanocyte chambers; long lateral alae completely 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, hairlike, long, slight central curvature, slightly reflexed or with straight points; distributed singly or in trichodragmata throughout mesohyl. Larvae. 28% of specimens examined contained incubated parenchymella larvae in varying stages of development; larvae oval to eliptical, 180825Rm long, 80-400Rm 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.) reinwardti and T. spongicola is probably widespread and facultative because specimens from all localities and material collected in 1965 (Bergquist & Tizard, 1967) and 1974 (AMZ4311) also contained infestations of this polychaete. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 373 Clat hria ( Thalysias) reinwardt i Vosmaer (specimen QMG303260). A, Choanosomal skeleton. B, Fibre FIG. 191. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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. MEMOIRS OF THE QUEENSLAND MUSEUM 374zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ microspined bases, respectively. Echinating acanthostyles dense, very heavily echinating 0 fibres (17%), moderate (43%), light (17%) or WET 3 very lightly echinating (23%). Microscleres: 8 PREDRY 26 Modified contort forms of palmate isochelae DRY 23 7 present in most specimens (although not prePREDRY 15 5 viously recorded in this species), with proportion of twisted larger isochelae ranging from 0% of Clat hr ia ( Thalysias) r einw ar dt i Vosmaer. FIG. 192. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Seasonal production of incubated larvae in the NT. spicules (46% of specimens), 1-4% (39%), 510% (13%), up to 20% (2%); in smaller isochelae proportion of twisted forms 0% of spicules (13% Variation. Shape: characteristic, consistent al- of specimens), 1-4% (29%), 5-10% (34%), up to though some variability in number of branch 20% (24%); toxas varied in abundance from very bifurcations, thickness of branches, degree of common in specimens (30% of specimens), comlateral flattening; 3 morphs recognised: thin and mon (36%), uncommon (26%) or rare (8%). evenly cylindrical; laterally flattened; or thickly cylindrical with knobbed and uneven surface; no Variability in spicule dimensions: Although some correlation found between variability in skeletal variability recorded in mean spicule dimensions architecture or spiculation and growth form; see for samples from different localities these were Kelly Borges & Bergquist (1988) for further not statistically significant for any spicule details on variability in growth form and coloura- categories (P > 0.05); similarly spicule dimention. Colour: relatively consistent range from sions were relatively homogeneous for samples grey, orange-brown to red-brown; density of pig- collected during different seasons, although only mentation may be related to exposure and water a small sample size was taken during the wet clarity where deeper and more turbid water season. populations are generally less heavily pigmented and shallow populations are brighter coloured REMARKS. Vosmaer (1880) erected C. Skeletal structure: Ectosomal skeleton typically (Thalysias) reinwardti for a specimen from the dense, continuous, discrete spicule brushes, oc- Moluccas incorrectly identified as Spongia cancasionally thin, paratangential ectosomal crust. nabina Esper, but his original diagnosis was inSubectosomal skeleton cavernous with long correct. Ridley (1884a) subsequently described plumose, non-echinated spicule tracts (51%) or two specimens (as var. subcylindrica) from Torwith choanosomal fibres close to surface (49%). res Strait where they were reportedly abundant. Choanosomal skeleton typically irregularly Vosmaer (1935a) redescribed the spiculation of reticulate, heavy and compact, with fully cored the holotype which agreed closely with Ridley's fibres forming rectangular meshes; but 4% of diagnosis, and hence emended the definition of specimens with curved fibres forming oval-elip- this species. From present data and published tical meshes; 2% with cavernous skeletons results of Kelly-Borges & Bergquist (1988) it is throughout and very few thin fibres and spicule shown that C. (T) reinwardti is a dominant tracts; 4% regularly reticulate with ladder-like species of the intertidal and shallow subtidal fibre anastomoses. Spicule skeleton lying outside fringing reef communities throughout the tropical fibres dense (48%) with abundant loose spicules Indo-west Pacific, particularly prevalent in more strewn throughout mesohyl, moderate (32%), or turbid waters. very light (20%) with few extra-fibre spicules. Aside from Kelly-Borges & Bergquist (1988) Mesohyl matrix typically light (47% of the species has been described as lacking any specimens), moderate (25%), heavy but only ectosomal specialisation (viz. Clathria condilightly pigmented (18%), or heavy, dark brown tion), whereas careful histological sectioning pigmented (10%). Megascleres: Subectosomal shows that it has a classical ectosomal skeleton of auxiliary subtylostyles typically with two distinct size categories of auxiliary microspined bases although most specimens had megascleres, localised in the ectosomal and subat least some smooth ones (0-4% of megascleres ectosomal regions respectively (viz. Thalysias were entirely smooth (in 4% of specimens), 5- condition). Bergquist & Tizard (1967) suggested 10% (15%), 11-20% (34%), 21-30% (15%), 31- that toxas were not previously recorded because 40% (15%), up to 64% (17%)). Choanosomal they are very slender and tend to be associated principal styles and smaller auxiliary ectosomal with larvae. However, all specimens examined in subtylostyles invariably with smooth and this study contained toxas, irrespective of SEASON TOTAL SAMPLES SAMPLES WITH LARVAE REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 375 very shallow subectosomal drainage canals radiating from large pores. Irregular and corrugated surface features were observed only in a small proportion of live samples, although 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 morphology 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. reinwardti var. pahnata)). Clathria reinwardti can be differentiated from all these species by its characteristic acanthostyle morphology, growth form, size and geometry of toxas, and ectosomal-subectosomal features. The species also differs from C. (T.) procera and C. (T.) FIG. 193. Clathria (Thalysias) ridleyi (Lindgren) (fragment of holotype spiculosa with similar growth BMNH1929.11.26.20). Section through peripheral skeleton. form in choanosomal architecture, spicule geometry and fibre characteristics. locality, seasonality or possession of larvae. Toxas are typically most abundant in the mesohyl matrix, occurring as both toxodragmata and as single spicules; they are probably also characC lath ria (Thalysias) ridleyi (Lindgren, 1897) (Fig. 193) teristic of the adult sponge. Bergquist et al. (1971) 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, p1.17, fig.8, were predominantly pale orange-brown, with p1.18, fig.14, p1.19, fig.17; Hallmann, 1912:187; smooth, turgid, membraneous surface and only Van Soest, 1984b:115. MEMOIRS OF THE QUEENSLAND MUSEUM 376zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Tenacia ridleyi; Levi, 196lb:522-524, text-fig.14. Clathria ridleyi; Hooper & Wiedenmayer, 1994: 273 cf. Clathria ramosa; Vosmaer, 1935a:611, 642, 669. MATERIAL. HOLOTYPE: NHRM (fragment BMNH1929.11.26.20): near Membalong, SW. of Belitung I., Java Sea, Indonesia, 3°09'S, 107°38'E, coll. C. Aurivillius (beach debris). HABITAT DISTRIBUTION. Depth 10-14 m; on dead or live coral substrate; Torres Strait (FNQ) (Ridley, 1884a); also Java Sea (Lindgren, 1897), and Taganak I., Philippines (Levi, 1961b). DESCRIPTION. Shape. Ramose, with thin cylindrical bifurcating and anastomosing branches, short stalk. Colour Red alive. Oscules. Unknown. Texture and surface characteristics. Harsh; surface 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 subectosomal auxiliary subtylostyles. Choanosome. Choanosomal skeleton irregularly reticulate, with light spongin fibres forming rectangular meshes and with both primary and secondary components; primary vaguely ascending fibres have multispicular core of 8-10 rows of choanosomal principal styles, whereas secondary elements less heavily cored; acanthostyles dispersed evenly over fibres. Megascleres. Choanosomal principal styles slightly curved, with rounded, smooth non-tylote bases. Length 150-300pm, width 8-15Rm. Subectosomal auxiliary subtylostyles straight, usually with microspined bases. Length up to 3001,im, width up to 12.5p.m. Ectosomal auxiliary subtylostyles are identical in morphology to subectosomal spicules. Length up to 120p,m, width up to 4pm. Acanthostyles subtylote, with blunt or rounded, profusely microspined points, with aspinose 'necks' proximal to base. Length 6875p,m, width 5-9.5p,m. Microscleres. Palmate isochelae unmodified, single size category. Length 9-18Rm. Toxas not recorded in Ridley's or Lindgren's material, but described as raphidiform by Levi, with slight central curvature and no apical flexion. Length 80-110p.m. REMARKS. Lindgren (1897, 1898) erected this species for Ridley's (1884a) unnamed specimen from Torres Strait, differentiating it from other ramose Clathria (Thalysias) in skeletal architecture, fibre characteristics, and Ectyoplasia-like acanthostyles (with recurved spines on the apex of spicules). However, the species is barely 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. Acanthostyles with apical spines have also been recorded for C. (T.) mutabilis and C. (T.) topsenti, and it is possible that this species is related to, or synonymous with one of these. Levi (1961b) recorded C. (T.) ridleyi from the Philippines, noting some similarities in growth form and surface 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, 1902a:212. Clathria rubra; Hooper & Wiedenmayer, 1994: 272. Thalassodendron paucispina Lendenfeld, 1888:224225; Whitelegge, 1901:86-87. Rhaphidophlus paucispinus; Hallmann, 1912:176188, 195, 202, 203, 300, p1.25, figs 1-2, p1.26, fig.1, text-fig.36; Guiler, 1950:8. Tenacia paucispina; Hallmann, 1920:770. Not Tenacia paucispina; Burton, 1934a:559. Thalassoendron rubens var. dura, in part; Lendenfeld, 1888:224; Whitelegge, 1901:87. Thalassodendron rubens var. lamella, in part; Lendenfeld, 1888:224, p1.7; Whitelegge, 1901:87. Not Thalassodendron rubens Lendenfeld, 1888:223. Clathria multipora Whitelegge, 1907:496, p1.45, cf. Microciona prolifera; Vosmaer, 1935a:637, 610, 670. MATERIAL. HOLOTYPE: AMG9048 (presently missing): Port Jackson, NSW, 33°51'S, 151°16'E. PAR ATYPE of E. rubra: AMG9049: unknown locality (label 'Clathria pumila var. rubra Lend, type ?'). LECTOTYPE of T. paucispina: AMG9121a: Port Jackson, NSW, 33°51'S, 151°16'E (dry, label `Thalysias paucispinus; type'). PARALECTOTYPES of T. paucispina: AMG9121b: same locality (dry, label 'cotype'). BMNH1887.1.27.1, 1954.2.10.71, 1954.2.12.54 (fragments AMG3557): same locality. AMZ961 (dry; presently missing): same locality. LECTOTYPE of T. rubens var. lamella: AMZ459: Port Jackson, NSW, 33°51'S, 151 0 16'E (wet). PARALECTOTYPES of T. rubens var. lamella: AMZ461: same locality (wet, fragment from figured specimen). BMNH1887.4.27.124 (1954.2.10.70): same locality (dry). BMNH1887.1.24.28 (fragment ZMB1147): same locality (wet). HOLOTYPE of T. rubens var. dura: AMG9123: same locality (dry). PARATYPES REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 377 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. 378zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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 Thaphidophlus paucispinus, var. multiporus'). PARALECTOTYPE of C. multipora: AMZ723: same locality. OTHER MATERIAL: NSW- AMZ458, AMZ117, AMZ4809, 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-lamellate 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 flabellatedigitate 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, compressible, flexible when preserved; surface smooth, with or without small conules, often with well developed subectosomal ridges and oscular areas. Ectosome and subectosome. Surface with disctinct 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; subectosomal 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 (200-550iim diameter), imperfectly differentiated into primary (pauci- or multispicular) fibres (up to 1601.1.m diameter) and secondary (uni- or paucispicular) fibres (up to 110Rm diameter); fibres substantially heavier and more regularly reticulate at core, more radial and plumo-reticulate towards periphery; peripheral fibres terminate 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 pigmented, containing few loose subectosomal and choanosomal megascleres dispersed between fibres; echinating acanthostyles sparse, confined mostly to larger fibres; choanocyte chambers oval, 50-80tLm diameter. Megascleres. Choanosomal principal styles relatively 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)-295p.m, width 9-( Subectosomal auxiliary subtylostyles long, slender, straight, with slightly subtylote, smooth or microspined bases, and fusiform points. Length 202-(227.1)-2811im, width 5-(6.6)-8Rm. Ectosomal auxiliary subtylostyles short, slender, straight or slightly curved near basal end, with subtylote spined bases and slightly hastate points. Length 105-(125.4)-154p.,m, width 3(4.4)-5Rm. Acanthostyles subtylote, tapering cylindrical, fusi form, with evenly distributed spines or fewer spines at 'neck' proximal to base, spines small, straight, erect. Length 75484.3)-98pm, width 4(6.8)-4,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)-24Rm. Toxas u-shaped, thick, with only slight central curvature and tapering, non-reflexed points. Length 45-(54.7)-84,m, width 1.5-(2.1)-3p.m. REMARKS. This species is very similar to C. (T) cactifonnis 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) cactifonnis), and the presence of plumose tufts of choanosomal principal styles, projecting through and echinating peripheral spongin fibres in C. (T) ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 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. MEMOIRS OF THE QUEENSLAND MUSEUM 380zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 distinguishing C. (T) rubra from other species is possession of small, peculiar u-shaped (oxea-like) toxas, 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) cactifortnis and C. (T) hirsuta). Judging from the number of specimens of this species collected by the early trawling expeditions (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) cf. rubra (Lendenfeld, 1888) (Figs 196) cf. Echinonema rubra Lendenfeld, 1888:221. Tenacia paucispina; Burton, 1934a:559. MATERIAL. SPECIMEN: GREAT BARRIER REEF, QUEENSLAND - BMNH1930.8.13.108: Penguin Channel, off Alexander Bay, 16°15'S, 145°31'E, 20-31m depth, 24.ii.1929, 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, compressible; 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 subtylostyles and long principal subtylostyles protruding from peripheral fibres supporting ectosomal skeleton; principal subtylostyles also occasionally protruding through surface; terminal spongin 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 (15035011m diameter), more cavernous in periphery than at core; spongin fibres imperfectly divided into primary, mostly ascending, multispicular fibres (60-801Jm diameter) and secondary, mostly transverse, paucispicular fibres (25-60pin diameter); echinating acanthostyles relatively sparse in deeper choanosome, more-or-less concentrated on exterior side of fibres and at fibre nodes; mesohyl matrix heavy, granular, with numerous microscleres and auxiliary spicules scattered throughout mesohyl between fibres; choanocyte chambers oval, up to 80pLm diameter. Megascleres. Choanosomal principal subtylostyles 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)-346p,m, width 5-(7.7)-10p.m. Subectosomal auxiliary subtylostyles straight, slender, subtylote microspined or smooth bases, fusiform points. Length 219-(262.0)-358p,m, width 3-(4.5)-6p.m. Ectosomal auxiliary subtylostyles very slender, straight, subtylote smooth or microspined bases, fusiform points. Length 121(152.8)-181p.m, width 2-(3.3)-4p.m. Acanthostyles long, slender, subtylote, evenly spined except for partially aspinose area at 'neck' proximal to base, spines small, recurved. Length 73-(82.3)-931..m, width 4-(4.8)-7p.m. Microscleres. Palmate isochelae large, unmodified. Length 15-(16.8)-19p,m. Toxas intermediate between wing-shaped and u-shaped, relatively thick, with gently rounded central curvature and reflexed points. Length 18(43.6)-106p,m, width 0.8-(1.8)4.0p.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 (1934a) indicated that it was most similar to Clathria multipora Whitelegge, subsequently demoted to a variety (or subspecies) of C. (T) rubra by Hallmann (1912), but comparison 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 REVISION OF MICROCIONIDAE 381 EzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA D_ 0 LO FIG. 196. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Thalysias) cf. rubra (Lendenfeld) (BMNH1930.8.13.108). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles. D, E,chinating acanthostyle. E, Palmate isochela. F, Wing-shaped toxas. G, Section through peripheral skeleton. H, Australian distribution. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 382zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA specimen consisted of a 'tangled, anastomosing mass of angular ... nodulose branches, each 35mm 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, p1.17, fig.7, p1.19, fig.18. Rhaphidophlus spinifer, Thiele, 1903a:958, p1.28, fig. 23; Hallmann, 1912:177. Clathria spinifera; Hooper & Wiedenmayer, 1994: 274. Not Clathria spinifera Sara, 1978:67-70, text-figs 4143; Desqueyroux-Faundez & Moyano, 1987:50 . cf. Microciona prohfera; Vosmaer, 1935a:611. MATERIAL. LECTOTYPE: ZMUU (not seen) (fragments BMNHI929.11.26.6, 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- NTMZI750.(fragment QMG300493). HABITAT DISTRIBUTION. Rocky reef and associated 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, 5-12mm 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 coppergreen 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 surface, 0.5-2mm diameter. Texture and surface characteristics. Basal stalk firm, almost rigid, branches firm, compressible, highly flexible; surface minutely rugose with distinct, shiny surface crust with numerous irregularly distributed microconules, dissected by minute ridges and canals. Ectosome and subectosome. Moderately well developed ectosomal skeleton, with more-or-less continuous palisade of discrete spicule brushes composed of smaller ectosomal auxiliary subtylostyles; 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 although peripheral skeleton predominantly plumose; spongin fibres moderately heavy, 58llOwn diameter (heavier in Indonesian specimen); fibres form oval to elongate meshes, 95-460u,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 protruding principal styles) forming characteristic stellate-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 subectosomal 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 383 FIG. 197. Clathria (Thalysias) spinifera (Lindgren) (SMF1815). A, Choanosomal principal style. B, Subectosomal 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. MEMOIRS OF THE QUEENSLAND MUSEUM 384zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 40. Comparison between spicule dimensions for type and other material of Clathria (Thalysias) spinifera (Lindgren). Measurements in p,m, and cited as range (and mean) (N=25). SPICULE Fragment of holotype (BMNH 1929.11.26.6) 153-(196.9)Choanosomal 242 x 8-(11.4)styles 14 Specimen NTMZ1750 (N=1) (Thiele, (NW Australia) 1903a) (Indonesia) 183-(206.0)242 x 12(16.1)-18 168-(258.2)S ubectosoma I 2034233.8)styles 256 x 5-(6.8)-8 291 x 5-(6.9)-9 Ectosomal styles 97-(132.0)171 x 3-(3.8)-5 Toxas 172-(196.4)210 x 446.4)8.5 92-(115.5)96-(132.2)151 x 3-(4.9)-6 158 x 2-(4.5)-6 69-(72.3)-77 x 7-(9.1)-11 102-(108.6)114 x 5-(8.6)-9 14-(14.7)-16 13-(15.2)-17 11-(13.9)-16 26479.7)-124 x 0.5-(0.9)-1.5 50-(149.0)194 x 0.8(1.2)-1.5 32484.2)-119 x 0.5-(0.8)-1.2 Acanthostyles 75-(84.8)-93 x 6-(7.6)-9 Chelae 155-(187.2)200x 8-(I6.2)20 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, unmodified, 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 curvature, 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 fibres are lighter; echinating acanthostyles are heavier on fibres; isochelae are poorly silicified, slightly sigmoid and have ill-defined alae; acanthostyles are slightly longer; principal styles are shorter and up to 50% have spined bases (Table 40). Conversely, skeletal structure, fibre characteristics, the distribution of spicules throughout fibres and spicule geometries are virtually identical. Clathria (Thalysias) spinifera has unusual fibre characteristics with only bases of principal styles enclosed in primary spongin fibres (forming multispicular 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 composed of choanosomal principal megascleres. In the spicata' group those tracts occur exclusively outside fibres, and usually ascend to the ectosomal 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, 1878:163 (nomen nudum). Not Echinonema typicum Carter, 1881a:362. Unidentified sponge-algae associate, 'possibly undescribed'; Scott et al., 1984:291-293. MATERIAL. HOLOTYPE: WAM649-81(1) (fragment NTMZ1729): Goss Passage, off Beacon Is, Wallabi 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 Abrolhos, 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 MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 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. I-J, Palmate and modified isochelae. 386zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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 to 5mm maximum thickness, with rounded or uneven, bifurcate margins. Colour Live colouration unknown, yellowbrown in ethanol. Oscules. Sparse, scattered over external surface, up to 2mm diameter; exhalant pores minute, dispersed over entire surface, giving ectosome slightly reticulate appearance. Texture and sutface characteristics. Texture rubbery, compressible; surface optically smooth, slightly uneven, with subdermal striations and grooves visible only near margins of lamellae. Ectosome and subectosome. Thin, disorganised ectosomal crust composed of acanthostyles erect on peripheral fibres, intermingled with paratangential or erect plumose brushes of auxiliary styles of 2 sizes (larger ones less common than smaller ones), together forming nearly continuous palisade of erect spicule brushes on surface; subectosomal skeleton absent entirely; choanosomal skeleton immediately subdermal. Choanosome. Skeleton structure irregularly reticulate, with very thick 'fibres' formed almost exclusively by Codiophyllum algal filaments, 701561.1.m diameter, with only a superficial layer of spongin covering surface of algal filaments; coring spicules excluded entirely from within 'fibres', although some auxiliary subtylostyles lie on surface of 'fibre' and many echinating acanthostyles embedded within surface and erect on 'fibre'; 'fibre' (algal filament) meshes usually form large nodes, and in peripheral skeleton nodes usually have tangential layer of subectosomal 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 small oval choanocyte chambers, 49-86Rm diameter. Megascleres. Choanosomal principal styles absent. Subectosomal auxiliary subtylostyles relatively uncommon, straight, relatively thick, fusiform, with slightly constricted, smooth bases. Length 211-(253.8)-292Rm. width 3-(5.2)-8Rm. Ectosomal auxiliary subtylostyles most common, straight or slightly curved at centre, thick, fusiform, with smooth subtylote bases. Length 92-(128.5)-1481J.m, width 446.2)-7.511m. Acanthostyles extremely abundant, short, thick, with slightly swollen subtylote bases, evenly spined except for aspinose points; some modified to acanthostrongyles; spines large, slightly recurved at point. Length 48456.4)63p,m, width 3.5-(7.2)-9Rm. Microscleres. Isochelae abundant, palmate, unmodified, 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)-16p.m. Toxas wing-shaped, short, relatively thick, slight angular central curves, slightly reflexed arms. Length 8443.5)-9611m, width 1.041.8)2.51.tm. Associates. Probable obligatory symbiotic relationship with red algae (Halymeniaceae), possibly Codiophyllum (identified from a superficial comparison with published descriptions and figures in Scott et al., 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 (Antho opuntioides (Lamarck) and A. frondifera (Lam.); Topsent, 1929). ETYMOLOGY. Greek protheco- from Topsent's (1929) term 'styloprothese' referring to the incorporation of algal filaments into the skeleton, displacing spongin fibres. REMARKS. This species is similar to C. (T.) cactifonnis in acanthostyle 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.) cactiformis). Carter's material has not been found in BMNH collections, but his description mentions the sponge-algae relationship as being a `pseudomorph' of the free living sponge. Carter's (1878) original report of Echinonema typicum is not valid (nomen nudum), and his subsequent description of the species (Carter, 1881a) is based on different material from his 1878 notice of the species. Hence the name typicum' is not available for this species. The external morphology of Clathria (Thalysias) styloprothesis appears to be identical to the red algae Codiophyllum flabellifornze (Sonder), redescribed and figured in detail by Scott et al. (1984) from the WA coast (lat. 28 0 35°S, 5-21m depth), particularly in the thickness REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 387 0 FIG. 199. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Thalysias) st yloprot hesis 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 PIB0004-345 (fragment QMG300043). J, Cross-section through algal lamella (pale area) showing spicules and collagen (darker areas). ^ MEMOIRS OF THE QUEENSLAND MUSEUM 388zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 200. Clathria (Thalysias) styloprothesis sp.nov. (paratype PIB0004-345 (QMG300043)). A, Algal filaments 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 389 ,f fe/ ' rrrrr;'rr7rZ TZ zyxwvutsrqponmlkjihgfedcbaZYXWVU (r ‘‘e 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 lotrochota baculifera. 390zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM and cortication of algal filaments. Scott et al. (1984) consider that this relationship involves an algae which has been invaded by a sponge, in which case C. (T.) styloprothesis is merely a thinly encrusting, excavating and opportunistic sponge species. By comparison, Topsent (1929) suggested that, in a similar case, the algal filaments are incorporated into the sponge ('styloprothese'). In his material the algal filaments were much thinner and acanthostyles varied from forms with rounded points (strongyloids), 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.) cactifortnis in the geometry and dimensions of its spicules, particularly microscleres, as well as incorporating algal filaments into fibres. In this latter respect it is easily differentiated from other rnicrocionids, although this statement is based on the assumption that the algal-sponge relationship is species specific, and that flabellodigitate sponges (or algae) with short squat acanthostyles, similar in geometry to those found in C. (T.) cactifonnis, are conspecific. Clathria (Thalysias) tingens sp. nov. (Figs 201-203, Plate 8C-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.iii.1985. NTMZ2530: Orontes Reef, mouth of Port Essington, Cobourg Peninsula, NT, 11°03.6'S, 132°05.4'E, 1 1m depth, 17.ix.1985, coll. J.N.A. Hooper (SCUBA). OTHER MATERIAL: WAQMG301154. NT- QMG300141 (fragment NTMZ2I 11). QLD - QMG303826, QM unreg. HABITAT DISTRIBUTION. Intertidal laterite rock, dead coral reef flats, in rock pools, encrusting on underside of dead faviid coral boulders, and on coral reef slopes in deeper waters, sheltered on the sides of faviid coral boulders or Acropora thickets; 0-32m depth; known only from Australia: Darwin Harbour, Port Essington, Cobourg Peninsula (NT); Hibernia Reef, Sahul Shelf (WA); Raine I. (FNQ), Hook Reef (MEQ) (Fig. 201H). DESCRIPTION. Shape. Thickly encrusting, 530mm thick, gelatinous lobate, following contours of substrate with prominent surface folding and ridge-like sculpturing. Colour Ectosome pale pink and white mottle to pale red-orange alive (Munsell 5RP 8/4-2.5R 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 live or preserved specimens. Texture and surface characteristics. Compressible, gelatinous, slimy in situ, producing abundant clear mucous upon exposure to air; surface optically smooth, lobate, with occasional folds and minute subectosomal canals and ridges, although striations not prominent; mottled or speckled external appearance superficially resembles a compound ascidian. Ectosome and subectosome. Minutely hispid, light, poorly differentiated series of ectosomal 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 below peripheral skeleton but development variable, 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 both sizes of auxiliary spicules; subectosomal skeleton with larger auxiliary subtylostyles forming extensive, plumose, and discrete paratangential tracts originating close to substrate, ultimately diverging and piercing ectosomal skeleton; individual subectosomal auxiliary subtylostyles also scattered throughout mesohyl; mesohyl 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. Choanosome. Skeletal architecture hymedesmoid, with spongin fibres reduced to a basal layer, up to 3011m thick, lying on coralline substratum, with bases of choanosomal principal styles and acanthostyles embedded and perpendicular to substrate; no folding of basal spongin or fibre nodes observed; choanosomal skeleton occupies only small percentage of sponge thickness with major portion being extensive plumose subectosomal and ectosomal skeletons in periphery of sections; mesohyl substantially lighter in deeper choanosomal skeleton, closer to substrate, than in peripheral skeleton; choanosomal chambers 52-65iim diameter. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 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. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 392zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 203. Clathria (Thalysias) tin gens 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 393 Megascleres. Choanosomal principal megascleres long, thick subtylostyles, with distinctive curvature towards basal end, prominent tylote bases abundantly microspined, tapering towards fusiform point. Length 198.54374.3)524.0p.m, width 8-(12.9)-19.1p,m. 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 259.34377.4)-502.5 p.m , width 4 .547 . 2 )10.5p.m. Ectosomal auxiliary subtylostyles short, slender, straight, fusiform, with less prominent tylote bases, usually microspined occasionally smooth. Length 117.9-(170.0)-208.6p,m, width 2.3-(3.8)-5.6p,m. Acanthostyles long, slender, straight, subtylote, fusiforrn pointed, evenly spined, spines small, sharp, slightly recurved. Length 79.9(108.3)-150.9p,m, width 4.6-(8.0)-12.6p,m. Microscleres. Palmate isochelae abundant, unmodified, with lateral and front alae approximately smae length, front ala completely detached along lateral margin, lateral alae fused to shaft. Length 9.7-(15.3)-20.7 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)-3.2p.m. Associations. Growing next to, or covering other encrusting sponges (lotrochota, Placospongia, Ulosa, Mycale), compound and simple ascidians, and coralline algae. ETYMOLOGY.Latin tin gens, refering to its superficially tinted pink live colouration. REMARKS. Clathria (Thalysias) tingens is similar to C. (T.) toxifera in 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 (possession of accolada toxas). In addition, live colour, differential pigmentation between ectosomal 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. (Wilsonella) tuberosa, and yellow morphs of C. (T.) abietina, both from NW Australia, and the yellow C. (T.) venosa from the West Indies. Clathria (Thalysias)tingens should also be compared with C. (Microciona)hymedesmioides Van Soest from Curacao in secondary colouration, hymedesmoid skeletal architecture (seen in thinly encrusting portions of C. (T.) tingens), and similarities in the morphology of some of their spicule categories. De Laubenfels (1954:135) recorded a specimen of C. (T.) cervicomis from the Marshall Is, which also had differentiated ectosomal (pale orangebrown) 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 8E) Hymeraphia toxifera Hentschel, 1912:382-383, p1.20, fig.40. Microciona toxifera; Burton, 1938a:31, p1.5, fig.30; Vacelet & Vasseur, 1977:116. Clathria toxifera; Hooper & Wiedenmayer, 1994: 274. Not Stylostichon toxiferum Topsent, 1913a:621. MATERIAL. HOLOTYPE: SMF967T: Bei Mimien, Aru I., Arafura Sea, Indonesia, 6°S, 134°50'E, 15m depth, 8.iv.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 Z3681. HABITAT DISTRIBUTION. Intertidal laterite rock, dead coral reef flats, in rock pools, on sublittoral faviid coral heads, usually exposed at ELWS tides; usually encrusting 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); Hibernia Reef, Sahul Shelf (WA) (Fig. 204H); also Aru Is, Indonesia (Hentschel, 1912), Ko Samui, Gulf of Thailand (present study), Madras (Burton, 1938a). DESCRIPTION. Shape. Thinly encrusting, 1- 5mm thick, often covering substantial areas of subtrata. Colour. Pale orange-brown to darker red-brown alive (Munsell 5YR 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. 394zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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, 150-450Rm diameter, scattered over surface, raised slightly above surface (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 substrate, 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, composed of 2 layers: outer layer with smaller ectosomal 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 surface brushes, up to 300p.,rn in thin sections, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 395 whereas in thicker sections principal megascleres barely pierce surface, surrounded at their points by ectosomal spicule brushes in classical Thalysias architecture; development of ectosomal skeleton variable, ranging from continuous dense palisade of spicule brushes in thicker growths, to sparse, irregularly paratangential, discrete brushes in thinner sections; foreign debris sometimes incorporated into ectosomal skeleton with particles surrounded by spicule brushes. In subectosomal region are also thin longitudinal bands, forming dense tracts, composed of subectosomal auxiliary subtylostyles 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 originate approximately halfway along length of principal spicules whereas in thicker sections subectosomal brushes do not diverge until peripheral skeleton; subectosomal region 0.5-3mm thick containing abundant, moderately heavily pigmented. Choanosome. Choanosomal skeletal hymedesmoid in thinner sections with single megascleres embedded in basal spongin lying flat on substratum, or microcionid in thicker regions with basal spongin slightly raised nodes (='fibres'); peripheral skeletal architecture distinctly plumose; choanosomal principal megascleres and echinating acanthostyles perpendicular to substrate with bases embedded in basal spongin or in erect fibre nodes where present; basal spongin moderately heavy, yellow-brown, lying directly on calcareous substrate, 8-20p,m thick in hymedesmoid sections, up to 451J,m in microcionid sections; few choanocyte chambers observed only in thicker sections, 13-55p,m diameter, usually lined by toxas and/or isochelae. Megascleres. Choanosomal principal subtylostyles 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)-685.1p,m, width 5.1 (14.1)-25.5p,m. Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, with distinctly swollen tylote bases, usually lightly microspined, less frequently smooth ((0-10% of spicules in individual specimens). Length 228.2-(354.3)-494.3p,m, width 2.0-(6.5)-12.9p.m. Ectosomal auxiliary subtylostyles short, straight, fusiform, less markedly tylote than larger auxiliary subtylostyles, with smooth or basal spines. Length 123.3-(171.1)-229.6p,m, width 1.5-(3.6)-6.6p.m. Acanthostyles thick, slightly curved towards basal end, with subtylote bases, long tapering points, mostly evenly, lightly spined except for bare 'neck' proximal to base, spines large, recurved, sharply pointed. Length 121.94154.5)208.0p,m, width 3.0-(7.9)-14.0p,m. Microscleres. Palmate isochelae usually abundant (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-(22.6)-30p,m. Toxas verging on oxhorn, very abundant, relatively thick but variable in length, with wide central curvature and straight or only slightly reflexed points. Length 16-(89.8)-241[1,m, width 0.8-(2.9)-5p,m. Associations. Growing over or in proximity to other encrusting sponges (Desmanthus, Mycale, Ulosa, Haliclona, Petrosia, and other microcionids), compound ascidians, coralline algae, Platygyra and faviid corals, barnacles (Chthamalus), and metal debris (aluminium cans) on the reef flat (with some evidence of etching on the metal substrate). It is probable that the species plays an active role in coral bioerosion. REMARKS. This species is perfectly recognisable from Hentschel's (1912) original description and material described here from Northern Territory, Western Australia and Thai waters do not differ markedly from the Indonesian population. Conversely, we do not know any details of Burton's (1938a) specimen from Madras since he merely repeated verbatim the original description, 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 MEMOIRS OF THE QUEENSLAND MUSEUM 396zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 397 of the mineral skeleton, particularly spicule geometries. The possession of stellate subectosomal surface sculpturing (i.e., subectosomal drainage canals radiating towards oscules) is known for C. (T.) 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 taxonomic importance given that it represents an ecological adaptation: viz, in thinly encrusting sponges subectosomal canals represent the primary means of water circulation whereas in thicker specimens the primary aquiferous system is predominantly internal. In any case C. (T.) toxifera differs from the Caribbean species in spicule geometry and ornamentation 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 include numerous species from the Arafura Sea: C. (T.) aruensis (Hentschel, 1912:381), C. (T.) calochela (Hentschel, 1912), C. (T.) distincta (Thiele, 1903a), C. (T.) longitoxa (Hentschel, 1912), C. (Microciona) rhopalophora (Hentschel, 1912), C. (M.) hentscheli nom. nov., C. (M.) simills (Thiele, 1903a), C. (M.) tetrastyla (Hentschel, 1912) and C. (M.) thielei (Hentschel, 1912) (all of which lack isochelae). Species from other localities are: C. (T.) michaelseni (Hentschel, 1911) from Shark Bay, WA (with sigmoid anchorate-like (bidentate) isochelae); C. (M.) aceratoobtusa (Carter, 1886g) from the Mergui Archipelago, Burma, and from Shark Bay, WA (Hentschel, 1911) (with smooth echinating megascleres); C. (M.) affinis (Carter, 1880a), C. (M.) bulboretorta (Carter, 1880a), C. (M.) fascispiculifera (Carter, 1880a) (with sigmoid palmate isochelae), and C. (M.) quadriradiata (Carter, 1880a), all from the Gulf of Manaar, Sri Lanka; C. (T.) dubia (Kirkpatrick, 1900a) from Christmas I., 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 in a forthcoming monograph on the Indo-Malay microcionids (Hooper et al., in prep.). From re-examination of the relevant type material of each of these species, 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 8F) Spongia vulpina Lamarck, 1814:449; Lamarck, 1814:376 Rhaphidophlus vulpitza; Ridley, 1884a:615. Rhaphidophlus vulpinus; Topsent, 1932:110, p1.5, fig.3. Clathria vulpina; Hooper & Levi, 1993a:1246-1250, figs 11-12, table 6; Hooper & Wiedenmayer, 1994: 274. Halichondria frondifera Bowerbank, 1875:288-289 Amphilectus frondifera; Vosmaer, 1880:115. Clathria frondifera; Ridley, 1884a:448-449, 612, p1.42, fig.i, p1.53, fig.j; Ridley & Dendy, 1887:149, 160, 178, 246, 254; Topsent, 1892b:23, p1.1, fig.4; Lindgren, 1897:480, 483; Lindgren, 1898:309-310; Dendy, 1916a:128; Dendy, 1922:65; Hentschel, 1912:360-361; Row, 1911:382, 389, 396; Burton, 1938a:27-28, p1.3, fig.21; Burton, 1959:243; Levi, 1961c:21-22; Thomas, 1970b:206-207, text-fig.11; Thomas, 1973:33-34, p1.2, fig.6, p1.8, fig.4; Tanaka et al., 1976:801-805; Tanaka et al., 1977:767-772; Tanaka et al., 1978:1283; Thomas, 1979a:26-27, p1.2, fig.!; Hoshino, 1981:161; Liaaen-Jensen et al., 1982:171. Clathria frondzfera var. seto-tubulosa; Wilson, 1925:439. Clathria frondifera var. dichela; Hentschel, 1912:360361. Tenacia frondifera var. dichela; Hallmann, 1920:771. Tenacia frondifera; Burton & Rao, 1932:337-339; Burton, 1934a:559; Levi, 196lb:521-522, text-figs 12,13. Rhaphidophlus frondifera; Thiele, 1903a:958, textfig.23. Thalysias frond ifera; de Laubenfels, 1954:138-139, text-fig.88. Clathria dichela; Vacelet et al., 1976:71, p1.3, fig.b; Vacelet & Vasseur, 1977:114. Rhaphidophlus dichela; Van Soest, 198413:115. Clathria corallitincta Dendy, 1889b:85, p1.4, fig.8; 1916b:128; Dendy, 1922:65. Rhaphidophlus seriatus Thiele, 1899:14, p1.1, fig.6, p1.5, fig.7. Clathria reinwardti var. palmata; Ridley, 1884a:447; Bergquist & Tizard, 1967:186. ? Clathria typica; Vacelet & Vasseur, 1971:94. Not Rhaphidophlus filifer var. spitufera; Lindgren, 1898:311-312, p1.17, fig.7, p1.19, fig.18; Thiele, 1903a:958. Not Clathria nuda; Hentschel, 1912:298, 359, 364365, p1.19, fig.28. Not Clathria frondifera var. major; Hentschel, 1912:361. cf. Microciona prolifera; Vosmaer, 1935a:609, 629. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 398zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA A 0 zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPON CD 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 I., Indonesia, 3°10'S, 107°15'E. HOLOTYPE and PARATYPE of C. frondifera var. setotubulosa: USNM21256, 21257: specific locality unknown, Philippines. HOLOTYPE of T. frondifera var. dichela: SMF1673 (fragment MNHNDCL2230): Straits of Dobo, Aru I., Indonesia, 6°S, 134°50'E, I 6m depth, 20.iii.1908, coll. H. Merton (dredge). HOLOTYPE and PARATYPE of R. seriatus: NMB16, 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 I., Torres Strait, Qld, 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°50'N, 79°40'E, coll. E. Thurston (dredge). OTHER MATERIAL (Hooper & Levi, 1993a for additional list): INDONESIA - QMG303689 (NCIOCDN-1388-S), QMG303682 (NCIOCDN1252-U). PHILIPPINES - QMG300298 (NCIQ66C5715-C), QMG300310 (NCIQ66C-5716-C), QMG300321, QMG300332. PNG- QMG300368 (NCIQ66C-4446-X). VIETNAM - PIB00O5-191 (fragment QMG300047). MADAGASCAR PIBOCB12-200 (fragment QMG300056). NTQMG3 03587, QMG303323, QMG303378, NTMZ3918, NTMZ3931, QMG300760 (NCIQ66C4776-1), QMG300560 (NCIQ66C-4825-L). WA - ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 5MF1673. FI, Hentschel's (1912) Aru I. specimen SMF1699B. I, NTMZ1810, deeper water, NW Australia. J, QMG300047, shallow water, Gulf of Thailand . MEMOIRS OF THE QUEENSLAND MUSEUM 400zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 208. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Clat hria ( Thalysias) vulpina (Lamarck) (NTMZ2691). A, Choanosomal skeleton. B, Fibre characteristics (x260). C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Bases of auxiliary subtylostyles. G, Palmate isochelae. H, Accolada - wing-shaped toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 401 MONTH TOTAL SAMPLES SAMPLES WITH LARVAE JANUARY 4 0 FEBRUARY 3 0 MARCH 4 0 APRIL 9 0 MAY 9 0 JUNE 3 0 JULY 3 0 AUGUST 5 0 SEPTEMBER 9 9 OCTOBER 19 19 NOVEMBER 12 0 DECEMBER 6 0 FIG. 209. Clathria (Thalysias) vulpina (Lamarck) Incidence of larval production in NT populations. PIB0004-457 (fragment QMG300053). QLDQMG3 0 3 5 2 2, QMG300861, QMG303038, QMG304758, QMG304409, QMG304370, QMG303898. NSW- QMG301376, QMG301385, QMG301405. HABITAT DISTRIBUTION. Mostly on rock or dead coral reefs, exposed to currents or sheltered between coral heads; 0-80m depth; widely distributed throughout Indo-west Pacific; Bynoe Harbour, Darwin Harbour, Orontes reef, Melville I., Wessel Is (NT); Shark Bay, Carnarvon, Barrow I., Exmouth Gulf, Port Hedland, Monte Bello Is, Amphinome Shoals (WA); Gulf of Carpentaria, Thursday I., Bird 1., Home Is, Green I., Frankland Is, Low Is, Cook Reef, Shelburne Bay, Howick Is, Lizard I. (FNQ); Hook Reef (MEQ); Byron Bay (N. NSW) (Fig. 206H); also Mozambique (Thomas, 1979a), Madagascar (Vacelet et al., 1971, 1976, 1977; present study), Amirante Is (Ridley & Dendy, 1887), Seychelle Is (Ridley & Dendy, 1887; Thomas, 1979a), Aldabra Is (Levi, 1961c), Red Sea (Topsent, 1892b; Burton, 1959a), Kattiawar W. coast of India, Madras, Gulf of Manaar and Sri Lanka (Dendy, 1889b, 1916b; Lindgren, 1897; Burton, 1938a; Thomas, 1970b), Mergui Archipelago and Andaman Is (Burton & Rao, 1932), Straits of Malacca, Malaysia and Gaspar Straits (Bowerbank, 1875), Aru Is, Arafura Sea, Java Sea and Sulewasi, Indonesia (Thiele, 1889; Lindgren, 1898; Hentschel, 1912; present study), Hon Rai 1., Vietnam (present study), Negros Orientate, Bohol Sea, Mindinao, and S. Philippines (Wilson, 1925; Levi, 1961b; present study), Guam, Micronesia (de Laubenfels, 1954), S. Japan (Hoshino, 1981), New Caledonia (Hooper & Levi, 1993a), Madang, Papua New Guinea (present study) DESCRIPTION. Shape. Tubulo-digitate, variable in size ranging from small single digits (80mm high) to massive multiple digitate lobes 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 absent in deeper water specimens, beige brown alive (7.5YR 8/4). Oscules. Oscules scattered between surface conules, not confined to any particular region, 26mm diameter, raised slightly above surface with distinct membraneous lip; ostia 0.4-2.2mm diameter flush with surface. Texture and surface characteristics. Soft, rubbery, 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 cavernous; when intact ectosomal membrane stretched across adjacent branches, through which subectosomal canals can be seen. Ectosome and subectosome. Conspicuous palisade of tangential, paratangential and erect spicule brushes composed of smaller auxiliary subtylostyles, with choanosomal principal megascleres protruding through surface, overlaying 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 (60-135p.m diameter) and transverse connecting fibres (25-391km diameter); fibre anastomoses form rectangular to square meshes lined by oval choanocyte chambers (22-58pLm); fibres heavy, slightly heavier in axis than towards periphery; primary fibres cored by multispicular tracts of both choanosomal principal and subectosomal auxiliary megascleres, and coring spicules occupy less than 66% of primary fibre diameter; secondary fibres paucispicular, and coring spicules comprise only 50% 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 subectosomal auxiliary spicules and ultimately ectosomal 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 113.34184.4)253.0p,m, width 3.6-(11.2)-19.4p.m (holotype 156.5-(173.8)-193.3 x 10.1-(13.1)-15.9p,m). MEMOIRS OF THE QUEENSLAND MUSEUM 402zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 41. Summary of results of one-way ANOV's (model I), testing for variability in spicule lengths and Clat hr ia ( Thalysias) vulpina, between groups subdivided on the basis of locality, season and widths of zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA bathymetric distribution. SPICULE LOCALITY' DEPTH3 SEASON2 (N) F Prob. (N) F Prob. (N) F Prob. Choanosomal styles L (850) 3.78 P<0.001 (600) 0.69 P>0.05 (675) 1.49 P>0.05 W (850) 3.94 P<0.001 (600) 1.48 P>0.05 (675) 0.17 P>0.05 Subectosomal styles L (850) 1.82 P>0.05 (600) 1.06 P>0.05 (675) 0.55 P>0.05 W (850) 1.28 P>0.05 (600) 1.01 P>0.05 (675) 0.70 P>0.05 styles L (850) 1.27 P>0.05 (600) 0.20 P>0.05 (675) 0.33 P>0.05 W (850) 0.88 P>0.05 (600) 0.23 P>0.05 (675) 0.52 P>0.05 Acanthostyles L (850) 6.68 P<0.0005 (600) 1.73 P>0.05 (675) 0.69 P>0.05 W (850) 1.18 P>0.05 (600) 0.29 P>0.05 (675) 0.96 P>0.05 Chelae I L (850) 2.80 P<0.01 (600) 0.43 P>0.05 (675) 1.04 P>0.05 Chelae II L (850) 0.12 P>0.05 (600) 0.18 P>0.05 (675) 0.56 P>0.05 Toxas L (850) 2.83 P<0.01 (600) 0.36 P>0.05 (675) 0.81 P>0.05 W (850) 0.40 P>0.05 (600) 0.09 P>0.05 (675) 0.04 P>0.05 Ectosomal Number of groups: 1. 7 locality groups (Northwest Shelf, Darwin, Cobourg Peninsu a, Great Barrier Reef, Indonesia, Philippines, Guam). 2. 4 seasonal groups (wet (FMA), pre-dry (on), dry (ASO), pre-wet (NDJ), for Darwin & Cobourg Peninsula matenal only). 3.4 depth groups (0-4m, 4-10m, I 0-40m, 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(221.4)-310.9p,m, width 1.0-(4.7)-10.4p.m (holotype 148.2-(196.8)-231.8 x 3.646.1)8.8p,m). Ectosomal auxiliary subtylostyles styles straight, short, thin, subtylote, invariably microspined, fusiform pointed. Length 64.2(99.1)-161.1vm, width 1.0-(3.3)-6.9p,m (holotype 65.9-(87.8)-116.4 x 1.1-(3.4)-6.1p,m). 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 51.8473.1)94.7pm, width 1.1-(6.6)-11.6p,m (holotype 59.7(74.3)-86.6 x 4.5-(7.8)-10.7p,m). Microscleres. Palmate isochelae abundant, scattered 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 completely to shaft. Length I: 7.1-(12.4)-17.5Rm (holotype 9.1412.8)-17.111m), length II: 1.1(3.8)-6.9p,m (holotype 1.1-(3.7)-6.11km). Toxas common to uncommon, distributed singly 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)-199.5p.m, width 0.5-(1.2)-3.2p,m (holotype 16.7-(55.8)94.6 x 0.6-(1.7)-2.3p,m). Larvae and reproductive periodicity. 28% of all specimens examined (including material reported by Hooper & Levi, 1993a) contained incubated (viviparous) parenchymella larvae in various stages of development; young larvae spherical, more mature larvae oval-elongate, 170-420Rm diameter; cilia not observed (preserved material); undifferentiated forms (170-3401km diameter) contain whispy juvenile megascleres in axis, with mesohyl matrix similar in colouration to adult; more advanced larvae (230-4201km diameter) contain juvenile styles and toxas at centre, isochelae towards periphery, and distinct ectosomal layer of differentiated cells. In Northern Territory populations sexual ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 403 TABLE 42. Latitudinal gradients in spicule dimensions for populations of Clathria (Thalysias) vulpina. Measurements (in p.m) are mean lengths (L) and widths (W) I SE. SPICULE (N) Choanosomal styles L LOCATION/LATITUDE NCAL 22°S NWS 19°S GBR 16°S DAR 12°S CF I 1°S INDO 6°S PHIL 10°N MICRO 14°N (175) (275) (250) (300) (300) (175) (125) (125) 156.8±9.2 167.6±10.9 169.7±10.9 189.9±9.6 185.6±9.7 185.8±9.7 197.9±10.1 196.9±13.5 W 6.2±0.9 11.0±1.7 8.0±0.9 11.5±1.3 11.4±1.3 10.7±0.9 12.0±1.4 7.9±0.8 Acanthostyles L 51.1±2.8 73.1±4.0 69.0±2.3 74.4±3.7 72.3±3.9 73.1±3.4 77.9±3.9 53.2±2.5 W 3.5±I.0 7.I±0.9 5.5±0.9 6.6±0.8 6.6±0.8 6.7±1.5 6.6±0.8 5.9±0.4 - I1.7±0.9 10.8±0.4 12.9±0.9 I2.3±0.9 11.9±I.1 12.6±0.7 13.3±0.7 79.3±17.2 79.1±21.8 66.8±19.3 94.1±22.2 I I 1.7±18.2 96.4±18.6 104.4±26.2 129.8±23.9 0.6±0.2 1.2±0.2 1.2±0.2 1.2±0.2 1.2±0.2 I.2±0.2 1.0±0.2 0.9±0.2 Chelae L Toxas L W Location: NCAL = New Caledonia; NWS = Northwest Shelf,WA; GBR = northern Great Barrier Reef,Q1d; DAR = Darwin region,NT; CP = Colyourg Peninsula region,NT; INDO = Southeast Indonesia; PHIL = southern Philippines; MICRO =ivlarianas & Caroline Islands, Micronesia. - reproduction was distinctly seasonal, with incubated 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 observed. 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 subectosomal auxiliary spicules and choanosomal styles protruding through ectosome; 31% of specimens had tangential skeletons in some sections 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 subectosomal skeleton of larger auxiliary megascleres arising directly from ultimate choanosomal fibres; 28% had distinctly plumose tracts of subectosomal spicules, outside of fibres, supporting ectosomal skeleton; 14% had long plumose subectosomal spicule brushes and cavernous peripheral skeleton. 64% of specimens had regular choanosomal skeleton forming square (renieroid) or sometimes triangular (isodictyal) meshes at core, more irregular in periphery, whereas 36% were regularly (sub)renieroidreticulate throughout skeleton, forming cavernous 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 pigmented (41%), abundant, moderately heavily pigmented (17%), or heavily pigmented (28% of specimens). Choanosomal principal styles rare in 3% of specimens, largely replaced by subectosomal 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%), 2130% (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 10% of specimens (between 4-8% 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 bathymetfic distributions of samples (for all northwest Australian populations combined) (i.e., variation was equally consistent within and between groups). Conversely, popula- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 404zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA tions collected from various localities (Northwest Shelf, Darwin, Cobourg Peninsula, Great Barrier Reef, Indonesia, Philippines, Guam, New Caledonia) showed significant statistical differences 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 Clathriafrondifera, is a widely distributed, predominantly Indian Ocean species and a major component of the tropical macrobenthos. The cavernous, insubstantial tubulo-digitate growth form, regularly rectangular skeletal construction and deep red colouration are characteristic for the species, although spicule geometry is unremarkable. The holotype is identical to NW. Australian material, and it is probable that original material collected by Perron & Lesseur was obtained from WA (although not specified by Lamarck, 1814). Neither Ridley (1884a), Wilson (1925) nor Topsent (1932) recorded toxas in the holotype, but these spicules are definitely present, relatively abundant albiet thin. Topsent (1932) compared the species with C. (T.) clathrata (Schmidt) and suggested that although spiculation was closely comparable, the two species could be differentiated by the presence of basal spination on both categories of auxiliary megascleres and the absence of toxas in C. (T.) vulpina. These characters are shown here to be of little systematic importance and erroneous, respectively, but these two species are otherwise differentiated by their skeletal construction and growth form. Wilson (1925) also noted that C. (T.) vulpina showed considerable intraspecific variation in growth form (and size), but his statement overemphasises this apparent variation. From the numerous published records of this species from Australasian and Indo-Malay regions in particular (most appearing under the name of C. frondifera), it is apparent that this species has a number of consistent and characteristic features including its cavernous growth form, regular (sub)renieroid choanosomal skeletal structure and individual spicule geometries, whereas ectosomal development is much more variable. To illustrate this variability, of the previously published material, Wilson's (1925) variety of C. setotubulosa from the Philippines and de Laubenfels'(1954) specimen of C. frondifera from Guam have well developed Thalysias special 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 difficulty in clearly differentiating Clathria and Thalysias species based on this variability in ectosomal 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 redescribed from New Caledonian specimens (Hooper & Levi, 1993a), 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 & Levi (1993a). The synonymy presented above is mostly new, corroborated by re-examination of the relevant type material, although Clathria corallitincta Dendy (1889b) was already merged with C. frondifera by Dendy (1905), who also commented on the fact that ectosomal skeletal development varied substantially in the Gulf of Manaar population. Similarly, Clathria reinwardti var. palmata Ridley was synonymised with this species by Bergquist & Tizard (1967), and that decision is supported in the present study. 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 justification in maintaining the two species separately. Conversely, Rhaphidophlus filifer var. spinifera Lindgren, proposed as a synonym of C. frondifera by Thiele (1903a), is rejected here given that they differ substantially in most characters 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) construction, nearly completely lacks echinating acan- ^ REVISION OF M1CROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 405 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 I., 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, Wessel Is, Gove Peninsula, NT, 11°27.6'S, 136°28.7'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 12nun 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 sulface characteristics. Compressible, fibrous, basal skeleton firm, flexible. Ectosome and subectosome. Thick crust of ectosomal auxiliary subtylostyles erect on surfaec, in discrete plumose bundles, forming continuous palisade, incorporating some scattered sand grains, and abundant, granular, dark brown collagen in peripheral skeleton; subectosomal skeleton thick, plumose brushes of large subectosomal auxiliary subtylostyles supporting ectosomal brushes, arising from ends of peripheral choanosomal fibres; subectosomal region extensive; ectosomal and subectosomal skeletons detachable, up to 250p.m thick. Choanosome. Skeletal structure irregularly reticulate, slightly more compressed at core than in periphery; spongin fibres moderately light, 65-130p.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- ly closer to surface; primary fibres in axial region of skeleton close-set, short, irregularly interconnected by short secondary fibres, 30-50p,m diameter; all fibres cored by choanosomal principal subtylostyles (but barely different in morphology from auxiliary spicules); primary longitudinal fibres cored by multispicular tracts of principal spicules, up to 25 abreast, occupying less than 60% fibre diameter; secondary fibres cored by uni- or paucispicular tracts of principal spicules, occupying up to 40% of fibre diameter; all primary fibres heavily echinated by small acanthostyles but sparse on secondary fibres; mesohyl matrix heavy, granular, lightly pigmented, with abundant microscleres and some auxiliary megascleres scattered between fibres; choanocyte chambers large, oval, 50-75p,m diameter, lined by isochelae; some detritus incorporated into mesohyl but mainly in peripheral skeleton. Megascleres. Choanosomal principal subtylostyles long, slender, straight, with subtylote bases, usually microspined, tapering fusiform points. Length 307-(358.3)-395p.m, width 4-(5.4)-7p.m. Subectosomal auxiliary styles very similar in geometry but slightly shorter, more slender than principal spicules; long, slender, straight, subtylote smooth or microspined bases, fusiform points. Length 207-(248.8)-265p,m, width 2(2.7)-4p.m. Ectosomal auxiliary subtylostyles short, slender, straight, subtylote smooth or occasionally microspined bases, fusiform points. Length 151-(172.8)-186p.m, width 1.5-(2.1)-3p.m. Echinating acanthostyles short, slender, straight or slightly curved near distal end, subtylote, sharply pointed or blunt, more-or-less evenly spined; spines long, slender, prominently recurved. Length 78-(84.4)-95p,m, width 343.9)4.5p.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)-20p,m. Toxas very abundant, very slender; longer accolada toxas with slight angular curvature at centre, straight arms Length I: 1464306.8)415p,m, width 0.5-(1.2)-2.0p.m.; shorter wingshaped toxas moderately rounded at centre, reflexed arms. Length II: 33-(52.9)-72p.m, width 0.540.8)-1.5 gm. Reproductive products. Numerous, small, eliptical embryos present in holotype, 150-185p,m MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVU 406zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ AzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA C D FIG. 210. Clathria (Thalysias) wesselensis sp.nov. (holotype NTMZ3952). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C. Ectosomal auxiliary subtylostyles. D, Accolada and wingshaped 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.) cactifonnis 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 ljuniperina' 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.) cactifonnis and C. (T.) placenta where only primary fibres are cored by principal spicules and ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 407 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. MEMOIRS OF THE QUEENSLAND MUSEUM 408zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 principal 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.) cervicomis are most derived having lost their principal spicules completely. Axociella bitoxifera Koltun, 1970:202-204, text- OTHER SPECIES OF CLATHR1A Rhaphidophlus coriocrassus Bergquist & Fromont, (THALYSIAS) 1988:111-112, pI.51, figs e-f, p1.52, figs a-b; Dawson, 1993:39 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPORI13. New Zealand. Clathria (Thalysias) amabilis (Thiele, 1905) Stylotellopsis amabilis Thiele, 1905:456-457, textfig.72a-d [Punta Arenas]; Burton, 1932a:326 [Falkland Is]; Burton, 1940:115 [Argentina]; Koltun, 1964a:66 [Antarctica]; Sara, 1978:64-66 [Tierra del Fuego]. Rhaphidophlus amabilis; Van Soest, 1984b:129 [generic synonymy]. MATERIAL. HOLOTYPE: ZMB3309. SE. Pacific rim, SW. Atlantic, Antarctica. Clathria (Thalysias) anomala (Burton, 1933) Rhaphidophlus anomalus Burton, 1933:252-253, fig.3 [Natal]; Levi, 1963:67 [Natal]. MATERIAL. Holotype: NM1410. South Africa. Clathria (Thalysias) araiosa Hooper & Levi, 1993 Clathria (Thalysias) a raiosa Hooper & Levi, 1993a:1256-1259, figs 17-18, table 9 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300694 (fragments NTMZ3886, ORSTOMR1370). SW Pacific. Clathria (Thalysias) arteria (de Laubenfels, 1954) Axociella arteria de Laubenfels, 1954:148-149, textfig.96 [NW. Ponape]. MATERIAL. HOLOTYPE: USNM22876. central W. Pacific. Clathria (Thalysias) aruensis (Hentschel, 1912) Hymeraphia aruensis Hentschel, 1912:381, p1.20, fig. 38 [Am I., Arafura Sea]. Eurypon (Hymeraphia) aruensis; Levi, 1958:27. MATERIAL. HOLOTYPE: SMF955T. Indonesia. fig.29, p1.8, figs 1-2 [Kurile-Kamchatka Trench, NW. Pacific]. MATERIAL. HOLOTYPE: ZIL. NW Pacific. Clathria (Thalysias) calochela (Hentschel, 1912) Hymeraphia calochela Hentschel, 1912:383-385, p1.20, fig.41 [Am I., Arafura Sea]. MATERIAL. HOLOTYPE: SMF1679. Indonesia. Clathria (Thalysias) coriocrassus (Bergquist & Fromont, 1988) Clathria (Thalysias) corneolia Hooper & Levi, 1993 Clathria (Thalysias) corneolia Hooper & Levi, 1993a:1253-1256, figs 15-16, table 8 [New Caledonia]. MATERIAL. HOLOTYPE: QM0300691 (fragments NTMZ3877, ORSTOMR527). SW. Pacific. Clathria (Thalysias) cratitia (Esper, 1797) Spongia cratitia Esper, 1797:195, 196, 221, p1.53 ['East Indies']. Rhaphidophlus cratitius; Ehlers, 1870:18-19, 31; Ridley, 1884a:450-451; Ridley & Dendy, 1887:151152; Noll, 1888:51; Thiele, 1899:13; Thiele, 1903a:957-959 [Ternate, Moluccas]; Hallmann, 1912:177, 187. Thalysias cratita; de Laubenfels, 1954:137-138, textfig.87 [NW. Ponape, Caroline Is]. Microciona cratitia; Hartman, 1955:176-177. Desmacidon cratitia; Vosmaer, 1880:159. MATERIAL. HOLOTYPE: Unknown. Indonesia, central W. Pacific. Clathria (Thalysias) cullingworthi Burton, 1931 Clathria cullingworthi Burton, 1931a:345, p1.23, figs 3-4, text-fig.4 [Natal coast]; Levi, 1963:66. Thalysias cullingworthi; de Laubenfels, 1936a:105. MATERIAL. HOLOTYPE: NM1270 (fragments BMNH1933.7.4.68-70). South Africa. Clathria (Thalysias) delaubenfelsi (Levi, 1963) Rhaphidophlus delaubenfelsi Levi, 1963:60 - 62, textfig.70 [Cape of Good Hope]. Clathria (Thalysias) basiarenacea (Boury-Esnault, 1973) MATERIAL. HOLOTYPE: MNHNDCL618. South Africa. Rhaphidophlus^basiarenacea Boury-Esnau It, Clathria (Thalysias) amirantiensis sp. nov. 1973:287, fig.49 [San Antonio Bay, Brazil] MATERIAL. HOLOTYPE: MNHNDNBE972. SW Atlantic. Colloclathria ramosa Dendy, 1922:74-76, p1.7, fig.2, Clathria (Thalysias) bitoxifera (Koltun, 1970) p1.14, fig.4 [Amirante, Coetivy and Seychelles, Indian Ocean]. Rhaphidophlus ramosus; Van Soest, 1984b:99, 115. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 409 Not Rhaphidophlus ramosus Kieschnick, 1896:533; Kieschnick, 1900:569-570, p1.45, figs 47-50. Not Clathria ramosa Lindgren, 1897:482-483. MATERIAL. HOLOTYPE: BMNH1921.11.7. 64.Note: C. (T.) ramosa (Kieschnick, 1896) has priority. NW. Indian Ocean. Clathria (Thalysias) distincta (Thiele, 1903) Hymeraphia distincta Thiele, 1903a:956-957, fig. 21 [Ternate, Moluccas]; Hentschel, 1912:378-379. MATERIAL. Holotype: SMF789T. Indonesia. Clathria (Thalysias) encrusta Kumar, 1925 Clathria encrusta Kumar, 1925:221, fig.4 [India]. Thalysias encrusta; de Laubenfels, 1936a:105. MATERIAL. HOLOTYPE: IMP196/1. India. Not Rhaphidophlus filifer, Topsent, 1897b:425, 447; Desqueyroux-Faundez, 1981:758, table2. Not Rhaphidophlus filifer, Lindgren, 1898:283, 311, 312, p1.17, fig.7, p1.19, fig.17a-e'. Not Rhaphidophlus filifer var. spinifera; Lindgren, 1897:483; Hallmann, 1912:187; Levi, 1960a:55. Not Rhaphidophlus filifer var. mutabills; Topsent, 1897b:447-448, p1.20, fig.24, p1.21, fig.33. Not Rhaphidophlus filifer var. cantabrica; Orueta, 1901:331-335, text-figs 1-5, pls 3-4. cf. Microciona probfera; Vosmaer, 1935a:637. MATERIAL. HOLOTYPE: BMNH1887.5.2. 104. Philippines, Indo-Malay region. Clathria (Thalysias) flabellata (Burton, 1936) Clathria (Thalysias) eurypa Rhaphidophlus flabellata Burton, 1936:145, fig .4 [Oudekraal, South Africa]; Levi, 1963:67 [note]. MATERIAL. HOLOTYPE: BMNH1935.10. 21.3. South (de Laubenfels, 1954) Africa. Dictyociona eutypa de Laubenfels, 1954:143, fig.91 [Palau Is]. Microciona eurypa; Bergquist, 1965:164, 165, figs 21a-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 fascicularisTopsent, 1889:35-37, fig.3 [Banc de Campeche] Pseudanchinoe fascicularis; de Laubenfels, 1936a:109. Rhaphidophlus fascicularis; Van Soest, 1984b:108, 111, 122, table 4. ClathriadentataTopsent,1889:37-38, fig.4A [Banc de Campeche]. MATERIAL. HOLOTYPE: MNHN missing (Van Soest, 1984b:108). NE Atlantic. Clathria (Thalysias) fasciculata Wilson, 1925 Clathria (Thalysias) flabellifera Hooper & Levi, 1993 Clathria (Thalysias) flabellifera Hooper & Levi, 1993a:1250-1253, figs 13-14, table 7 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300693 (fragments NTMZ3884, ORSTOMR1416). SW Pacific. Clathria (Thalysias) hartmani (Simpson, 1966) Axocielita hartmani Simpson, 1966:2393; Simpson, 1968a:63-65, p1.15, text-figs 6-7, table 24 [San Juan I., Washington]. MATERIAL. HOLOTYPE: PMNH. NE Pacific. Clathria (Thalysias) hechteli sp. nov. Microciona microchela Hechtel, 1965:41-42, textfig.7 [Port Royal, Jamaica]; Wintermann-Kilian & Kilian, 1984:134 [Colombia]. Not Dictyociona microchela; de Laubenfels, 1953a:528. cf. Rhaphidophlus schoenus; Van Soest, 1984b:122. MATERIAL. HOLOTYPE: PMNH 5040. PARATYPE Clathria fasciculata Wilson, 1925:442, p1.42, fig.6, p1.49, 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,1981:20-21; Van Soest, 1989b:1-2, fig. 47. Thalysias fasciculata; de Laubenfels, 1936a:105. MATERIAL. HOLOTYPE: USNM21326. Indonesia, Philippines, central W Pacific, Japan. (Van Soest, 1984) Rhaphidophlus isodictyoides Van Soest, I984b: 118120, p1.8, fig.6, text-fig.47, table 4 [Curacao]. MATERIAL. HOLOTYPE: ZMAPOR4781. Caribbean. Clathria (Thalysias) filifera Clathria (Thalysias) jolicoeuri USNM24498. Caribbean. C. (C.) microchela (Stephens, 1916) has priority. Clathria (Thalysias) isodictyoides (Ridley & Dendy, 1886) (Topsent, 1892) Rhaphidophlus filifer Ridley & Dendy, 1886:475; Ridley & Dendy, 1887:152, 247, 255, p1.28, fig.2, p1.46, fig.9 [Masbate, Philippines]; Thiele, 1899:13-15; Kirkpatrick, 1900a:136; Thiele, 1903a:958; Dragnewitsch, 1905:3, 16-17; Dragnewitsch, 1906:441 [Singapore]; Whitelegge, 1907:503; Weltner, 1910a:33; Ferrer Hernandez, 1914:4,42. Rhaphidophlus jolicoeuri Top sent, 1892c:25 [Banyuls, Mediterranean]; Topsent, 1893d:446; Topsent, 1894a:19; Loisel, 1898:38; Topsent & Olivier, 1943:2 [Monaco]; Topsent, 1925:658-660, text-fig.14 [Gulf of Naples]; Levi, 1960b:55,65 [Dakar, N. Atlantic, Mediterranean, Naples and Monaco]; Boury-Esnault, 1971:327 [Banyuls]; MEMOIRS OF THE QUEENSLAND MUSEUM 410zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Pulitzer-Finali, 1983:610; Pansini & Pronzato, 1985:5 [Mediterranean]. Tenacia jolicoeuri; Levi, 1959:133-134, text-fig.26 [Sao Tome, Gulf of Guinea]. cf.Microciona prolifera; Vosmaer, 1935a:641. MATERIAL. HOLOTYPE: MOM (fragment BMNH1953.11.9.42). NW Atlantic, Mediterranean. Clathria (Thalysias) kilauea (de Laubenfels, 1951) Axocielitakilauea de Laubenfels, 1951a:262-263, textfig.9 [Coconut I., Hawaii]. Axociella kilauea; Hechtel, 1965:43-44 [note]. MATERIAL. HOLOTYPE: USNM22779. Central Pacific. Clathria (Thalysias) lambda (Levi, 1958) Leptoclathria lambda Levi, 1958:38, text-fig.35 [Marmar, 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, textfig.102 [Ailing-lap-lap, Truk]. Axociella linda; Hechtel, 1965:43-44 [note]. MATERIAL. HOLOTYPE: USNM22860. NW central Pacific. Clathria (Thalysias) lissoclada (Burton, 1934) Rhaphidophlus lissocladus Burton, 1934b:32-33, 5152, p1.4, fig.!, text-figs 4-5, 16 [Falkland Is]; Levi, 1963:62, p1.9, figs H,J, text-fig.71 [Cape of Good Hope, South Africa]. MATERIAL. HOLOTYPE: ZRS955 (fragment BMNH1933.3.17.176). PARATYPES BMNH 1933.3.17.30, 31, 32, 38, 154. South Africa, SW Atlantic. Clathria (Thalysias) longitoxa (Hentschel, 1912) Hymeraphia longitoxa Hentschel, 1912:381, p1.20, fig.39 [Am I., Arafura Sea]. Microciona longitoxa; Burton, 1938a:30-31, p1.5, 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-fig.6 [Coconut and Hawaii Is, Hawaii]; de Laubenfels, 1957:240 [Oahu, Hawaii]; Bergquist, 1977:65 [Hawaii]. 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, 1936a:113 [note]; Hechtel, 1965:43 [note]. MATERIAL. HOLOTYPE: ZMB3303 (paratypes ZMB3304, BMNH1930.11.28.21). SW Atlantic, SE Pacific. Clathria (Thalysias) micropunctata (Burton & Rao, 1932) Tenacia micropunctata Burton & Rao, 1932:340-341, text-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, text-fig.45, table 4 [Curacao]; Kobluk & Van Soest, 1989:1216; Meesters et al., 1991:195 [Curacao, Bonaire]; Muricy et al., 1991:1187 [SE. Brazil]. MATERIAL. HOLOTYPE: ZMAPOR4796. Caribbean. Clathria (Thalysias) mutabilis (Topsent, 1897) Rhaphidophlus filifer var. mutabilis Topsent, 1897b:447, p1.20, fig.24, p1.21, fig.33 [Ambon, Banda Sea]. Rhaphidophlus mutabilis; Desqueyroux-Faundez, 1981:743, figs 49-54, 116. MATERIAL. HOLOTYPE: MHNGC-12/27 (fragment MNHNDTI834). Indonesia. Clathria (Thalysias) naikaiensis (Hoshino, 1981) Eurypon naikaiensis Hoshino, 1981:153-155, p1.6, fig.8, text-fig.8 [Sasajima, Japan]. MATERIAL. HOLOTYPE: MMBSSIS090-4-a. Japan. Clathria (Thalysias) nervosa (Levi, 1963) Axociella nervosa Levi, 1963:65-66, p1.9E, 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 Hentschel, 1912:298, 359, 364-365, p1.19, fig.28 [Aru I., Arafura Sea]. Tenacia nuda; Hallmann, 1920:771. Thalysias nuda; de Laubenfels, 1936a:105. cf. Microciona prohfera tropus senta; Vosmaer, I 935a:649. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 411 MATERIAL. HOLOTYPE: SMF1576 (fragment MNHNDCL2278). Indonesia. Clathria (Thalysias) ongulensis (Hoshino, 1977) Axociella ongulensis Hoshino, 1977a:45, text-fig.3, p1.1, fig.3 [fossil demosponge; W. Ongul I., LutzowHolm Bay, Antarctica]. MATERIAL. HOLOTYPE: MMBS. Antarctica. Clathria (Thalysias) orientalis (Brondsted, 1934) Rhaphidophlus schoenus; Van Soest, 19846:112-113, p1.8, figs 1-4, text-fig.44, table 4 [Curacao, Bonaire, Puerto Rico]; Chen & Mok, 1993: 278 [probable misidentification, Taiwan]. Not Microciona microchela Hechtel, 1965:41, textfig.7 [Curacao, Bonaire, Puerto Rico, Jamaica]. MATERIAL. HOLOTYPE: USNM22404. Carribean. Clathria (Thalysias) tener Carter, 1887 Thalysias tener Carter, 1887a:70 [Mergui Archipelago]. MATERIAL. HOLOTYPE: IMEN14 (Reniera fibrosa') Rhaphidophlus orientalis Brondsted, 1934:20-22, textfigs 20-22 [Aru I., Arafura Sea]. MATERIAL. HOLOTYPE: Unknown. Indonesia. (fragment BMNH1887.6.1.9). Andaman Sea. Imperfectly known. Clathria (Thalysias) originalis Rhaphidophlus Pifer, in part, Topsent, 1897b:425, 447, p1.20, fig.22 [Ambon, Indonesia]; Desqueyroux-Faundez, 1981:758, table 2. Not Rhaphidophlus filifer Ridley & Dendy, 1886:475. Rhaphidophlus topsenti Thiele, 1899:15, p1.2, fig.3 [ Sulawesi, Indonesia]; Whitelegge, 1907:503; Hallmann, 1912:177. cf. Microciona prolifera; Vosmaer, 1935a:611,643 MATERIAL. HOLOTYPE: NMB20 (dry) (fragments (de Laubenfels, 1930) Esperiopsis originalis de Laubenfels, 1930:27; de Laubenfels, 1932:70-72, text-fig.38 [California]. Axocielita originalis; Lee & Gilchrist, 1985:24-32 [biochemistry]; Sim & Bakus, 1986: I 1 [California]; Bakus & Green, 1987:71 [S California]. MATERIAL. HOLOTYPE: USNM2 1441, paratype BMNHI929.8.22.54. NE Pacific. Clathria (Thalysias) oxeota (Van Soest, 1984) Rliaphidophlus oxeotus Van Soest, 1984b:120-122, text-fig.48, table 4 [Curacao]. MATERIAL. HOLOTYPE: ZMAPOR4880. Caribbean. Clathria (Thalysias) oxitoxa Levi, 1963 Clathria oxitoxa Levi, 1963:54-56, text-fig.62 [Humansdorp, South Africa]. Rhaphidophlus oxitoxa; Van Soest, 1984b:115, 122. MATERIAL. HOLOTYPE: MNHNDCL610. South Africa. Clathria (Thalysias) pachyaxia (Levi, 1960) Axociella pachyaxia Levi, 19606:763-764, text-fig. 16, [Senegal, W. Africa]. MATERIAL. HOLOTYPE: MNHNDCL787. NW Africa. Clathria (Thalysias) robusta (Dendy, 1922) Microciona strepsitoxa var. robusta Dendy, 1922:6061 [Amirante 11. Tenacia robusta; Burton & Rao, 1932:339-340 [Singapore]. Not Clathria robusta Koltun, 1959:186, p1.25, 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 Arndt, 1927:148, p1.1, fig.3, text-fig.9 [Curacao]. Aulospongus schoenus de Laubenfels, 1936a:100, p1.13, 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]. Clathria (Thalysias) topsenti (Thiele, 1899) ZMB2903, BMNH1908.9.14.167). Indonesia. Clathria (Thalysias) tricurvatifera (Carter, 1876) Thalysias tricurvatifera Carter, 1876:311-312 [Cape 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]; Kobluk & Van Soest, 1989:1216. Rhaphidophlus venosus; Meesters et al., 1991:194-195 [Curacao, Bonaire]. Rhaphidophlus raraechelae Van Soest, 1984b:116118, p1.8, fig.5, text-fig.46, table 4 [Curacao]; Pul itzer-Finali, 1986:151 [West Indies]. MATERIAL. HOLOTYPE: Cuba. Holotype of raraechelae: ZMAPOR4874. Caribbean. Clathria (Thalysias) virgultosa (Lamarck, 1814) Spongia virgultosa Lamarck,1814; Duchassaing & Michelotti, 1864:86, p1.23, fig.3. Thalysias virgultosa; Duchassaing & Michelotti,1864:86, p1.23,fig.3 [St. Thomas, Caribbean]; Tortonese,1962:3; de Laubenfels,1936a:104,106; Hartman,1955:173; Levi, 1960a:52. Microciona plana Carter, 1876:238, 472. Clathria copiosa Topsent, 1889:40-41, 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 [Antilles, Caribbean]; Carter, 1875:195; Hallmann, 1920:769; de Laubenfels, 1936a:106; Desqueyroux-Faundez & Stone, 1992: 73 [list]. MEMOIRS OF THE QUEENSLAND MUSEUM 412zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Clathria clathrata; Vosmaer, 1880:153; Ridley & Dendy, 1887:147; Wilson, 1902:397; Alcolado, 1976:5. Not Rhaphidophlus clathratus; Hal!mann, 1912:209; Topsent, 1920b:17-18; Topsent, 1932:97, p1.5, fig.6, text-fig.3. Pandaros juniperina; Duchassaing & Michelotti, 1864:90, p1.19, fig.3; de Laubenfels, 1936a:106. Thalysias juniperina; de Laubenfels, 1936a:105-107; Hartman, 1955:171-177; Lëvi, 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, p1.29, figs 3-5, p1.30, figs 1-3, text-figs 146-147; Carballeira, Shalabi & Maldonado, 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:109III, p1.7, fig.11, text-fig.43, table 4; Meesters eta!., 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 prohfera; Vosmaer, 1935a:608-611, 627, 628, 667, 630, 638, 644. MATERIAL. HOLOTYPE: Fragments of holotype of S. virgultosa: 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 Atlantic. 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, p1.17, fig, p1.19, fig.2 [St.Thomas]; Carter, 1882a:282, p1.11, 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. Thalysias hians Duchassaing & Michelotti, 1864:86, p1.16, 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, p1.18, 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), paralectotypeZMAPOR2373 (fragment MNHNDNBE1341). Referred to Tedaniidae. Thalysias massalis Carter, 1886a:50 [Port Phillip, Vic]. Reniera massalis; Dendy, 1895:236. MATERIAL. HOLOTYPE: BMNH1886.12. 15.433. Referred to Haplosclerida, Chalinidae. Thalysias proxima Duchassaing & Michelotti, 1864:84, p1.18, 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, p1.11, text-fig.I0 [Puerto Cabello and Antigua, West Indies]. Xestospongia ^subtriangularis; Wiedenmayer, 1977:257, table 51. MATERIAL. SYNTYPES: BMNH (3 specimens, unregistered). Referred to Haplosclerida, Petrosiidae. Thalysias rugosa Duchassaing & Michelotti, 1864:84, p1.18, 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. HOLOTYPE: no extant type material (Wiedenmayer, 1977; Van Soest et al., 1983)].Unrecognisable. Dictyocylindrus sessilis Carter, I 880a:38, p1.4, fig.2 [Gulf of Manaar, Ceylon]. Aulospongus sessilis; Dendy, 1905:176 [note]. MATERIAL. HOLOTYPE: LFM destroyed. Unrecognisable. MATERIAL. HOLOTYPE: MNHNDNBE1340 (fragment: MNHNDNBEI324, BMNH1928. 11.12.44, 56). Referred to Haplosclerida, Chalinidae, Haliclona. Thalysias subtriangularis Duchassaing, 1850; Duchassaing & Michelotti, 1864:85, p1.17, fig.! [St. Thomas, Caribbean]; Carter, 1879:287 [Kerguelen Is]; Carter, 1885c:196; Dendy, 1889a:58. Thalysias coccinea Duchassaing & Michelotti, 1864:84, p1.18, fig.5 [St.Thomas]. ? Isodictya mirabilis Bowerbank; Carter, 1878:159. 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. ? 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; lectotype of var. lyriformis: ZMAPOR2375, paralectotype of REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 413 var. lyriforrnis: 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, p1.5, fig.2. MATERIAL. HOLOTYPE: BMNH not found (poorly preserved fragments BMNH1887.5.21. 1331, 1332). Uncertain placement; possible Raspailiidae. Thalysiasvarians Duchassaing &Michelotti, 1864:86, p1.13, fig.6 [St. Thomas, Caribbean]. Anthosigmella varians; Topsent, 1918:557; de Laubenfels, 1957:242-243; Pang, 1973:47-50, textfig.14 [Jamaica; plus synonymy]; Wiedenmayer, 1977:255, table 50; Van Soest et al., 1983:204. MATERIAL. LECTOTYPE: TM POR 71 (fragments U5NM31048, 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 (rectangular) or isodictyal (triangular) choanosomal skeleton composed of acanthostyles and/or acanthostrongyles; (2) secondary (extra-axial, subectosomal) 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 spongin fibres, with or without axial compression; spongin fibres relatively poorly developed; additional 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. REMARKS. Under Van Soest & Stone's (1986) system all microcionids having a renieroid (and/or isodictyal) basal (or axial) skeleton composed of acanthose megascleres are grouped in Antho. This system is supported here with subgenera 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)strongyles forming the renieroid skeleton, less often acanthostyles, and a special category of echinating acanthostyles overlap the main skeleton); (3) A. (Isopenectya) (with an axially compressed and extra-axially renieroid reticulate skeleton composed of 2 forms of choanosomal spicules inside spongin fibres, overlaid by a second extra-axial plumose skeleton. Isopenectya (s.s.) could also be included in Echinoclathria, given the close resemblance in growth form and renieroid skeletal structure with E. leporina, but in A. (Isopenectya) 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 extraaxial (renieroid) regions, overlaid by a second extra-axial (plumose) skeleton composed of larger, smooth principal styles. In Echinoclathria, megascleres of the renieroid skeleton are exclusively smooth, and the larger, smooth principal 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; Levi, 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; PulitzerFinali, 1973: 40 (not Topsent, 1928a: 63). Quizciona de Laubenfels, 1936a: 111. TYPE SPECIES. Myxilla involvens Schmidt, 1864: 37 (by monotypy). MEMOIRS OF THE QUEENSLAND MUSEUM 414zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ E DO 0 tr) zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 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, Paralectotype MNHNDT3418. ^ 415 REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP DEFINITION. Primary basal renieroid (and/or isodictyal) choanosomal skeleton composed of acanthostyles and/or acanthostrongyles; secondary extra-axial (subectosomal) skeleton plumose, plumoreticulate, or simply composed of choanosomal styles echinating (project from) basal renieroid skeleton; spongin fibres poorly developed; special category of echinating acanthostyles absent; ectosomal skeleton with tangential, paratangential, or plumose tracts of 1 size of auxiliary styles or subtylostyles; microscleres include diverse forms of isochelae and toxas. REMARKS. Twenty one species have been included in, or referred to, Antho (Antho), although only 11 are widely accepted, 2 of which are known from Australia. Antho (Antho) opuntioides (Lamarck, 1815) (Figs 212-213) Unidentified sponge; Turgot, 1758: p1.24, fig.e. Alcyonium opuntioides Lamarck, 1815: 164. Anomoclathria opuntioides; Topsent, 1929: 21 26, text- figs 1-9. Antho opuntioides; Hooper & Wiedenmayer, 1994: 255. Not Anomoclathria opuntioides var. frondifera; Topsent, 1929: 26-29, text-figs 10-14; Topsent, 1932: 103, p1.1, figs 6-7. - MATERIAL. LECTOTYPE: MNHNDT654: Precise locality unknown, SW. Australia, Peron & Lesueur 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, possibly oscules, scattered over sides of branches. Texture and surface characteristics. Harsh, brittle in dry state, even, unornamented surface. Ectosome and subectosome. No ectosomal membrane intact (dry material), although remnants of sparse, tangential and paratangential skeleton composed of subectosomal auxiliary subtylostyles scattered near periphery; points of choanosomal styles and acanthostyles in peripheral skeleton protrude through surface; entire peripheral skeleton dense, virtually undifferentiated from deeper choanosomal skeleton (although spicule tracts with more sparse spongin component in periphery than at core), clearly dominated by close-set renieroid reticulation of acanthostyles. Choanosome. Skeleton with two distinct components: plumose extra-axial skeleton composed of multispicular or paucispicular continuous tracts of large choanosomal principal styles extending from centre of skeleton to ectosome; renieroid skeleton regular, tight meshed, rectangular and triangular meshes, 80-150pAn diameter, even mesh size throughout skeleton; spongin fibres heavier, slightly more compressed at centre of skeleton than in periphery, with oval meshes 6O-135m 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)-108p.m, width 4-(11.2)-1411m. Acanthose choanosomal styles of renieroid skeleton straight or slightly curved at centre, evenly spinose except for aspinose points and sometimes aspinose base; spines large, recurved, sharply pointed. Length 93-(104.5)-112p,m, width 8-(13.3)-161.m. Subectosomal auxiliary subtylostyles long, slender, straight, smooth or microspined bases, fusiform orr occasionally with telescoped points. Length 102-(118.4)-152p.m, width 2-(3.4)4.5m. Microscleres. Palmate isochelae large, unmodified, with lateral and front alae approximately equal length, lateral alae completely used to shaft, front ala detached along entire length. Length 16-(19.4)-2l p.m. Toxas wing-shaped, generously curved at centre, with only slightly reflexed points. Length 36-(68.9)-148p,m, width 1.5-(2.1)-3.0p.m. REMARKS. I initially thought this species conspecific with C. (T) styloprothesis (see above), based on Topsent's (1929) description of styloprothese', whereby spongin fibres are replaced by algal filaments, but the two species differ in spicule geometry and skeletal architecture (see also A. (P.) frondifera below). It differs from the allied A. (A.) tuberosa in growth form, MEMOIRS OF THE QUEENSLAND MUSEUM 416^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 417 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, WAM64881(1). 418zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM spicule geometry (particularly its acanthostyle morphology), and spicule sizes (refer to discussion in A. (A.) tube rosa). Antho (Antho) tuberosa (Hentschel, 1911) (Figs 214-215, Plate 9B-C) Lissodendoryx tuberosa Hentschel, 1911: 326-328, text-fig.21. lsociona tuberosa; Hallmann, 1920: 768; Burton & Rao, 1932: 341-342. Antho tuberosa; Hooper & Wiedenmayer, 1994: 255. Clathria hartmeyeri Hentschel, 1911: 379-381, text- fig.50. Thalysias hartmeyeri; de Laubenfels, 1936a: 105. cf. Microciona prolifera; Vosmaer, 1935a: 611, 648, 664. MATERIAL. HOLOTYPE: HM (fragment ZMB4417): 4km SW. of Denham, Shark Bay, WA, 25°56.5'S, 113°30.0'E, 3m depth, 10.vi.1905, coll. W. Michaelsen & R. Hartmeyer (dredge).0THER MATERIAL: WA- WAM645-81(1) (fragment NTMZ1722) QMG300203 (fragment NTMZ2958), NTMZ3214 (fragments PIB0004-352, QMG300044), QMG300678 (NCIQ66C-4229-N), NTMZ1466. S AUST- SAMTS4050 (fragment NTMZ1637). HABITAT DISTRIBUTION. Encrusting on macrophytes, coralline algae, corals, or other sponges; 3-40m depth; Shark Bay, Straggler Rocks, Houtman Abrolhos, 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 flattened 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 (2.5YR 8/8); yellow-brown or light brown in ethanol. Oscules. Indetectable in thinly encrusting specimen, scattered on lateral and upper surfaces of lobate specimens, up to 2.5mm diameter; surface minutely porous on lobate-flabellate specimen, pores 0.5-1.2mm diameter. Texture and surface characteristics. Texture compressible; surface optically even (encrustinglobate specimens) or striated and pitted, with ridges most prominent near margins (flabellate specimen). Ectosome and subectosome. Hispid, with tracts of smooth choanosomal styles from plumose skeleton protruding singly or in brushes; smaller subectosomal auxiliary subtylostyles form tangential tracts (encrusting specimen), paratangential tracts (lobate specimen), or distinct plumose brushes on ectosome, surrounding protruding choanosomal styles (flabellate specimen); subectosomal region undifferentiated from choanosomal skeleton; smaller acanthose choanosomal styles of renieroid skeleton terminate in uni- or paucispicular brushes just below surface; mesohyl 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 42-125p.m diameter; meshes more open in lobate specimens (92-197p.m) and flabellate specimens (97-208p.m maximum diameter); vaguely differentiated primary and secondary components of skeleton, best developed in flabellate specimens, with primary 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 differentiation of primary and secondary lines; plumose skeleton of larger, smooth, choanosomal principal styles forming pauci- or multispicular tracts; plumose tracts continuous, originating from basal attachment extending to peripheral skeleton in encrusting-lobate material, but not obviously continuous in flabellate specimens, prominent only near periphery; true echinating acanthostyles absent; mesohyl matrix light, containing few loose extra-fibre spicules; spongin fibres 45-108p.m diameter relatively light, only barely differentiated from mesohyl matrix in flabellate specimens (with only primary ascending elements and small interconnecting secondary fibre components obvious); spicule skeleton only minimally associated with fibre skeleton, each forming more-or-less independent support systems; choanocyte chambers relatively large, 155-65211m diameter, ovoid, often paired, lined by isochelae and rarer toxas. Megascleres. Smooth choanosomal principal subtylostyles of plumose skeleton slightly curved, fusiform, with tapering, rounded or subtylote smooth bases, occasionally slightly microspined; very variable in length. Length 164(213.0)-337p.m, width 6.5-(9.9)-14.0p.m (holotype 126-295 x 6.5-9p.m). Acanthose choanosomal styles of renieroid skeleton slightly curved or straight, subtylote, ^ 419 REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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. fl, Wing-shaped toxas. 420^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MEMOIRS OF THE QUEENSLAND MUSEUM fusiform, with evenly distributed small spines; acanthose choanosomal spicules include intermediates between larger smooth megascleres of plumose skeleton and entirely spined spicules of renieroid skeleton. Length 86-(114.9)-136p,m, width 3.5-(6.3)-10.0Rm (holotype 63-128 x 2.56 p.m). 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)-274p.m, width 1.8-(3.1)-4.5p.m (holotype 129-214 x 2.5-411m). Microscleres. Palmate isochelae common, unmodified, 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)-18p,m (holotype 10-15p,m). Toxas wing-shaped, relatively common, ranging from small thin forms to long relatively thick forms, all with evenly rounded, wide central curvature, and straight or very slightly reflexed points. Length 37494.5)-23211m, width 0.8(1.8)-3.6Rm (holotype 6-58 x Larvae. Larger flabellate-lobate specimen (WAM645-81(1)) contained ovoid-elongate, incubated embryos, 210-240Rm diameter, with some cellular differentiation and few larval megascleres. Associations. Several specimens had parasitic zooanthids on surface. REMARKS. This species is relatively polymorphic 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 lobateencrusting 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 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 discovered in any museum and this synonymy cannot 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.) involvens) the two genera are close in skeletal architecture, but A. (A.) tuberosa has entirely monactinal megascleres, whereas A. (A.) involyens has basal (renieroid) monactinal and/or diactinal spicules (acanthostyles, acanthostrongyles). Also included in Isociona 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 dimensions (smooth choanosomal styles 152-238x1115p.m, acanthose choanosomal styles 129-166x8-14p.m, subectosomal subtylostyles 133-293x3-61.Lm, palmate isochelae and toxas 18-178x0.8-3Rm). 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, 1986zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJI Van Soest & Stone, 1986: 42-44, figs 1-3 [Norway] Ant ho brat t egardi MATERIAL. HOLOTYPE: ZMAPOR5190. PARATYPE BMNH1982.9.6.1. NE. Atlantic. species offia de Laubenfels. Antho (Antho) brondstedi Bergquist & Fromont, 1988 Bergquist & Fromont, 1988: 97, pls 46d-f, 47a-c [New Zealand]; Dawson, 1993: 44 [index to fauna]. Ant ho brondst edi MATERIAL. HOLOTYPE: NMNZPOR111. New Zealand. Antho (Antho) dichotoma (Esper, 1794) REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 421 Spongia dichotoma Esper, 1794: 202, p1.10 [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, p1.3, fig.4, p1.6, fig.8 [Norway]; Thiele, 1903b: 394; Koltun, 1959: 184, p1.29, fig.!, 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, p1.12, fig.3, p1.15, fig.25a-b [N. of Scotland]. Clathria abyssorum; Vosmaer, 1880: 154 [Faroe Is, N.Atlantic]; Arndt, 1913: 119. Raspailia abyssorum; Fristedt, 1885: 48, p1.4, fig.!. Antho dichotoma; Alander, 1942: 63 [Sweden]; Van Soest & Stone, 1986: 44 [Norway]. MATERIAL. HOLOTYPE: unknown; holotype of D. abvssorum: BMNH1898.5.7.39. NE. Atlantic. Antho (Antho) graceae (Bakus, 1966) Burtonanchora lacunosa; de Laubenfels, 1961: 195197. Not Myxilla lacunosa Lambe, 1892: 70-71. Anthoarcuata graceae Bakus, 1966: 431-432, textfig.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, 1936a: 78. MATERIAL. HOLOTYPE: MOM (fragment MNHNDT1884). NE Atlantic. Antho (Antho) heterospiculata(Brondsted, 1924) Microciona heterospiculata Brondsted, 1924: 465, text-fig.20 [Colville Channel, NZ]. Quizcionaheterospiculata; de Laubenfels, 1936a: Ill. 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, p1.4, 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]; Levi, 1960a: 57, 76-80, text-figs 19-22 [var. inconstans; Atlantic, Mediterranean]; Vacelet, 1960: 267 [Mediterranean]; Vacelet, 1961: 41 [Corsica, Mediterranean]; Sara, 1961: 48 [Adriatic]; Sara & Siribelli, 1962: 10, 36, 48; Levi, 1963: 62-63, text-fig.72 [Mossel Bay, South Africa]; Sara, 1964: 228-229 [Ligurian Sea, Mediterranean]; Poggiano, 1965: 3, 7; Riitzler, 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 SolOrzano & Rodriguez Babie, 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 [Alboran Sea]; Uriz et al., 1992: 104 [Balearic Is]; SolOrzano et al., 1991: 177 [Galicia, Spain]; Ackers, Moss & Picton, 1992: 139 [Ireland]. Desmacodes involvens; Vosmaer, 1880: 108; Vosmaer, 1885: 235. Myxilla banyulensis, in part; Topsent, 1892b: 23; Topsent, 1902: 351, 363, 366; Cotte, 1903: 423. Clathria morisca Schmidt, 1864: 37, 45 [Adriatic]; Schmidt, 1868: 9, 41, 43, p1.2, fig.7 [Mediterranean]; Vosmaer, 1880: 150-151 [Algiers]; Topsent, 1902: 329. Dictyoclathria morisca; Topsent, 1920b: 18-21; Topsent, 1928a: 301-302, p1.3, fig.3 [Porto Santos, Azores]; Levi, 1959: 134, text-fig.27, p1.5, fig.! [Rio de Oro, Gulf of Guinea]; Levi, 1960b: 761-762, text-fig.15 [var. anisotyla; SW. Cape of Naze, W. Africa]; Sara, 1960a: 462 [Ischia, Mediterranean]; Desqueyroux-Faundez & Stone, 1992: 35 [index]. Plocamia zyxwvutsrqponmlkjihgfedcbaZYXWVUTSR inconst ans Topsent, 1925: 661-664, textfig.15 [Gulf of Naples]; Topsent, 1939: 6; PulitzerFinali, 1983: 610 [list]. Plocamilla inconstans; Burton, 1935a: 402. Holoplocamia inconstans; de Laubenfels, 1936a: 75. Antho inconstans; Ackers et al., 1992: 140 [Ireland]. lsodictya beani Bowerbank, 1866: 274, 334, 335 [Britain]; Gray, 1868: 164; Schmidt, 1870: 77; Bowerbank, 1874: 147, p1.58, 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 tnediterranea Babic, 1921: 87 [Adriatic]; Babic, 1922: 258-259, text-fig.B; Burton, 1930a: 528; Levi, 1960a: 57, 76-80; Maldonado, 1992: 1154 [possible synonym of A. (P.) novizelanica]. Microciona virgula Sara & Siribelli, 1960: 77-79, textfig.22 [Bay of Naples]; Siribelli, 1960: 16-17, textfig.7A [Naples]; Sara, 1964: 228-229 [Mediterranean]. ? Artemisina paradoxa Babic, 1921: 87; Babic, 1922: 260-261, p1.8, fig.6, text-fig.c [Adriatic]; Topsent, 1925: 660; Levi, 1960a: 85-86 [Adriatic]; Ristau, 1978: 585-586 [note on affinities]. MEMOIRS OF THE QUEENSLAND MUSEUM 422zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Clathria paradoxa; Burton, 1930a: 528. Antho paradoxa; Pulitzer-Finali, 1983: 610. ? Raspailia incrustans Svarcevskij, 1906: 52, p1.5, fig.6, p1.7, fig.l. 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 involvens and inconstans. A nt h o (Antho) oxeifera(Ferrer-Hernandez, 1921) Clathria oxetfera Ferrer-Hernandez, 1921: 171, p1.1 [Mediterranean]; Levi, 1960a: 84-85 [incertae sedis; Mediterranean]; Pulitzer-Finali, 1983: 610 [list]. Labacea oxetfera; de Laubenfels, 1936a: 125 [?]. Antho oxeifera; Uriz & Maldonado, 1993: 359-362, figs 6-9 [W. Mediterranean]. MATERIAL. HOLOTYPE: Madrid. Mediterranean. Antho (Antho) paucispinaSara & Siribelli, 1962 Antho paucispina Sara & Siribelli, 1962: 48-51, textfig.12 [Mediterranean]; Pulitzer-Finali, 1983: 610. MATERIAL. Holotype: IZUG. Mediterranean. Antho (Plocamia) Schmidt, 1870 Plocamia Schmidt, 1870: 62. Dirrhopalum Ridley, in Ridley & Duncan, 1881: 477. Plocamiopsis Topsent, 1904a: 155. Heteroclathria Topsent, 1904b: 93. Lissoplocamia Brondsted, 1924: 470. Plocamilla Topsent, 1928a: 63; Levi, 1960a: 80. Anomoclathria; in part, Topsent, 1932: 103 (not Topsent, 1929: 26). Holoplocamia de Laubenfels, 1936a: 75. TYPE SPECIES. Plocamia gymnazusa Schmidt, 1870: 62 (by subsequent designation of Burton, 1935a: 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 by acanthostyles at spongin fibre nodes; basal renieroid skeleton overlays leptoclathriid or microcionid main skeleton composed of echinating (acantho-)styles and/or choanosomal styles, standing perpendicular to base or axis, joining with echinating megascleres to produce ascending plumose skeletal tracts; extra-axial (subectosomal) skeleton plumose, dendritic, or subisodictyal, composed of choanosomal styles, originating from substrate or simply confined to periphery, forming tangential, paratangential or plumose extra-axial tracts; ectosomal skeleton with or without specialised spiculation (1 or 2 categories of auxiliary styles); microscleres include 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. (P.) 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 frondifera Lamarck, 1814: 445; Lamarck, 1816: 374. Anomoclathria frondifera; de Laubenfels, 1936a: 108. Antho frondifera; Hooper & Wiedenmayer, 1994: 256. Anomoclathria opuntioides var. frondifera; Topsent, 1929: 26-29, text-figs 10-14; Topsent, 1932: 103, p1.1, figs 6-7. Hymeniacidon chftoni Bowerbank, 1862a: 773, p1.30, fig.9; Bowerbank, 1864: 276, figs 70, 291. Acarnia ftont; Gray, 1867: 515. Not Alcyonium opuntioides Lamarck, 1815: 164. MATERIAL. LECTOTYPE: MNHNDT565: Precise locality unknown, suspected to be SW. Australia (Turgot collection). PARALECTOTYPE: MNHNDT3356: same details. HOLOTYPE of H. chftoni: BMNH1877.5.21.608 (fragments BMNH1877.5.21. 616, 1185, 218): Precise locality unknown, SW Australia, coll. H. Clifton. HABITAT DISTRIBUTION. Ecology unknown; SW Australia (WA) (Fig. 216G). DESCRIPTION. Shape. Lobate, thickly flabellate, 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 converging 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 (although rarely intact in dry specimens), with some embedded detritus and sparse tangential and paratangential tracts or single auxiliary subtylostyles scattered near periphery, sometimes forming bundles protruding through ectosome; points of (smooth) choanosomal principal styles from ascending plumose tracts protrude only slightly ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 423zyxwvutsrqp AzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 216. Antho (Plocamia) frondifera (Lamarck) (lectotype MNHNDT565). A, Choanosomal principal subtylostyles. B, Acanthostrongyles of renieroid skeleton. C, Subectosomal auxiliary subtylostyles. D, Wingshaped toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Paralectotype MNHNDT3356. through collagenous surface membrane; subectosomal 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 components: 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 anastomosing; tracts associated with, but not necessarily coring, heavy, dark brown, spongin-coated algal filaments (ostensibly Ficus (Topsent, 1932)), which dominates skeleton; filaments up to 250ttm diameter, 300-400p.m apart, branching, diverging from base of sponge through sponge surface; renieroid skeleton composed of 1 or 2 acanthostrongyles abreast forming square or triangular meshes up to 120Rm diameter, even mesh size throughout skeleton, overlaying plumose skeleton; some detritus scattered between renieroid skeletal meshes, usually coated with spongin; mesohyl not intact although some granular collagen containing microscleres scattered between spicule meshes; choanocytes not observed. Megascleres. Choanosomal principal styles entirely smooth, short, robust, slightly curved at centre, with rounded or slightly subtylote bases, fusiform points. Length 88-(103.4)-118Rm, width 4-(8.1)-13p.m. Acanthostrongyles of the renieroid skeleton thick or thin, rounded or slightly subtylote at both MEMOIRS OF THE QUEENSLAND MUSEUM 424zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 217. Antho (Plocamia) frondtfera (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. ^ 425 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR TABLE 43. Comparison between species of Antho (Plocamia). Measurements in 1.1.m. A.(P.) A.(P.) A.(P.) frondifera A.(P.) ridleyi (Hentschel) (Lamarck) coriacea elegans (Hentschel) Holotype Holotype (Bowerbank) I (Ridley & Specimens 2 (SMF1709) (MNHNDT Dendy) (N=13) 565) ridleyi CHAR - ACTER Shape encrusting encrustinglobate Choanosomal styles 212-388 x 14-20 183-562 x 7-25 88-118 x 4-13 85-103 x 3-14 (encrusting) digitate 120-660 x 11-16.4 160-550 x 8.2-15 A.(P.) A.(P.) A.(A.) A.(P.)burtoni manaarensis ovizelanicum n circonflexa 6 5 (Levi) (Dendyf (Ridley)4 (Levi) digitate 475 x 20.6 Renieroid spicules 92-104 x 6.5-10 99-142 x 4.5-15 70-120 x 234 x 19 3-10 (strongyles) (strongyles (strongyles (strongyles ^ (strongyles Ectosomal spicules 129-209 x 1.8-4 100-252 x 1-6 Subectosomal spicules 298-388 x 4-7 Echinating spicules 170-202 x 8-13 231-473 x 2.5-13 107-248 x 5-15 encrusting encrusting 500 x 25 150-450 x 10 100-350 x 5-7 130-170 x 75-100 x 7 177 x 15.8 (strongyles) 10 (styloid) (strongyles) 130 120-184 x 1-2.5 Chelae 11-18 8-19 15-20 Toxas I 24-46 x 0.8-1.2 21-86 x I4 40-116 x 1-2 82-211 x 63-354 x Toxas II 62-117 x 7.9 (strongyles) digitate 425-430 x 2.7 75-158 x 7.9-12 14-16 20-30 x 1-3 150-460 x 2-6 140-180 x 5-11 14-20 44-200 x ? (spined) 190-230 x 2-4 2-11 2.7-10 (spined) (spined) (spined) 316.7 x 6.3 190-360 x 24.7 150-400 190-260 x 2-3 114 x 9.5 272.4 x 17.4 80-120 'present' 19 19 10-12 70 x 2.5 63.3 x 2.1 25-70 x 8 13-14 15-75 x 0.52 85-130 x 3 (spined) Sources: 1. Ridley & Duncan (1881: 481); Dendy (1922: 76); Levi (1960a: 81). 2. Ridley & Dendy (1887: 158); Dendy (1922: 77); Levi (1960b: 760); Pulitzer-Finali (1973: 35). 3. Ridley & Duncan (1881: 482). 4. Ridley & Duncan (1881: 483). 5. Levi (1960a: 81). 6. Levi (1952: 53) ends, heavily spined particularly at points, spines large, conical or slightly recurved, sharply pointed. Length 85-(95.2)-103pLm, width 347.8)1411m. Subectosomal auxiliary subtylostyles long, very slender, curved at centre or sinuous, subtylote usually microspined bases, fusiform or occasionally telescoped points. Length 1204135.9)-184p.m, width 1-(1.7)-2.51.m. Microscleres. Palmate isochelae large, unmodified, with front and lateral alae approximately same length, lateral alae entirely fused to shaft, front ala detached along lateral margin. Length 15-(17.4)-2011m. Toxas wing-shaped, short, moderately thick, with large central curvature and slightly reflexed points. Length 40-(67.6)-116p,m, width 141.3)2pAn. REMARKS. Spongin fibres are excluded from the skeleton and replaced entirely by algal filaments ('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 symbiosis A. (A.) opuntioides, and A. (P.) fronchfera 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. (A.) opuntioides, differing in spicule geometry (acanthostrongyles 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 Hentsche1,1912: 387-388, p1.20, fig.44. Holoplocamia ridleyi; de Laubenfels, 1936a: 75. Antho ridleyi; Hooper & Wiedenmayer, 1994: 256. MATERIAL. HOLOTYPE: SMF1709 (fragment MNHNDCL2183): Mimien Bay, Am 1., Arafura Sea, Indonesia, 6°S, 134°50'E, 15m depth, 8.iv.1908, coll. H. Merton (dredge). OTHER MATERIAL: NTNTMZ0299, NTMZ2108, NTMZ2110, NTMZ2112, NTMZ2131, NTMZ2142, NTMZ2201, QMG300146 (fragment NTMZ2212), NTMZ2221, QMG300507 (fragment NTMZ2230), QMG303295, NTMZ2556, NTMZ2378. WA- QMG301185. 426zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 43. (continued) CHARACTER Shape Choanosomal styles Renieroid spicules _ A.(P.) A.(P.) A.(P.) A.(P.) A.(P.) A.( P.) A(P.) A.(P.) A.(P.) illgizyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA barbadensis gymnazusa omata lambei aub inconstanszyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC penneyi (de plena del(I, / Si 8 itienfezyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 1e )7 (Bakus) (Topsent) 9 Laubenfels) I 9 (Sollas)" (Burton)" (Van Soest)" (Schmidt)" encrusting encrusting encrusting flabellate encrusting encrusting encrusting 15 225-884 x 18-37 190-340 x 13-17 385 x 12 1004 x 4449 687 x 1619 200-500 x 8 213-248 x 4 112-153 x 9 120-283x 10-30 130-150x 10-12 (strongyles (strongyles encrusting 199-306 x (strongyles) 122 x 7 184 x 12 100 x 8 159-301 x 5- 479 x 6 (strongyles) (strongyles (strongyles 8 (styles) (strongyles Ectosomal spicules Chelae Toxas I encrusting 372 x 14 68-136 x 810 (stongyles) _ 368 x 10 176 x 3 220-304 x 1.5-3 97 x 9 245 x 24.5 183 x 9-13 57-73 x 3-4.5 16-20 12 7-14 13 6-9 19 50-200 x 3 30-80x 0.5-2 7-10 65 40-108 82 x 3 159-733 x 4-17 230-290 x 4-5 - 177-358 x 10-29 'present 11-16 17-27 Subectosomal spicules Echinating spicules zyxwvutsrqponmlkji (Dendy)" 48-103 11-133 99 x 6 172 x 10 Toxas II Sources: 7. Little (1963: 45). 8. Bakus (1966: 440); Simpson (1968a:43). 9. Topsent (1925: 662). 10. de Laubenfels (1936a: 76). 11. Sollas (1879: 44). ^12. Bakus (1966); Lambe (1895: 124). 13. Van Soest (1984b: 125). ^14. Ridley in Ridley & Duncan (1881: 478).^15. Dend^(1924:351) HABITAT DISTRIBUTION. Holotype collected from a sand substrate, encrusting on an Oceanapia species (Niphatidae; incorrectly identified as Phloedictyonfistulosa (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 Harbour (NT); Hibernia Reef, Sahul Shelf (WA) (Fig. 2181); also Aru Is, Indonesia (Hentschel, 1912). DESCRIPTION. Shape. Thinly, thickly or bulbous-encrusting, 2-14mm thick, forming extensive overgrowths on coral substrata. Colour Live colouration consistent, even bright red or blood red (Munsell 5R 5-3/10), turning grey or pinkish grey in ethanol (5R 8/2-4). Oscules. Large oscules on exterior of bulbous lobes, 0.4-1.6mm diameter, with slightly raised membraneous lip; minute pores, up to 150p,m 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 preservation, but thicker specimens retain shape. Ectosome and subectosome. Surface microscopically hispid, with points of smooth choanosomal principal styles protruding and surrounded by plumose brushes of mostly smaller ectosomal auxiliary subtylostyles (with fewer larger subectosomal megascleres contributing to ectosomal skeleton); subectosomal region structurally variable; thinly encrusting specimens with peripheral skeleton not clearly delineated from choanosomal skeleton, containing only thick tangential or paratangential tracts, up to 14011m diameter, composed of larger subectosomal auxiliary subtylostyles; in thicker bulbous specimens subectosomal region cavernous, containing numerous plumose, stellate brushes composed 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, extrafibre tracts, 25-65tim diameter. Choanosome. Skeletal structure with 3 distinct components: hymedesmoid skeleton, with basal layer of spongin fibre lying on substrate, 60240Rm thick, with smooth choanosomal principal styles and echinating acanthostyles perpendicular to substrate; renieroid skeleton forming regularly reticulation of acanthostrongyles, overlaying hymedesmoid basal skeleton, in pauci- or multispicular tracts (vaguely ascend- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 427 ing) and uni- or paucispicular (irregularly transverse) tracts, producing triangular (isodictyal) or rectangular (renieroid) meshes, 60150p.m diameter, without any obvious spongin fibre component; echinating acanthostyles, occurring singly or in plumose brushes, at major nodes of renieroid skeleton, sometimes also forming irregularly plumose, discontinuous, ascending 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 leptoclathriid basal skeleton to peripheral region, becoming more plumose or dendritic towards periphery; mesohyl matrix heavy but virtually unpigmented, surrounding renieroid meshes; choanocyte chambers circular to oval, 63-95Rm 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 slightly subtylote bases, smooth or with lightly microspined bases, fusiform points. Length 183(317.9)-562p,m, width 6.54 14.8)-25p,m (holotype 212-388 x 14-20p,m). Acanthostrongyles of renieroid skeleton short, thick, straight or slightly curved, with either symmetrical subtylote bases, or asymmetrical ends (subtylote bases, rounded or slightly subtylote points); usually evenly microspined, spines small, conical, sharply pointed. Length 98(120.6)-142p,m, width 4.5-(10.1)-15p.,m (holotype 92-104 x 6.5-10p,m). Subectosomal auxiliary subtylostyles long, thin, fusiform, straight, with prominent tylote, subtylote or polytylote bases, bases microspined or less commonly smooth. Length 2314372.9)473p,m, width 2.5-(5.9)-12.5Rm (holotype 298388 x 4-7p,m). Ectosomal auxiliary subtylostyles identical in geometry but smaller than large auxiliary megascleres, with tylote or subtylote, smooth or microspined bases. Length 100-(192.7)-252p,m, width 1.1-(3.3)-6.5pLm (holotype 129-209 x 1.84p,m). 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 prin- cipal styles). Length 1074194.1)-24811m, width 5-(10.1)-15pm (holotype 170-202 x 8-13pLm). Microscleres. Palmate isochelae abundant, single size category, unmodified, with front and lateral alae approximately same length, lateral alae completely fused to shaft, front ala detached from front alae along lateral margin. Length 8-(l2.5)19p.m (holotype 11-18p,m). Toxas wing-shaped, incompletely differentiated into 2 size classes: smaller thin, extensive rounded central curvature, straight or slightly reflexed points; larger thick, sharply angular or slightly rounded central curvature, straight or very slightly reflexed arms characteristically bearing terminal bulbous swelling and a single apical spine surrounded by smaller spines. Length I: 21-(44.3)-8611m, width 1-(1.9)-4.1p.m (holotype 24-46 x 0.8-1.2p,m); length II: 62(169.2)-355p.m, width 2-(4.7)-10.6p,m (holotype 82-211 x 2-4p,m). Larvae. Parenchymella larvae oval-elongate, 195-410p,m long, 135-330p,m wide, orangebrown alive, incompletely ciliated with a bare posterior pole. Younger larvae poorly differentiated in cellular construction, but containing clearly visible, longitudinally disposed, whispy sinuous monactinal megascleres. Older larvae have distinct, oval, cellular differentiation, radially disposed thin tylostyles, and small toxas and isochelae. Incubated larvae were abundant in 33% of specimens, collected between January and May 1985 in the Darwin region, but 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 be taken every month). Associations. All NT specimens were encrusting dead faviid coral heads, growing next to, or over other sponges (e.g., Stelletta, Mycale, Placospongia, Ulosa, Clathria), coralline algae and ascidians. Variation. Ectosomal structure variable, ranging from Clathria condition (with thin or thick tangential crust; 17%), Thalysias condition (with a stellate plumose ectosomal architecture; 50%), to merely paratangential (with elements of both structures; 33%). Subectosomal region cavernous (with plumose tracts of choanosomal and/or subectosomal megascleres; 67%), to merely tangential or paratangential (without subdermal cavities; 33%). Extra-axial (non-renieroid) skeleton ranges from subisodictyal, distinctly plumose, ascending, diverging towards periphery MEMOIRS OF THE QUEENSLAND MUSEUM ^ 428zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 218. Antho (Plocamia) ridleyi (Hentschel) (NTMZ2142). A, Choanosomal principal style. B, Echinating acanthostyles. C, Acanthostrongyles of renieroid skeleton. D, Subectosomal auxiliary subtylostyles. E, Ectosomal auxiliary subtylostyle. F, Accolada and wing-shaped toxas. G, Palmate isochelae. H, Section through peripheral skeleton. I, Australian distribution. J, NTMZ299. (58%), to irregularly disposed, mostly longitudinal extra-fibre tracts (48%). Echinating 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 429 FIG. 219. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Ant ho ( Plocam ia) ridleyi (Hentschel) (QMG301185). A, Choanosomal skeleton. B, Renieroid skeleton (x437). C, Acanthostrongyles of renieroid skeleton. D, Acanthostrongyle 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. MEMOIRS OF THE QUEENSLAND MUSEUM 430zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ NO. SAMPLES WITH LARVAE SEASON TOTAL SAMPLES WET 3 3 PREDRY I 0 DRY 7 0 PREWET 4 2 FIG. 220. Antho (Plocamia) ridleyi (Hentschel). Incidence of incubated parenchymella larvae in NT specimens. (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 differs 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). synonymy is clearly wrong. In their spicule diversity and growth form the two taxa are similar (Table 43), whereas comparisons between field observations on living populations of A. (P.) ridleyi (present study) and A. (P.) coriacea (Ackers, Moss & Picton, 1992: 141) show that the two species have quite different surface features, live colouration, and some differences in spicule dimensions (Table 43) indicating at most a possible sibling species relationship. The separate category of echinating acanthostyles, a renieroid skeleton composed of diactinal or quasi-diactinal spicules, and a more-or-less plumose (non-renieroid), subisodictyal skeleton of smooth choanosomal and subectosomal spicules are typical of Antho (Plocamia). However, Antho and Plocamia are barely differentiated on that basis and they are formally merged here. Some Plocamia have a mixture of both acanthostyles and acanthostrongyles in the renieroid skeleton (Levi, 1960a). Megascleres echinating fibre nodes may vary from true acanthostyles, with different geometry from other choanosomal spicules (A. (P.) barbadensis (Van Soest, 1984b)), or smooth styles which are differentiated from choanosomal megascleres only by their marginally smaller size and light, irregular spination (most species including A. (P.) ridleyi, A. (P.) elegans (Ridley & Dendy, 1886) and A. (P.) coriacea (Bowerbank, 1874)), or echinating megascleres may be entirely undifferentiated from choanosomal styles or absent (e.g., A. (P.) novizelanicum (Ridley, in Ridley & Duncan, 1881), A. (P.) penneyi (de Laubenfels, 1936a), A. (P.) fronclifera (Lamarck)). Similarly, the extra-axial (non-renieroid) skeleton varies between specimens. Some thinly encrusting species, such as A. (P.) ridleyi, A. (P.) delaubenfelsi (Little, 1963) and A. (P.)burtoni (Levi, 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 Antho (Plocamia) barbadensis (Van Soest, 1984) Plocamilla barbadensis Van Soest, 1984b: 125-126, text-fig.50 [Barbados, West Indies]. Antho barbadensis; Van Soest & Stentoft, 1988: 123 [Barbados]. MATERIAL. HOLOTYPE: ZMAPOR3832. Province: Caribbean. Antho (Plocamia) burtoni (Levi, 1952) Plocamilla burtoni Levi, 1952: 53-54, text-fig.17 [Senegal, W. Africa]; Levi, 1960b: 760 [note]. MATERIAL. HOLOTYPE: MNHN missing. Province: NW. Africa. Antho (Plocamia) circonflexa (Levi, 1960) Plocamilla circonflexa Levi, 1960a: 81-83, text-figs 24-25 [Brest, France]; Sara & Siribelli, 1960: 80 [Bay of Naples, Mediterranean]; Sara & Siribelli, 1962: 51 [Gulf of Naples]; Descatoire, 1966: 242, text-fig.6B [Glenan Archipelago, Brittany]; Pulitzer-Finali, 1983: 610 [list]. MATERIAL. HOLOTYPE: MNHN missing. NE. Atlantic, Mediterranean. Antho (Plocamia) coriacea (Bowerbank, 1874) Isodictya coriacea Bowerbank, 1874: 136, 228, p1.76, figs 7-12 [Britain]. Dirrhopalumcoriaceum; Ridley, 1881: 481, p1.29, figs 3-7 [Ireland]. Plocamia coriacea; Hanitsch, 1894: 173 [Britain]; Dendy, 1922: 76-77 [Amirante, Indian Ocean]. Plocatnilla coriacea; Topsent, 1928a: 63; Burton, 1935a: 402; Burton, 1959b: 44 [Iceland]; Levi, 1960a: 80-81, text-fig.23 [Roscoff, English Channel; Atlantic]; Sara & Siribelli, 1962: 51 [with question; Gulf of Naples]; Poggiano, 1965: 3,7; Pulitzer-Finali, 1983: 610 [list]; SolOrzano et al., 1991: 177 [Galicia, Spain]; Ackers, Moss & Picton, 1992: 141-142 [Ireland]. Holoplocamia coriacea; de Laubenfels, 1936a: 75. MATERIAL. Holotype: BMNH1877.5.21. 761/1910.1.1.251. NE. Atlantic, Mediterranean, E. Africa. Antho (Plocamia) delaubenfelsi (Little, 1963) Holoplocamia delaubenfelsi Little, 1963: 45-48, textfig.18 [Gulf of Mexico]. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 431 MATERIAL. HOLOTYPE: USNM23596. NE. Pacific. Antho (Plocamia) elegans(Ridley & Dendy, 1886) Plocamia elegans Ridley & Dendy, 1886: 475 [var.]; Ridley & Dendy, 1887: 158-159, p1.29, fig.9, p1.31, fig.! [var. elegans; Azores]; Topsent, 1892a: 117, p1.7, fig.!! [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; PulitzerFinali, 1973: 35-41 [Azores]. Plocamilla coriacea var. elegans; Levi, 1960b: 760761, text-fig. 13 [W. coast of Africa]. Holoplocamia elegans; de Laubenfels, 1936a: 75. MATERIAL. HOLOTYPE: BMNH1887.5.2. 109. NE Atlantic, E & W Africa. A ntho (Plocamia) erecta(Ferrer-Hernandez, 1923) Plocamia erecta Ferrer-Hernândez, 1923: 248, textfigs 1-3 [Spain]. Plocamilla erecta; Burton, 1935a: 402. Holoplocamia erecta; de Laubenfels, 1936a: 75 [Santander, Atlantic]. Antho erect a; Levi, 1960a: 80. MATERIAL. HOLOTYPE: Madrid. NE Atlantic. Antho (Plocamia) gymnazusa (Schmidt, 1870) Plocamia gymnazusa Schmidt, 1870: 62-63, p1.4, fig. 17 [Florida]; Burton, 1935a: 401; de Laubenfels, 1936a: 76. Dirrhopalum gymnazon; Ridley, 1881:478-479, p1.29, figs 1-2. MATERIAL. HOLOTYPE: BMNH1870.5.3.70 (fragment MNHNDCL1105L). Caribbean. Antho (Plocamia) illgi (Bakus, 1966) Plocamilla illgi Bakus, 1966: 440-443, p1.1 A, figs 6a-j [ San Juan Archipelago, Washington]; Simpson, 1968a: 43-47, 93, text-fig.3 [San Juan Is, Washington]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Bakus & Green, 1987: 73-74 [S.California]. Dickinson, 1945: 23, p1.35, figs 69-70, p1.36, figs 71-72 [Pacific Grove, California]. Antho lithophoenix; Van Soest, 1984b: 129 [generic synonymy for Isociona]; Lee & Gilchrist, 1985: 24-32 [biochemistry]; Sim & Bakus, 1986: 11 [California]. MATERIAL. HOLOTYPE: USNM21460, paratype BMNH1929.8.22.42. NE Pacific. Antho (Plocamia) manaarensis (Carter, 1880) Dictyocylindrus manaarensis Carter, 1880a: 34, p1.4, fig.1 [Gulf of Manaar, Ceylon]. Dirrhopalum manaarense; Ridley, 1881: 482. Plocamia manaarensis; Dendy, 1905: 179, p1.8, fig.I [Gulf of Manaar, Ceylon]; Burton & Rao, 1932: 355 [Laccadive Sea, Mangalore and Karwar, India]. Not Plocamia manaarensis; Lambe, 1895: 124, p1.2, figs Ila g [California]. Plocamilla manaarensis; Burton, 1935a: 402; Burton, - 1959a: 252-253 [Arabian Sea]; Bakus, 1966: 512. MATERIAL. HOLOTYPE: LFM destroyed (fragment BMNH1986.4.29.1b). India, Arabian Sea Antho (Plocamia) novizelanica (Ridley, 1881) Dirrhopalum novizelanicum Ridley, 1881: 483-485, p1.29, figs 8-16 [Bay of Islands, New Zealand]. Plocamilla novizelanicum; Burton, 1935a: 402. Plocamilla novizelanica; Levi & Levi, 1983a: 965966, text-fig.27 [S. of New Caledonia]; [?] Uriz, 1988a: 90-91, text-fig.65 [Namibia; ? affinity]. Holoplocamia novizelanica; de Laubenfels, 1936a: 75. Plocamia novizelanicum; Bergquist & Fromont, 1988: 120-122, p1.56, fig.f, p1.57, 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); ?SW Africa, Mediterranean. Antho (Plocamia) ornata (Dendy, 1924) Bubaris ornata Dendy, 1924a: 351, p1.14, figs 25-27. Plocamia ornata; Burton, 1928: 129. Axoplocamia omata; Burton, 1935a: 402. MATERIAL. HOLOTYPE: BMNH missing (fragments BMNH1923.10.1.126, 322). W Indian Ocean. MATERIAL. HOLOTYPE: USNM23737. NE Pacific. Antho (Plocamia) lamhei (Burton, 1935) Plocamia manaarensis, in part; Lambe, 1895: 124 [California]; Lambe, 1900: 161. Not Dictyocylindrus manaarensis Carter, 1880a: 37. Heteroclathria lambei Burton, 1935a: 403. Plocamilla zimmeri Bakus, 1966: 512. Antho (Plocamia) penneyi (de Laubenfels, 1936) Holoplocamia penneyi de Laubenfels, 1936a: 76 [Tortugas, Florida]. Antho penneyi; Van Soest & Stentoft, 1988: 126 [table]. MATERIAL. HOLOTYPE: USNM22460. Caribbean. MATERIAL. HOLOTYPE: USNM6331. NE Pacific. Antho (Plocamia) plena (Sollas, 1879) Antho (Plocamia) lithophoenix (de Laubenfels, 1927) Plocamia lithophoenix de Laubenfels, 1927: 268. Isociona lithophoenix; de Laubenfels, 1932: 99-100, text-fig.59 [California]; Burton, 1935a: 400 [note]; Plocamia plena Sollas, 1879: 44, pls 6-7 [W Africa]; Topsent, 1894: 21. Holoplocamia plena; de Laubenfels, 1936a: 75 [note]. Clathria plena; Vosmaer, 1880: 154 [Angola]. Dirrhopalum plenum; Ridley, 1881: 480-481. MEMOIRS OF THE QUEENSLAND MUSEUM 432zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 221. Antho (Isopenectya) chartacea (Whitelegge) (holotype AMZ436). 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 433 MATERIAL. HOLOTY PE: Bristol (fragment BMNH1909.8.15.3). W Africa. Antho (Plocamia) prima (Brondsted, 1924) Lissoplocamia prima Brondsted, 1924: 470, fig.24a-d [North Cape, New Zealand]; Topsent, 1928a: 63; Levi, 1963: 63, fig. 73 [S.Africa]. Plocamia prima; Bergquist & Fromont, 1988: 122, p1.57c-e. MATERIAL. HOLOTYPE: possibly UZC. South Africa, NZ. Antho (Plocamia) signata (Topsent, 1904) Plocamiopsis signata Topsent, 1904a: 155-157, p1.14, fig.1 [Azores]; Topsent, I928a: 306-307, p1.10, fig.20 [et var. mitis; W. of Flores, Azores]; Burton, 1935a: 402 [note]. MATERIAL. HOLOTYPE: MOM (fragment collection unknown, coll. FIV 'Thetis' (trawl). HOLOTYPE of A. petforata: BMNH1886.8.27.459: Broughton I., Port Stephens, NSW, 32°36'S, 152°19'E, other details unknown.OTHER MATERIAL. NSW- NTMZ2831, AMZ3605, AMZ3604, AMZ3606, AMZ4216 (RRIMPFN1339), AMZ4256 (RRIMPFN1435), AMZ4255 (RRIMPFN1434), AMZ3207, AMZ3162, AMZ4569 (RRIMP-59PJP), QMG303711, QMG303713. HABITAT DISTRIBUTION. 12-100m depth; rock platform, heads or outcrops on sand substrate; known only from Australia: Port Stephens, Botany Bay, Coogee, Long Reef, Dee Why, N. Sydney, Port Hacking, 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 Antho (Isopenectya) Hallmann, 1920 thick, with slightly digitate or evenly rounded margins. Isopenectya Hallmann, 1920: 789. Colour. Bright red-orange alive (Munsell 5R 5/10 Clathriella Burton, 1935c: 73; Koltun, 1959: 186. - lOR 6/10), pale brown in ethanol. Oscules. Not observed. TYPE SPECIES. Clathria chartacea Whitelegge, 1907: 497 (by monotypy). Text ure and surface charact erist ics. Firm, barely compressible, flexible, slightly spiky; optically DEFINITION. Three skeletal components: (1) smooth, even surface. renieroid reticulation of acanthose styles, (2) Ect osom e an d su bect osom e. Ectosome overlayed by isodictyal or subisodictyal reticula- prominently hispid, with pauci- or multispicular tion of smooth styles coring spongin fibres, (3) plumose brushes of larger, smooth choanosomal surmounted by plumose or radial extra-axial principal styles protruding through surface, formskeleton of larger smooth styles, perpendicular to ing a vestigial radial extra-axial skeleton, arising axis, in peripheral region; skeleton may be slight- from pauci- or multispicular tracts of (marginally compressed at core, spongin fibres only ly) smaller smooth principal styles in subecmoderately developed; echinating megascleres tosomal region; subectosomal auxiliary absent; ectosomal skeleton with single category subtylostyles tangential, paratangential, or rarely of auxiliary subtylostyle forming tangential or plumose, at base of protruding choanosomal paratangential tracts; microscleres absent. spicule brushes; peripheral skeleton relatively cavernous in comparison to the central REMARKS. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA I sopenect ya contains 4 species, 3 from the SW Pacific and 1 from the NW Pacific. choanosomal skeleton, with moderately heavy All lack microscleres but this is interpreted as mesohyl matrix. Choanosom e. Skeletal structure with 3 distinct secondary loss. components: (1) slightly compressed spongin fibres forming close-meshed anastomoses at core Antho (Isopenectya) chartacea (Whitelegge, of skeleton, more cavernous towards surface, (2) 1907) renieroid skeleton composed of acanthose styles, (Figs 221-222, Plate 9F) overlaying other structures; (3) longitudinal, ascending tracts of smooth principal styles, marClathria (?) chartacea Whitelegge, 1907: 497. ginally smaller than those protruding through Isopenectya chartacea; Hallmann, 1920: 789. Antho chartacea; Rudman & Avern, 1989: 335; surface, forming subisodictyal tracts at core, Hooper & Wiedenmayer, 1994: 255. more plumose in periphery, and usually (but not Antherochalina pelforata Lendenfeld, 1887b: p1.22, invariably) associated with larger spongin fibres; fig.44. spongin fibres in axial skeleton heavy, 48-84,m Not Antherochalina petforata, in part; Lendenfeld, diameter, producing irregularly oval or elongate I887b: 788; Lendenfeld, 1888: 89-90. meshes, 32-120Rm diameter, cored by uni- or MATERIAL. HOLOTYPE: AMZ436: Off Coogee, bispicular tracts of smaller, smooth choanosomal NSW, 33°45'S, 151°20'E, 98-100m depth, date of principal styles; fibres closer to surface, 19-4211m BMNH1930.7.1.36). NE Atlantic. MEMOIRS OF THE QUEENSLAND MUSEUM 434^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA diameter, regularly anastomosing, wide-meshed, 75-162p.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 multispicular ascending tracts of smooth choanosomal styles standing perpendicular to axis, becoming increasingly plumose, larger, and typically multispicular towards periphery; echinating megascleres absent; mesohyl matrix lightly pigmented, with few auxiliary spicules scattered throughout; choanocyte chambers elongate-oval, 36-75 pm 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)-312p.m, width 6-( (holotype 168-274x13-17p,m). 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 completely smooth shaft. Length 74-(86.1)-H2p,m, width 4-(7.2)-8.5p.m (holotype 92-127x9-12.5pLm). 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)-203p.m, width 2.5(2.9)-3.8p.m (holotype 163-243x2-4.5p.m). Echinating megascleres absent. Microscleres. Absent. Larvae. Viviparous, parenchymella larvae oval to elongate, 340-420x180-360p,m, with central core ofjuvenile styles, well differentiated cellular construction. Associations. Obligatory (?) host for nudibranch Rostanga sp. (AMC150065) (W. Rudman, pers.comm.). REMARKS. Hallmann (1920) erected Isopenectya 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 Isociella) in having a compressed axis and more-or-less plumose extra-axial skeletons cored by smooth choanosomal (principal) styles, in one or more size categories, together with the usual renieroid structure overlaying the remainder of the skeleton composed of acanthose (or sometimes smooth) styles different from principal spicules. This species superficially resembles Ophlitaspongia tenuis (Carter) (= Echinoclathria leporina (Lamarck)) mainly due to the emphasis of the compressed central skeleton and subrenieroid skeletal structure in both species, whereas megascleres forming these skeletons are quite different. Choanosomal megascleres in A. (I.) 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 perpendicular 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 contrasted with the renieroid Atnphinomia (Raspailiidae), which also has acanthose structural spicules (Hooper, 1991). Antho (Isopenectya) punicea sp. nov. (Figs 223-224, Plate 10A) MATERIAL. HOLOTYPE: QM0304399: Mrs Watson's Bay, midway in bay, Lizard I., Old, 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- ^ 435 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR FIG. 222. Antho (lsopenectya) 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. MEMOIRS OF THE QUEENSLAND MUSEUM 436zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ face aquiferous canals, surrounded by collapsible membraneous lip. Texture and surface characteristics. Firm, compressible, not easily torn; surface turgid in life, with distinctive ectosomal membrane, arteriallike longitudinal aquiferous canals obvious on external surface, branching and interconnecting, opening into common oscules, porous surface between canals; canals, ridges and oscules collapse 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 ascending primary, plumose spicule tracts, extending 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 subectosomal regions; 3 distinct skeletal components: (1) renieroid skeleton composed of both acanthose styles and smaller smooth principal styles in uni-, bi- or paucispicular tracts, coring small, light spongin fibres up to 2511m diameter, producing rectangular or triangular meshes up to 90Rm diameter; (2) plumose, diverging skeleton 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 embedded in peripheral skeleton perpendicular to surface; echinating megascleres absent; mesohyl matrix light, without microscleres but few whi spy (? juvenile) auxiliary subtylostyles scattered between fibre meshes; choanocyte chambers small, oval, 25-45Rm diameter. Megascleres. Smooth choanosomal principal styles long 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)-4p.m. Acanthose styles of renieroid skeleton slender, slightly curved towards base, rounded, sparsely microspined bases, sparsely spined shaft, spines small, erect, conical; points of spicules fiisiform. Length 884114.9)-153 p.m, width 2-(3.6)-6p,m. Subectosomal auxiliary subtylostyles variable in length and thickness but only comprising a single category; bases subtylote, microspined, tuberculate (granular) or occasionally smooth, points fusiform or slightly telescoped; whispy juvenile forms present scattered throughout mesohyl. Length 78-(169.8)-296Rm, width 0.5(1.6)-3Rm. Echinating spicules absent. Microscleres. Absent. ETYMOLOGY. Latin puniceus , reddish. REMARKS. The bulbous growth form, red colour and production of abundant mucus is common to many other Indo-west Pacific microcionids (such as C. (Isociella) eccentrica, C. (Thalysias) vulpina, C. (T) hirsuta, Echinoclathria axinelloides, and Echinochalina (Protophlitaspongia) bargibanti), but this species belongs to Antho (Isopenectya) having a renieroid skeleton composed (mainly) of a special category of acanthose styles (geometrically different from choanosomal spicules), a secondary, diverging, plumose skeleton of smaller, smooth choanosomal styles, and larger smooth choanosomal styles embedded in the peripheral 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 (I.) punicea differs from A. (I.) chartacea in growth form, absence of axial skeletal compression, spicule geometry and spicule sizes. The bases of auxiliary spicules in this species are also unusual, varying from swollen bases with prominent terminal 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'E, 7m depth, 16.xi.I977, 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, 1 1 mm diameter (cylindrical portions), or up to 20mm diameter, 8mm thick (lamellate portions of digits), repeatedly bifurcate, rarely anastomosing, expanding towards ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 437 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. MEMOIRS OF THE QUEENSLAND MUSEUM 438zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 224. Antho (lsopenectya) 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 439 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, compressible, flexible; surface smooth, even, unornamented, 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 subtylostyles also protruding through surface in association 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, 40-60p,m diameter, slightly more bulbous at fibre nodes, 70-90Rm diameter; all axial fibres cored by unior paucispicular tracts of acanthose principal styles forming rectangular or less often triangular meshes, 70-100Rm diameter; extra-axial fibres lighter, with differentiated primary, ascending fibres, 20-40p.m diameter, cored by paucispicular tracts of both acanthose principal styles and smaller, smooth principal styles, becoming increasingly 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, 15-30p.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 ascending primary fibres; echinating megascleres absent; mesohyl matrix heavy but only lightly pigmented, with both fully formed and raphidiform subectosomal auxiliary subtylostyles scattered between fibre meshes; choanocyte chambers small, oval, 40-50ptm diameter. 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)-156p,m, width 4-(6.9)-10p.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)-9811m, width 4.5-(5.9)-8p.m. Subectosomal auxiliary subtylostyles short or long, slender or raphidiform, straight or slightly curved at centre, prominently subtylote, smooth bases, fusiform points. Length 664115.7)198t.tm, width 1-(1.7)-3p.m. Microscleres. Absent. ETYMOLOGY. For the type locality. REMARKS. This species resembles both Antho and Echinoclathria, having a renieroid architecture and larger, smooth principal styles protruding through the surface. Like A. (I.) punicea it is included in Antho because it has a special category of acanthose styles (geometrically different from choanosomal spicules) forming the renieroid skeleton, and a secondary, diverging, plumose skeleton of smaller, smooth choanosomal styles. Antho (I.) saintvincenti differs from A. (I.) punicea in its flattened or cylindrical-digitate, arborescent growth form, reminiscent of Echitzoclathria chalinoides, although spiculation and skeletal architecture differ substantially. OTHER SPECIES OF ANTHO (ISOPENECTYA) Antho (Isopenectya) primitiva (Burton, 1935)zyxwvutsrqpo 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. Clut hr iella pr im it iva - - Echinoelathria Carter, 1885 Echinoclat hr ia Carter, 1885f: 355. Ophlit aspongia; of authors; (not Bowerbank, 1866: 14). TYPE SPECIES. Echinoclat hria t enuis Carter, 1885f: 355 (by subsequent designation offlurton, 1934a: 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 440zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 441 FIG. 226. Antho (I sopenectya) saintvincenti sp.nov. (holotype SAMTS4035). A, Choanosomal skeleton. B, Fibre characteristics (x303). C, Acanthose subtylosty le of renieroid skeleton. D, Acanthostyle spines. E, Bases of principal subtylostyles. F, Base and point of subectosomal auxiliary subtylostyles. MEMOIRS OF THE QUEENSLAND MUSEUM 442zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ styles, echinated by identical spicules (occasionally 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 perched 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 paratangentially or embedded perpendicular to surface; microscleres include toxas and palmate isochelae. REMARKS. Sixty nine species have been included 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. Echinoclathria is similar to Antho (Isopenectya), as noted above, differing in having only 2 skeletal components: a relatively homogeneous renieroid choanosomal skeleton composed of smaller, smooth principal spicules, and vestigial radial extra-axial skeleton on the external surface. Isopenectya has in addition a renieroid 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. (I.) 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 the development of the extra-fibre skeleton. In some species (e.g., E. leporina, E. confragosa) 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 (e.g., E. egena, E. waldoschmitti)there is 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 classifications for Echinoclathria are rejected here. Echinoclathria axinelloides (Dendy, 1896) (Figs 227-228, Plate 10B) Ophlitaspongia axinelloides Dendy, 1896: 39; Hallmann, 1912: 268-270, p1.36, fig.3, text-fig.58; Burton, 1934a: 599. Echinochalina axinelloides; de Laubenfels, 1936a: 119. Echinoclathriaaxinelloides; Carpay, 1986: 22; Hooper & Wiedenmayer, 1994: 279. MATERIAL. HOLOTYPE: NMVG2318 (fragment BMNH1902.10.18.342): Port Phillip, Vic, 38°09'S, 144°52'E, 36m depth, coll. J.B. Wilson (dredge). OTHER MATERIAL: VIC- AMZ802, AMZ1593. TAS- QMG300269 (NCIQ66C-3655-0) (fragment NTMZ3804). HABITAT DISTRIBUTION. Rock reef; 20-36m depth; Port Phillip Bay (Vic); Fumeaux Is (Tas) (Fig. 227D). DESCRIPTION. Shape. Erect, club-shaped or arborescent, up to 115mm long, 75mm wide, with thick subcylindrical 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 2.5R 4/10), pale brown in ethanol. Oscules. Numerous, moderately small, 2-4mm diameter, mainly on lateral margins of lamellae or on 1 side of branches, slightly raised with membraneous lip. Texture and surface characteristics. Firm, compressible, rubbery in life; surface optically smooth, minutely reticulated, with distinct membraneous covering. Ectosome and subectosome. Membraneous, with minutely reticulate, skin-like membrane stretched over surface, microscopically hispid from protruding choanosomal styles forming well developed, multispicular plumose brushes just below surface producing a more-or-less continuous palisade; surface spicule brushes heavier at surface than at core of skeleton; fewer subectosomal auxiliary styles paratangential to surface in association with oscules. Choanosome. Skeleton more-or-less dendroreticulate, 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 (75-14811m diameter) multispicular, cored by distinctly plumose tracts of choanosomal principal styles, whereas secondary fibres (38-72Rm REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO ^ 443 FIG. 227. Echinoclathria axinelloides (Dendy) (holotype NMVG23 l8). 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 anastomoses form oval, elongate or rectangular meshes, 38-195ttm diameter, more compact in peripheral skeleton, relatively cavernous towards axis (130-275 p.m diameter), fibre nodes slightly enlarged, bulbous; choanosomal fibres and MEMOIRS OF THE QUEENSLAND MUSEUM 444zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ spicule tracts become more regular and more compacted towards periphery; mesohyl matrix relatively heavy, granular, with oval to eliptical choanocyte chambers (56-216Rm 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 764109.8)-14811m, width 449.8)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)-187Rm, width 2.544.1)-5 Microscleres. Absent. Larvae. Viviparous, parenchymella larvae oval to elongate, 155-275x120-170p.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 between 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 (1934a) the Saville Kent collection contains this species, but this record is questionable as the specimens have not been discovered in the BMNH collections. Echinoclathria bergquistae sp. nov. (Figs 229-230, Plate 10C) MATERIAL. HOLOTYPE: QMG303872: S. of Triangle 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: QMGL952 (fragment NTMZ1534): E. of Murdock I., 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, 65150mm 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, up to 3mm diameter, with slightly raised membraneous lip alive, scattered on exterior surface of lobate digits. Texture and surface characteristics. Soft, compressible, fibrous, difficult to tear, produces slight, clear mucus alive (on deck), stains ethanol orange; surface highly clathrous with large, flattened lobate or pointed conules covering exterior surface of digits, 5-I5mm long, up to 5mm wide; surface porous in preserved state, membraneous alive. Ectosome and subectosome. Surface prominently hispid with longer, smooth choanosomal principal styles embedded in peripheral fibres, extending nearly 70% of their full length through surface; near bases of protruding principal styles are relatively heavy multispicular tracts of subectosomal auxiliary subtylostyles, usually tangential 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 subectosomal auxiliary subtylostyles scattered throughout mesohyl; spongin fibres large, 4060iLm diameter, well developed but only lightly invested with spongin, without any marked differences 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 445 FIG. 228. Echinoclathria arinelloides (Dendy) (holotype NMVG2318). A, Choanosomal skeleton. B, Fibre characteristics. 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 9011,m diameter in axis, 160Rm 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, 35-5511m diameter. MEMOIRS OF THE QUEENSLAND MUSEUM 446^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 714149.8)309p,m, width 2.5-(5.6)-12p.m. Subectosomal auxiliary subtylostyles long, slender, straight, subtylote, smooth or less commonly microspined bases, fusiform points; numerous smaller and raphidiform styles also scattered through mesohyl presumably being younger forms. Length 203-(356.7)-480Rm, width 243.7)-611m. Microscleres. Palmate isochelae not common, relatively large, with short thin alae, lateral alae completely fused to shaft, front ala nearly completely detached from lateral alae, shaft straight. Length 18-(24.2)-3211,m. Toxas wing-shaped, relatively thick, with slightly rounded central curvature, slightly reflexed points. Length 32-(49.7)-68p,m, width Echinoclathria chalinoides (Carter, 1885) (Figs 231-232) Axinella chalinoides Carter, 1885f: 358; Carter, 1886g: 377 [et varr glutinosa, cribrosa]. Axinella cladoflagellata Carter, 1886g: 377. Echinochalina chalinoides; de Laubenfels, 1936a: 119. Ophlitaspongia chalinoides; Dendy, 1896: 36. Echinociathria chalinoides; Hooper & Wiedenmayer, 1994: 279. Not Ophlitaspongia chalinoides; Hal!mann, 1912: 270-272, text-fig.59. MATERIAL. LECTOTYPE: BMNH1886.12. 15.401 (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: BMNH1886.12.15.407: same locality as lectotype. HABITAT DISTRIBUTION. Ecology unknown; Port Phillip (Vic) (Fig. 231D). DESCRIPTION. Shape. Arborescent branching, 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 echinating 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 echinating fibre nodes, is reminiscent of Antho (Isopenectya). 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 choanosome. up to 250mm long, 170nun maximum width, with small basal stalk up to 45mm long, 22mm diameter, long cylindrical branches up to 105mm long, 9mm diameter, slightly flattened, bifurcating repeatedly, rarely anastomosing. Colour. "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, compressible but difficult to tear, flexible branches, more rigid stalk; surface slightly microconulose, with fine surface network of radiating spicules associated with aquiferous system. Ectosotne and subectosome. 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 paratangential to surface; subectosomal region slightly cavernous, meshes up to 450p,m diameter, substantially more wider-meshed than in axial region skeleton. Choanosome. Skeletal architecture dendroreticulate, vaguely subrenieroid and more regular towards periphery than axis, composed of heavy, well developed spongin fibres, 25-7011m diameter, thicker and slightly bulbous at fibre ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 447 FIG. 229. Echinoclathria berg quistae 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. MEMOIRS OF THE QUEENSLAND MUSEUM 448zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 between 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 predominantly of longer spicules whereas towards core of skeleton coring spicules generally smaller and contained mainly within fibres; transverse connecting fibres uni-, bi- or aspicular; echinating ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO 449 FIG. 231. Echinoclathria chahnoides (Carter) (lectotype BMNH1886.12.15.401). A, Principal styles and anisostyles (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, 90-3201.1m diameter in axis; mesohyl matrix heavy but only lightly pigmented, with ovoid choanocyte chambers (90- 120Rm 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 tapering, sometimes subtylote or telescoped, with has- MEMOIRS OF THE QUEENSLAND MUSEUM ^ 450zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 modified to quasidiactinal forms (symmetrical ends). Length 176-(264.7)-325pLm, width 8(10.8)-13p.m. Subectosomal auxiliary styles, anisostyles or anisoxeas with similar geometry to principal spicules but generally longer, more slender, straight or occasionally slightly curved or sinuous, with smooth rounded bases, sometimes REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 451 telescoped, fusiform or telescoped points. Length 2434315.5)-365 pin, width 4-(5.3)-8p.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 spongin 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, p1.35, fig.2, text-fig.53. Axociella confragosa; de Laubenfels, 1936a: 113. Echinoclathria confragosa; Hooper & Wiedenmayer, 1994: 279. MATERIAL. HOLOTYPE: AMZ992 (dry): Shoalhaven 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, bifurcating and anastomosing, forming loose reticulate mass, with shaggy lobate surface projections on points of branches. Colour Grey-brown in dry state. Oscules. Not seen. Texture and surface characteristics. Firm, compressible, brittle, fibrous; surface porous, pitted, slightly arenaceous. Ectosome and subectosome. Membraneous ectosome, 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; subectosomal auxiliary styles form tangential and paratangential tracts on surface; thickness of ectosomal skeleton ranges from tangential (three spicules abreast) to paratangential (tracts of up to 20 spicules at obtuse angles to surface, forming low microconules). Choanosome. Irregularly renieroid reticulate skeleton, more-or-less homogenous throughout; spongin fibres thin, relatively light, 18-48pm diameter, without any obvious differentiation between primary or secondary components; fibres cored by uni- or paucispicular tracts of choanosomal principal styles, occasionally aspicular, echinated by choanosomal styles sparsely and irregularly dispersed over fibres; fibre meshes predominantly rectangular (=renieroid), less often oval or triangular (=isodictyal), 112345 p.m diameter, slightly more cavernous at core and more compacted in peripheral regions of skeleton; mesohyl matrix heavy but only lightly pigmented, oval choanocyte chambers 52110p,m diameter; numerous microscleres, subectosomal auxiliary styles, and also few choanosomal styles scattered between fibres. Megascleres. Choanosomal principal subtylostyles (coring and echinating fibres) short, thick, straight, with smooth, slightly constricted subtylote bases, almost hastate points, slightly rounded, telescoped or pointed. Length 142(164.8)-197p,m, width 5-(9.6)-12p.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 1364214.4)-291 pan, width 3.545.1)6p,m. 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)-14p,m. Toxas oxhorn, small, thick, with slight to moderate, evenly rounded central curvature, straight arms, slightly reflexed points. Length 32458.2)-951_1,m, width 142.2)-3.5 MEMOIRS OF THE QUEENSLAND MUSEUM 452zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 important 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 paratangential ectosomal skeleton covering most of the surface; and distinctive oxhorn toxa morphology. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 453 Echinoclathria digitata (Lendenfeld, 1888) (Figs 235-236, Plate 10D) Thalassodendron digitata 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, bulbous, expanding and slightly flattened towards apical extremities, up to 32mm long, 6mm wide, bifurcating several times, occasionally anastomosing, with 1 or more blind branches; mass growing from a semi-encrusting common base. Colour Bright red alive (Munsell 5R 4/10), greybrown in ethanol. Oscules. Small, up to 3mm diameter, on apex of each digit. Texture and surface characteristics. Firm, compressible, flexible, fibrous; surface bulbous, 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 auxiliary styles tangential to surface. Choanosome. Skeletal architecture more-or-less renieroid reticulate, with heavy spongin fibres producing wide-meshed rectangular reticulation; primary ascending fibres long, multispicular 6090iim diameter, interconnected by numerous, shorter, secondary fibres, 30-55Rm diameter cored by 1 or few principal spicules; fibres sparsely echinated by principal subtylostyles, identical to but marginally thicker than those coring fibres, confined mostly to distal margins (periphery) of fibres; fibre anastomoses produce cavernous meshes, 80-320Rm diameter; mesohyl matrix sparse in choanosome, lightly pigmented, granular, containing numerous whispy auxiliary styles and fewer microscleres; choanocyte chambers elongate 20-30Rm diameter. Megascleres. Principal subtylostyles coring and sparsely echinating fibres short, moderately thick, straight or slightly curved at centre, entirely smooth, with slightly subtylote bases, bluntened or slightly telescoped points. Length 186(214.4)-238Rm, width 4-(4.6)-711.m. 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 1824204.2)246p,m, width 1.541.9)-2.511m. Microscleres. Palmate isochelae small, unmodified, long lateral alae approximately same length as front ala, entirely fused to shaft, front alae nearly completely detached. Length 10(11.7)-13Rm. REMARKS. Lendenfeld's (1888) original material is not extant in the collections of either the AM or BMNH, but we know from his brief 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. Echinoclathria digitata is similar to E. berquistae in its digitate growth form and having a cavernous, predominantly renieroid, reticulate skeletal architecture. It differs from E. bergquistae in geometry and dimensions of all its spicules, lacking toxa microscleres, and having relatively homogeneous megascleres throughout the skeleton, coring and echinating tracts and scattered interstitially. In possessing relatively homogeneous megascleres E. digitata also resembles E. levii, although the latter has completely different skeletal architecture, growth form and toxa microscleres. Echinoclathria egena Wiedenmayer, 1989 (Figs 237-238, Plate 10E) Echinoclathria egena Wiedenmayer, 1989: 64-66 p1.6, fig.8, p1.24, fig.6, p1.25, figs 1-2, text-fig.44; Hooper & Wiedenmayer, 1994: 279. MATERIAL. HOLOTYPE: NMVF51978: Winter Cove, E. side of Deal I., Kent Group, Bass Strait, Tas, 39°29'S, 147°20'E, 26.iii.1981, 3-6m depth, coll. F. Wiedenmayer et al. (SCUBA).0THER MATERIAL: TAS- QMG300669 (NCIQ66C-3721-L) (fragment NTMZ3817). MEMOIRS OF THE QUEENSLAND MUSEUM 454^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 234. Echinoclathria confragosa (1-lallmann) (holotype AMZ992). A, Choanosomal skeleton. B, Fibre characteristics (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, Oxhom toxa. HABITAT DISTRIBUTION. On granite boulders and rock reef in sand substrate; 3-6m depth; Kent Is, E. St Patrick's Head (Tas) (Fig. 237D). DESCRIPTION. zyxwvutsrqponmlkjihgfedcbaZYXWVUT Shape. Erect, digitate, ranging from young forms thickly encrusting basal mat, up to 5mm thick, with irregularly bifurcate and REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 455 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 subterrninal processes along branch length, and with short stalk, up to 50mm long, 21rrirn diameter, and enlarged basal attachment. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 456zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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. Compressible rubbery texture, difficult to tear; surface contorted, macroscopically even but microscopically reticulate. REVISION OF MICROCIONIDAE 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, micropapillose 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 458zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM peripheral skeleton become more compacted and oval (70-90Rm 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-echinating fibres) small, thin, straight, with smooth, tapering, slightly subtylote or rounded bases, fusiform or slightly telescoped points. Length 38447.7)53p.,m, width 2.342.5)3.11.1m. Subectosomal auxiliary megascleres thin, straight, or rarely sinuous, strongylote styles (asymmetrical, anisostyles), with evenly rounded points and slightly thicker bases. Length 88-(158.2)-178p.,m, width 1-(1.5)-2.4iLm. Microscleres. Absent. Larvae.^Viviparous parenchymella larvae, 150195 im diameter, in various stages of development througout mesohyl. REMARKS. This species FIG. 238. Echinoclathria egena Wiedenmayer (QMG300669). A, Choanosomal is placed with Burton's skeleton. B, Fibre characteristics. C, Principal subtylostyle. D, Ends of principal (1959a) group of Babelspicules. E, Base of subectosomal auxiliary stongylote style. late, massive, ramose Echinoclathria sponge s elements, cored by uni- or multispicular tracts of lacking microscleres (axinelloides, chalinoides, choanosomal principal subtylostyles, fibres beinornata, leporina, nodosa; none of which are coming swollen and compressed towards synonyms contrary to Burton's (1959a: 247) periphery (27-52p.m diameter), and secondary transverse, usually aspiculose fibres (16-40iLm 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 (270-420iLm diameter), whereas meshes towards 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 characteristics, but differs from these species in spicule geometry and compression of peripheral spongin fibres. It should also be compared with E. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 459 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) Ophlitaspongia inornata Hallmann,1912: 265 268, - p1.36, fig.2, text-fig.57; de Laubenfels, 1954: 163; Coombe et al., 1987: 381; 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).0THER MATERIAL WA- WAM622-81(1) (fragment NTMZ1714). S AUST- SAMTS4055 (fragment NTMZ1656), AME938, AME768. - HABITAT DISTRIBUTION. Rock reef; 31-60m 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 lobate branches, bifurcate, expanded and bulbous at their ends, 7-25mm diameter; long cylindrical basal stalk, 12-35mm long, up to 15mm diameter, enlarged basal attachment. Colour Live colouration unknown, pale brown or yellowish grey in ethanol. Oscules. Small, 1-2mm diameter, only seen on upper portions of digits. Texture and sulface characteristics. Firm, compressible, 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 ectosome, surrounding bases of protruding principal 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, 60-90p,m diameter, bulbous, not clearly divided into primary or secondary elements; axial fibres contain only 1-2 smaller principal styles per tract, producing nearly regular renieroid skeleton, whereas fibres running longitudinally through branches (seen in cross-section in skeletal preparations) are multispicular, partially obscuring renieroid appearance of axial skeleton; axial fibre anastomoses form tight oval meshes, 40-90p,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, 30-50p.m diameter, bifurcating and anastomosing, cored by 1-3 smaller principal styles and very heavily echinated by the same spicules, particularly near surface; secondary connecting, transverse fibres 20-40p.m diameter, 1-2 spicules per tract, occasionally aspicular, also heavily echinated; extra-axial fibre meshes not bulbous as in axial region, forming large oval cavernous meshes up to 250p.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 but very heavily echinated, eventually producing plumose bundles of larger principal styles protruding through surface; mesohyl matrix heavy, granular, with few extra-fibre spicules; choanocytes large, oval, 55-90p,m diameter. Megascleres. Choanosomal principal styles (coring and echinating fibres) long or short, relatively thick, slightly curved towards basal end, rounded or faintly subtylote bases, predominantly smooth, rarely slightly microspined bases, fusiform points. Length 88-(124.2)-199p,m, width 3-(6.8)-10p,m. Subectosomal auxiliary subtylostyles long, slender, straight, slightly curved or sinuous, prominently subtyolote, smooth or microspined bases, hastate points. Length 1704205.6)235p,m, width 0.5-(1.9)-3Rm. Microscleres. Raphidiform toxas uncommon, found in only 2 of 5 specimens (presumably associated with larvae; impossible to retain intact on SEM stubs); very long, hair-like, with slight angular central curvature, straight arms, straight points. Length 205-(225.4)-238p.m, width up to 0.5pan. Associations. Three of the five known specimens of this species are covered with a zoanthid, allegedly a cornulariid (Hallmann, 1912). REMARKS. Hallmann (1912) erected this species mainly by comparison with E. leporina (as O. tenuis), remarking on their close MEMOIRS OF THE QUEENSLAND MUSEUM ^ 460zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 239. Echinoclathria inomata (Hal!mann) (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 consistently different and are distinct species. The skeleton of E. inomata 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 461 FIG. 240. Echinoclathria inornata (Hal!mann) (holotypc 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 echinating spicules are sparse. Echinoclathria inomata also has a thick palmate-digitate or lobate- digitate growth form (whereas E. leporina is thinly flabellate), principal megascleres are generally larger, and some specimens of E. inomata have thin raphidiform toxas (although these are not 462zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM consistently present and might be associated with larvae). Echinoclathria leporina (Lamarck, 1814) (Figs 241-242) Spongia leporina Lamarck, 1814: 444, 373. Echinoclathria leporina; Topsent, 1932: 101, p1.6, fig.!; de Laubenfels, 1936a: 119; de Laubenfels, 1954: 163; Wiedenmayer, 1989: 61-63, p1.6, fig.!, p1.24, figs 4-5, text-fig.42; Carpay, 1986: 24; Hooper & Wiedenmayer, 1994: 279. Ophlitaspongia leporina; Burton, 1934a: 558, 562, 599. Echinoclathria tenuis Carter, 1885f: 355. Ophlitaspongia tenuis; Dendy, 1896: 37; Hallmann, 1912: 261-265, p1.35, fig.1, text-fig.56; Topsent, 1932: 101, p1.6, fig.1; Burton, 1934a: 558, 562, 599. Not Clathria tenuis Hentschel, 1911: 377-379, textfig.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: MNHNDT567: 'Australian Seas', Peron & Lesueur collection. HOLOTYPE of E. tenuis: BMNH1886.12.15.147: 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 P. papyracea: BMNHI886.12.15.232 (dry): same locality. HOLOTYPE of A. tenuispina: BMNH1886.8.27.448 (dry) (fragment AMG3467): Westemport Bay, Vic, 38°26'S, 145°08'E. OTHER MATERIAL: S AUST- AME826. VICNMVRN1075, AMZ518, AMZ1167. NSWAMZ1642, AME820. TAS- AMZ2136, AMZ2210. HABITAT DISTRIBUTION. 5-42m depth in shallow coastal waters on rock reef substrate; known only from Australia: Coogee (NSW); Port Phillip, Westemport Bay (Vic); N. coast (Tas); Cape Martin (SA) (Fig. 241D). 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 bifurcate palmate digits growing in more than I 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 2.5R 5/8-5/10), light brown in ethanol. Oscules. Relatively small, up to 2.5mm diameter, dispersed over margins of digits, without associated subectosomal drainage canals. Texture and surface characteristics. Firm, flexible, moderately difficult to tear; surface even, without pronounced sculpturing; some with distinct radial growth lines on lamellae. Ectosome and subectosome. Microscopically hispid, with larger sizes of principal style/subtylostyle protruding through surface for up to 10011m, singly or in brushes, forming a vestigial plumose or radial extra-axial 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. Choanosome. Skeletal architecture with 3 components differentiated: (1) irregularly isodictyal, slightly compressed axis; (2) more open reticulate 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 multispicular fibres, 28-65p,m diameter, and secondary, mostly transverse uni- or paucispicular fibres, 28-36Rm 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); echinating principal spicules located predominantly on primary fibres; fibre anastomoses in axis form predominantly triangular meshes (=isodictyal), less often rectangular or oval meshes, 75-18011m 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 pigmented; choanocyte chambers ovoid, 43-112tim 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 quasioxeote, rarely microspined bases, usually with fusiform points. Length 62-(185.3)-305p,m, width 4-(9.6)-141J.m. - ^ REVISION OF MICROC1ONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO 463 FIG. 241. Echinoclathrialeporina (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, occasionally microspined, subtylote bases, hastate points. Length 148-(265.8)-321 p.m. width 1.543.1)4.41,m. Microscleres. Absent. REMARKS. This species is better known under the junior names Echinoclathria or Ophlitaspongia 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 characters 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. (I.) chartacea, the MEMOIRS OF THE QUEENSLAND MUSEUM 464zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ .FIG. 242. Echinoclathria leporina (Lamarck) (holotype MNHNDT567). 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- are homogeneous, entirely smooth shafts, and the ing the renieroid skeleton, and much larger renieroid/ isodictyal skeleton is cored and smooth choanosomal styles protruding through echinated by the same smooth principal spicules, the ectosome. Choanosomal styles in zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB E. leporina with differentiated primary and secondary fibre REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 465 structures, there is less pronounced axial compression, 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 Antho. The two species may be confused and hence care has been taken in checking all available voucher specimens to ascertain conspecificity. Echinoclathria leporina is probably widespread throughout temperate SE Australia although some published records (in the Zoological Record) are not corroborated by voucher samples and are not included in the present synonymy. Echinoclathria levii sp. nov. (Figs 243-244, Plate 10F) MATERIAL. HOLOTYPE: QMG300675 (NCIQ66C37644) (fragment NTMZ3832): Trap Reef, Bicheno, Tas., 41°51.7'S, 148°18.6'E, 30m depth, 26.ii.1990, coll. NCI (SCUBA). HABITAT DISTRIBUTION. Large boulder 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, 5mm diameter; irregular digits and small lamellae ('buds') also arising from lateral sides of fan; stalk long, thickly cylindrical, 45mm long, 9mm diameter, with expanded basal attachment. Colour Dark red-orange alive (Munsell 5R 5/10), brown in ethanol. Oscules. Large, up to 2.5mm diameter, scattered over 1 side of fan, without any obvious membraneous lip, collapsing in air. Texture and surface characteristics. Stalk stiff, lamella firm, flexible, slightly compressible; surface membraneous, optically hispid, relatively even, slightly lumpy but without any conules or other ornamentation. Ectosome and subectosome. Surface microscopically hispid, with longer principal styles protruding through surface in thick, bushy, erect brushes arising from ascending choanosomal fibres in peripheral skeleton; subectosomal auxiliary subtylostyles in sparse tangential brushes on surface; mesohyl matrix heavy but only lightly pigmented in peripheral skeleton. Choanosome. Skeleton with 2 components: irregularly renieroid renieroid reticulate and plumo-reticulate, without any marked axial com- pression but clearly differentiated axial and extraaxial regions; (1) axial skeleton more-or-less renieroid, with heavy spongin fibres divided into primary and secondary elements; primary fibres ascending, 70-90p,m diameter, cored by pauci- or multispicular tracts of generally smaller choanosomal principal styles; secondary fibres transverse, short, thinner 30-40p,m diameter, interconnecting primary fibres, cored by unispicular tracts of smaller principal styles; fibre anastomoses in axial region produce cavernous oval or elongate meshes, 150-250p,m diameter, fibre nodes heavy, slightly bulbous, containing multispicular tract ascending through longitudinal plane of lamellae; axial fibres sparsely echinated by smaller principal styles, mainly at fibre nodes; (2) extra-axial skeleton distinctly plumo-reticulate, with disctinctly different primary and secondary spongin fibre systems; primary ascending fibres very thick, up to 130p.m 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 increase towards surface; secondary, connecting fibres in extra-axial skeleton small, less than 3011m diameter, uni- or aspicular; echinating spicules in extra-axial region mostly obscured by ascending plumose spicule tracts; mesohyl matrix heavy, with numerous auxiliary megascleres (sometimes also incorporated into fibres) and numerous toxas scattered throughout; choanocyte chambers small, oval, 40-60p,m diameter. Megascleres. Choanosomal principal styles (coring 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)-341p,m, width 546.1)-7 p.m. Subectosomal auxiliary styles relatively short, slender, slightly curved at centre, with slightly subtylote, smooth bases, fusiform or hastaterounded points. Length 1544187 .8)-205p.m, width 243.1)-4.5 p,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-(14.8)-1611m. Toxas oxhorn (although smaller wing-shaped forms also present), thick, with prominent, wide, even central curvature, slightly reflexed arms, MEMOIRS OF THE QUEENSLAND MUSEUM 466^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA .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 23491)-135p.m, ETYMOLOGY. For Professor Claude Levi, MNHN width 1-(1.4)-41..m. Paris, for his contributions to Indo-Pacific sponges. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 467 FIG. 244. Echinoclathria levii sp.nov. (holotype QMG300675). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Ends of principal and auxiliary spicules. E, Oxhorn toxas. F, Palmate isochela. 468zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM REMARKS. This species is an unusual Echinoclathria having well developed extraaxial plumo-reticulate skeleton in which the thick plumose spicule tracts completely dominate the thin unispicular connecting fibres, partially obscuring the renieroid skeleton (seen only clearly at the core of the skeleton). Although the thickness 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 lamellae or surface digits (growing edges) and the latter predominant towards the basal stalk region), it is a prominent feature of skeletal structure. This species also differs from related flabellate species such as E. ridddlei sp. nov. and E. 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 lamellae). Echinoclathria nodosa Carter, 1885 (Figs 245-246) Echinoclathria nodosa Carter, 1885f: 356; Ridley & Dendy, 1887: 160; Carpay, 1986: 25; Hooper & Wiedenmayer, 1994: 280. Ophlitaspongia nodosa; Dendy, 1896: 37. Litaspongia nodosa; de Laubenfels, 1954: 162. MATERIAL. HOLOTYPE: BMNH1886.12.15.96 (fragment AMG2770): Port Phillip, Vic, 38°09'S, 144°52'E, 8m depth, coll. unknown (dredge).0THER MATERIAL: VIC- NMVRN264, NMVRN628. HABITAT DISTRIBUTION. Sand and shell grit, 838m depth; Port Phillip (Vic) (Fig. 245E). DESCRIPTION. Shape. Bulbous-digitate, up to 55mm long, 30mm maximum width, 35mm breadth, small cylindrical basal stalk 5-15mm long, lOmm diameter; with bifurcate and occasionally anastomosing, irregularly cylindrical, slightly swollen branches, up to 22mm long, 8mm diameter. Colour 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, compressible, difficult to tear; surface nodulose, granular. Ectosome and subectosome. Membraneous, 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 palisade; subectosomal auxiliary styles form sparse paratangential, tangential or occasionally erect layers below protruding skeleton of principal spicules. Choanosome. Skeleton dendro-reticulate, moreor-less renieroid and homogeneous throughout, without any axial compression and only slight differentiation between peripheral skeleton and core (the former with plumose primary tracts 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, 35-6511,m diameter, ascending, multispicular with 3-8 spicules per fibre, spicules confined to within fibres at core but become plumose in subectosomal and peripheral regions; secondary fibres uni- or paucispicular, 15-40p,m diameter, predominantly transverse; fibre nodes usually bulbous, up to 801im 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, 150-270Rm diameter, slightly more cavernous in subectosomal region than at core; mesohyl matrix light, unpigmented, with few interstitial spicules; choanocyte chambers small, oval, 15-25iLm diameter. Me gascleres. Choanosomal principal styles short, thick, straight or very slightly curved at centre, smooth, evenly rounded bases, occasionally slightly subtylote, fusiform points; sometimes modified to quasi-oxeote spicules. Length 109-(128.6)-164Rm, width 648.4)131.tm. Subectosomal auxiliary styles long, slender, straight or very slightly curved at centre, smooth rounded bases or very slightly subtylote, long tapering fusiform points. Length 1164161.1)242[Em, width 2-(3.6)-6Rm. Microscleres. Absent. REMARKS. Similar to E. thielei and E. notialis sp. nov. this species lacks noticeable size differences between principal styles coring choanosomal fibres and those protruding through ectosome. 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. axinelloides in its spicule skeleton, which becomes increasingly dense and plumose towards the surface, but this resemblance is superficial, whereas characters such as growth form, bulbous fibres, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 size of both categories of megascleres together differentiate it within the genus. Kangaroo I., SA, 35°46.9'S, 137°46.5'E, 6m depth, 31.i.1989, coll. NCI (SCUBA). 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, clubshaped, 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, ^ 470zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 246. Echinoclathria nodosa Carter (NNIVRN264). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Ends of principal and auxiliary spicules. REVISION OF MICROCIONIDAE 471 fused to adjacent digits; attached directly to substrate, no stalk or basal `holdfase. Colour Pale redbrown alive (Munsell 5R 5/6), pale brown in ethanol. Oscules. Large, 36mm diameter, on apex or subapex of digits, raised above (.1 surface with prominent 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 lmm diameter, scattered evenly over surface), superficially resembling a red-brown Acropora coral. Ectosome and subectosome. Fibrous, with tips of ascending primary fibres protruding a long way through surface, with bundles of choFIG. 247. Echinoclathria notialis sp.nov. (holotype QMG300614). A, Principal styles anosomal spicules (coring and echinating fibres). B, Subectosomal auxiliary style. C, Section through also protruding peripheral skeleton. D, Australian distribution. E, Holotype. slightly from ends of MEMOIRS OF THE QUEENSLAND MUSEUM 472zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 subectosomal auxiliary styles lying tangential, paratangential or erect on surface; auxiliary spicules heaviest between protruding fibres, presumably surrounding surface pores and collapsing upon exposure to air and preservation. Choanosome. Skeleton regularly renieroid reticulate, relatively homogenous skeletal tracts, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 473 without any differentiation between axial and extra-axial regions; spongin fibres moderately well developed, 15-25p.m diameter; fibre meshes cavernous throughout, rectangular, square or occasionally triangular, 125-250p.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 becoming 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 lightly pigmented, subectosomal auxiliary styles scattered throughout; choanocyte chambers small, round or oval, 30-50Rm diameter. Megascleres. Choanosomal principal styles (coring and echinating fibres) short, slender, straight, smooth rounded bases, fusiform or slightly telescoped points. Length 1074123.6)151p.m, width 1.5-(2.7)-4p.m. Subectosomal auxiliary styles long, slender, straight, rounded smooth bases, fusiform or prominently telescoped points. Length 123(163.2)-191p.m, width 1-(1.9)-2.5p.m. Microscleres. Absent ETYMOLOGY. Greek notos, south. REMARKS. In growth form, texture, colouration, surface sculpturing and conules E. notialis is quite different from all other Echinoclathria, yet its skeletal structure, spicule geometry and distribution of spicules are more difficult to differentiate 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). 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 NTMZ1601, QMG300473): 20km due W. of Outer Harbour, Adelaide, St. Vincent Gulf, SA, 34 0 455, 138°20'E, 23-25m depth, 23.iv.1975, coll. J. McPhalain. PARATYPE: SAMTS4097 (fragments NTMZ1649, QMG300128): same locality. HABITAT DISTRIBUTION. Gravel, rock reef; 2325m depth; St. Vincent Gulf (SA) (Fig. 249E). DESCRIPTION. Shape. Massive, subsphericalbushy or erect digitate growth forms, 75-130mm 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 lamellae and basal mass, up to 15mm long, 9mm wide, tapering at 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 sutface characteristics. Firm, compressible, not easily torn, harsh due to incorporation 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 paratangential 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, irregularly anastomosing, incompletely divided into primary and secondary fibre systems; axial skeleton with very heavy primary fibres, 90I201.Lm diameter, running longitudinally (through longitudinal sections of surface digits), ascending to apex of digits, interconnected by heavy but shorter, thinner secondary fibres, 4295p,m diameter, with slightly bulbous nodes at their junction, 155-195Rm diameter; fibre anas- MEMOIRS OF THE QUEENSLAND MUSEUM 474^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA tomoses in axial skeleton close-meshed, 60155p,m diameter; primary axial fibres cored by multispicular tracts of shorter choanosomal principal styles, secondary fibres unispicular, occasionally paucispicular; all fibres echinated by smaller principal styles; extra-axial skeleton more cavernous, fibre meshes 110-360p,m diameter, with more-or-less evenly reticulate fibres and irregularly renieroid reticulate spicule tracts; extra-axial primary fibres heavy, thick, 65-95 p.m diameter, clearly ascending and arising perpendicularly from primary fibres in axis, multispicular, with spicule tracts composed of smaller and larger principal styles diverging increasingly towards periphery; secondary connecting fibres heavy, short, mainly transverse, 18-43p,m diameter, unispicular; echinating spicules moderately common in extra-axis; mesohyl matrix heavy, darkly pigmented, containing many scattered subectosomal auxiliary styles mostly near surface; choanocyte chambers oval, 30-50p,m diameter. Megascleres. Choanosomal principal styles (coring and echinating fibres) long or short, relatively thick, slightly curved at centre, with smooth, rounded or slightly subtylote bases, fusiform points. Length 1244196.6)-312pm, width 5-(7.1)-10p,m. Subectosomal auxiliary subtylostyles thin (occasionally raphidiform), straight, slightly curved or sinuous, smooth subtylote bases, fusiform points. Length 123-(238.4)-355p,m, width 1.5(2.9)-4.5 p.m. Microscleres. Chelae absent. Toxas accolada, moderately short, thin, with slightly rounded central curvature, straight arms, straight points. Length 63-(129.4)-175 p.m, width ETYMOLOGY. For the late Dr Shane Parker of the SA Museum, bryozoologist, sponge enthusiast, ornithologist 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 is irregularly digitate, lacks the specialised surface sculpturing and porous reticulation peculiar to the prominently bulbous E. notialis. Its 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 unispicular and renieroid in construction whereas in E. subhispida secondary fibres are aspicular. The two species differ in growth form, fibre thickness, spicule geometry and size. Only one other species (E. inornata) has toxas that approach the accolada geometry of E. parkeri. In E. inomata 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 architecture 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 11B) MATERIAL. HOLOTYPE: QMG305005 (NCIQ66C3637-1) (fragment NTMZ3801): Channel between Chappell I. and Badger I., Furneaux Is, Bass Strait, 40 16.8S, 147°54.4'E, 15m depth, 22.11.1990, coll. M. Riddle, NCI (SCUBA). PARATYPE: QMG300271 (NCIQ66C-3752-T) (fragment NTMZ3827): Trap Reef, Bicheno, E. coast Tas., 41°51.7'S, 148°18.6'E, 30m depth, 27.ii.1990, coll. NCI. OTHER MATERIAL: TAS- QMG300664 (NCIQ66C-3556H) (fragment NTMZ3781). 0 HABITAT DISTRIBUTION. Rock reef, 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 attachment; fan very thin walled, 1-4mm thick, flat, growing face-on to current, or convoluted growing in several planes; margins of fan digitate, palmate-digitate or heavily convoluted, never even. Colour Red or orange-brown alive (Munsell 2.5R 5/10 - SYR 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 sculpturing, or with only faint ridges on surface near margins of convoluted fan; surface porous in preserved state, collagenous alive. Ectosome and subectosome. Ectosome microscopically hispid with protruding large ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 475 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 300vm from surface, surrounded by irregular bundles of smaller thin subtylostyles lying tangential, paratangential or rarely forming brushes on ectosome; mesohyl matrix light in subectosomal region, ectosome membraneous.zyxwvutsrqpon Choanosom e. Skeletal architecture renieroid reticulate, without any division between axial or peripheral regions except for larger principal 476^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 250. Echinoclathria parkeri sp.nov. (holotype SAMTS4091). A, Choanosomal skeleton. B, Fibre characteristics. C-D, Ends of principal and auxiliary styles. E, Accolada toxas. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 477 spicules protruding through surface; spongin fibres light, imperfectly divided into primary and secondary elements; primary fibres ascending, 30-401im diameter, cored by pauci- or multispicular tracts of smaller subectosomal auxiliary subtylostyles; secondary fibres connecting primary elements, mainly transverse, 10-2011m diameter, cored by uni- or paucispicular tracts of smaller principal spicules; primary and secondary fibre anastomoses form rectangular, irregular, or occasionally triangular meshes, 40-8011m 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, 15-45 fun diameter. Megascleres. Choanosomal principal subtylostyles 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 844146.4)34811m, width 4-(6.8)10-p.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)-2821J,m, width 141.9)-3p.m. Microscleres. Absent. ETYMOLOGY. Named for Dr Martin Riddle for collecting this and many other undescribed sponge species throughout Australasia during the AIMS NCI contract 1984-1991, and providing the author with many opportunities to collect from remote sites throughout the Indo-west Pacific. REMARKS. This species has a very thinly flabellate growth form and renieroid skeletal structure similar to Antho (I.) 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 terminal ascending tracts, and the entirely spined styles forming extra-axial renieroid skeletal tract; principal styles are longer and thicker, with evenly 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 comparison, E. riddlei sp. nov. has its principal styles with an entirely smooth shaft, bases are smooth or microspined (and spines are morphologically quite different from those in A. chartacea), bases of principal spicules are subtylote, the longer principal styles are embedded only in external surface (i.e., there is no secondary, longitudinal 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, p1.36, fig.!, text-figs 5455. 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: BMNH 1886.12.15.45 (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, AMZI 142, AMZ1144, AMZ1763. S AUSTQMG300240 (NCIQ66C-2194-Z) (fragment NTMZ3534). MEMOIRS OF THE QUEENSLAND MUSEUM 478zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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. 253H). DESCRIPTION. Shape. Erect, digitate, up to 220mm high, 90mm maximum width; digits long, whip-like, thinly cylindrical or slightly flattened, up to 120mm long, 12mm maximum thickness, 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 redbrown 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 irregular in shape. Texture and surface characteristics. Digits compressible, rubbery, flexible; surface optically smooth, even except for microscopic conules ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 479 FIG. 252. Echinoclathria riddlei sp.nov. (paratype QMG300271). A, Choanosomal skeleton. B, Fibre characteristics. 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 terminal fibres, and fibres themselves also protrude through surface; few subectosomal auxiliary styles scattered tangential or paratangential to ectosome. Choanosome. Skeleton regularly dendro-reticulate, composed of heavy spongin fibres more compressed in axial than extra-axial regions; axis irregularly reticulate, heavy fibres incompletely differentiated into primary and secondary elements, 46-182p.m diameter, with markedly bulbous axial fibre nodes; extra-axial skeleton nearly 480zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM perfectly regular, dendro-reticulate, with clearly differentiated primary ascending, radial, paucispicular fibres branching at outer surface (30-621i.m diameter), interconnected at regular intervals by thinner transverse, aspicular or rarely unispicular secondary fibres (18-53p.m diameter); primary fibres both cored and echinated by choanosomal principal styles, slightly smaller than those at the surface; echinating spicules marginally more abundant on peripheral fibres; fibre anastomoses form oval or elongate meshes in axis (160-295p.m diameter), square, rectangular or oval meshes towards periphery (125-290p.m diameter); mesohy 1 matrix heavy but only lightly pigmented, large ovoid choanocyte chambers 75-110p.m diameter, numerous loose choanosomal styles and fewer subectosomal auxiliary megascleres scattered between 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)-168p.m, width 4-(5.4)-7Rm. Subectosomal auxiliary subtylostyles long, thin, straight or slightly curved at centre, with smooth subtylote bases, fusiform points. Length 89-(183.1)-235p.m, width 1-(2.2)-311sn. Microscleres. 1sochelae 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)-95p.m, width 0.841.2)-1.5 Larvae. Viviparous, spherical or elongate parenchymella larvae, 180-280x 150-210p.m, heavily pigmented, and with toxas and juvenile styles dispersed. REMARKS. AMZ948 of Whitelegge (1902b) and Hallmann (1912), was labelled `Ceraochalina multiformis var. dura Lendenfeld (no. 332)', but is now labelled (in Hallmann's handwriting) 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 Chalmers, 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 (18876) descriptions. The specimen Thalassodendron viminalis Lendenfeld, no.365 missing from AM (pers. obs.) was rejected as type 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. intennedia are given below under E. intermedia. 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-orless 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 E. 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 welldeveloped 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-differentiated axial and extra-axial fibres), has virtually 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, multispicular spicule tracts whereas transverse fibres are often aspicular in E. subhispida). OTHER SPECIES OF ECHINOCLATHRIA Echinoclathria arborea (Tanita, 1968)zyxwvutsrqponmlkjihg Tanita, 1968: 49-50, p1.1, fig.7, text-fig.9 [Ariake Sea]. Not Echinoclat hria arborea; Hallmann, 1912: 202. Lit aspongia ar bor ea MATERIAL. HOLOTYPE: MMBS. Japan. Echinoclathria arcifera (Schmidt, 1880) ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 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. 482zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM Tenacia arcifera Schmidt, 1880: 81, p1.10 [Gulf of Mexico]; Hallmann, 1920: 771. Ophlitaspongia arcifera; Burton, 1959a: 247 [key to species]. MATERIAL. HOLOTYPE: LMJG. Caribbean. Echinoclathria atlantica Sara, 1978 Echinoclathria atlantica Sara, 1978: 76-79, text-figs 49-51 [Cape Domingo, Tierra del Fuego]; Desqueyroux-Faundez & Moyano, 1987:50 [Tierra del Fuego, Argentina]. MATERIAL. HOLOTYPE: IZUG34, fragment MNHNDCL2739. SW. Atlantic. del Fuego]; Desqueyroux-Faundez & Moyano, 1987: 50 [Tierra del Fuego, Argentina]. MATERIAL. HOLOTYPE: IZUG124, fragmentMNHNDCL2750. SW. Atlantic. Echinoclathria noto (Tanita, 1963) Ophlitaspongia now Tanita, 1963: 124-125, p1.4, fig.3, text-fig.3 [Sea of Japan]; Tanita, 1964: 17-18, 21, p1.1, 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 beringensis (Hentschel, 1929) Echinoclathria oxeata(Bergquist & Fromont, 1988) Phakellia papyracea, in part; Lambe, 1892: 76 [Alas- Ophlitaspongia oxeata Bergquist & Fromont, 1988: ka]. Phakellia beringensis Hentschel, 1929: 975. Echinoclathria beringensis; de Laubenfels, 1954: 164 113, p1.52, figs c-e [New Zealand]; Dawson, 1993: 38 [index to fauna]. MATERIAL. HOLOTYPE: NMNZPOR117. New Zealand. [note]. Echinoclathria schmitti de Laubenfels, 1942: 264 [Baffin Bay, Arctic]. MATERIAL. HOLOTYPE: USNM. Arctic. Echinoclathria contexta Sara, 1978 Echinoclathria contexta Sara, 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 (Levi, 1963) Ophlitaspongia dichotoma Levi, 1963: 59, p1.9G, text- fig.68 [South Africa]. MATERIAL. HOLOTYPE: MNHNDCL615. South Africa. Echinoclathria digitiformis (Row, 1911) Ophlitaspongia digitiformis Row, 1911: 351-353, p1.37, figs 14-15 [Shab-ul-Shubak, Red Sea]. Echinochalina digitiformis; de Laubenfels, 1936a: 119 [note]. MATERIAL. HOLOTYPE: BMNH1912.2.164. Red Sea. Echinoclathria hjorti Amesen, 1920 Echinoclathria hjorti Arnesen, 1920: 21-22, p1.2, fig.5, p1.5, fig.3 [off Cape Bojador, Sahara, N. Atlantic]. Axociella hjorti; de Laubenfels, 1936a: 119 [note]. Ophlitaspongia hjorti; Burton, 1959b: 43-44 [Iceland]. MATERIAL. HOLOTYPE: ZMUU. NW. Africa, NE. Atlan- Echinoclathria reticulata(Bergquist & Fromont, 1988) Ophlitaspongia reticulata Bergquist & Fromont, 1988: 113-114, p1.52, fig. f, p1.53, 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, text-figs 20-22 [Tyrrhenian Sea]. MATERIAL. HOLOTYPE: BMNH1977.7.6.9. Mediterranean. 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, p1.80 [Shark Bay, WA]. Terpiosella fucoides; Burton, 1930b: 675. MATERIAL. HOLOTYPE: BMNH1877.5.21.8. Note: referred to Suberitidae. tic. Echinoclathria minor (Burton, 1959) Ophlitaspongia minor Burton, I959a: 246, text-fig.27 [Arabian Sea; key to species]; Sim & Kim, 1988:24, p1.1, figs 1-2 [S. Korea, doubtful conspecificity]. MATERIAL. HOLOTYPE: BMNHI936.3.4. 609. Arabian Gulf. Echinoclathria mutans (Sara, 1978) Ophlitaspongia mutans Sara, 1978: 73-76, text-figs 47-48 [Cape San Sebastiano, Cape Domingo, Tierra Holopsamma Carter, 1885 Holopsamma Carter, 1885c: 211 Plectispa Lendenfeld, 1888: 225. Aulena; in part, Lendenfeld, 1888: 228; (not Lenden- feld, 1885c: 309). [Ha/me] Lendenfeld 1885c: 285 (preocc.). TYPE SPECIES. Holopsamma crassa Carter, 1885c: 211 (by subsequent designation of de Laubenfels, 1936a: 98). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 483 FIG. 254. Echinoclathria subhispida Carter (holotype BMNH1886.12.15.70). 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 form of tightly anastomosing flattened fibrebranches (lacunae); choanosomal skeleton simply reticulate, without any axial compression or differentiation between axial and extra-axial skeletons; principal spicules core and echinate fibres, those inside fibres sometimes partially or completely replaced by detritus although those echinating fibres usually always present. MEMOIRS OF THE QUEENSLAND MUSEUM 484^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 Ha/me have been used interchangeably for all have been confused throughout the literature and species. Holopsamma is separated from EchinoEchinoclathria, Ophlitaspongia, Plectispa, clathria by its honeycombed reticulate growth ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 485 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 (i.e., 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. MEMOIRS OF THE QUEENSLAND MUSEUM 486^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Holopsamma arborea (Lendenfeld, 1888) (Figs 255-256, Plate 11C) Plectispa arborea L,endenfeld, 1888: 226. Echinoclathria arborea; Hallmann, 1912: 202, 280282, p1.29, fig.2, text-fig.63. Holopsamma arborea; Hooper& Wiedenmayer, 1994: 281. Not Clathria (Plectispa) arborea; Whitelegge, 1901: 65, 88, 89, p1.11, figs 15-15a; Hallmann, 1912: 211 [nomen nudum]. Not Litaspongia arborea Tanita, 1968: 49. Not Thalassodendron reticulata Lendenfeld, 1888: 227; Whitelegge, 1902a: 212. Echinoclathria elegans; Whitelegge, 1901: 90. Not Plectispa elegans Lendenfeld, 1888: 226. Not Clathria multipes Whitelegge, 1901: 88. ? Echinoclathria laxa; de Laubenfels, 1954: 164. MATERIAL. HOLOTYPE: AMZ946/G9160 (dry): I Ilawan a, NSW, 34°32'S, 150°50'E, other details unknown (old specimen label 'Halme alborea, type or cotype', new label `Echinoclathria arborea Lendf. ? type').0THER MATERIAL: NSW- QMG303756, QMG303760, QMG303797, QMG303808. VICQMG311287 (NCIQ66C-3499-T) (fragment NTMZ3772). TAS- QMG311450 (NCIQ66C-3759A) (fragment NTMZ3826). S AUST- QMG300239 (NCIQ66C-2148-Z) (fragment NTMZ3526). WAQMG300192 (fragment NTMZ2907), NTMZ3368. - HABITAT DISTRIBUTION. Widely distributed in temperate coastal waters usually on rocky reefs; shallow subtida1-50m depth; Illawarra, N. & S. Sydney (NSW); Westernport Bay (Vic), Bicheno (Tas), Kangaroo I. (SA), Houtman Abrolhos, Barrow I. (WA) (Fig. 255D). DESCRIPTION. Shape. Arborescent, cavernousreticulate, 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 anastomose; branches 120-330mm long, 15-38mm diameter, often tapering towards ends. Colour White or off-white alive, grey or greybrown in ethanol. Oscules. Large, up to 7mm diameter, recessed between honeycomb surface meshes, surrounded by membraneous lip. Texture and surface characteristics. Firm, compressible, flexible branches, slightly harsh surface; surface regularly or irregularly honeycomb reticulate, with fibre-branches (lacunae) forming close hexagonal meshes, 8-22mm diameter; lacunae ridge-like, projecting up to lOmm above surface, with thin membrane stretched between adjacent lacunae. Ectosome and subectosome. Ectosome membraneous, collagenous, with few subectosomal auxiliary megascleres lying tangential or paratangential to surface, and single or paucispicular plumose brushes of principal styles protruding through surface; subectosomal skeleton undifferentiated from choanosomal skeleton, immediately below surface Choanosome. Skeletal architecture regularly reticulate, with relatively light, small spongin fibres differentiated into ascending primary, pauci- or multispicular (30-52iim diameter), and transverse bi-, uni- or rarely aspicular connecting elements (12-22Rm diameter); fibre anastomoses form oval to elongate meshes, 60-14011m diameter; fibres cored and echinated by choanosomal principal styles, and echinating spicules most common on peripheral fibres; mesohyl matrix light, virtually unpigmented, with few scattered subectosomal auxiliary spicules between fibres; choanocyte chambers small, oval, 22-35Rm diameter. Megascleres. Choanosomal principal styles or subtylostyles short, thick, with tapering or slightly subtylote bases, typically smooth, sometimes vestigially microspined bases, points tapering fusiform. Length 78-(104.8)-118p.m, width 4(5.6)-7.5Rm. Subectosomal auxiliary megascleres thin, straight or slightly curved at centre, rounded smooth bases and telescoped hastate-pointed or rounded strongylote points sometimes resembling anisostrongyles. Length 1094133.2)14311m, width 1.5-(2.2)-2.5Rm. Microscle res. Absent. REMARKS. There is only a single type specimen in the AM collections, despite Hallmann's (1912: 282) remarks to the contrary, and I 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 H. arborea were found in the AM collections although there are several specimens labelled Clathria or Plectispaarborea that Hallmann may have been referring to, but these do not belong here (see remarks for C. multipes). 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 in most other species the surface reticulation is generally wider and the lacunae thicker). This species is closest to H. ramosa and H. macropora, with comparable REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 487 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, p1.8, figs 1,2, 5, 6, 22, 23, p1.9, figs 5,9. Echinoclathria crassa; Hallmann, 1912: 287. Not Antherochalina crassa Lendenfeld, 1887b: 787. Ha/me micropora Lendenfeld, 1885c: 303; Lenden- feld, 1889a: 461, p1.9, fig.12, p1.11, fig.4. Ha/me globosa Lendenfeld, 1885c: 303; Lendenfeld, 1889a: 456-457, p1.9, fig. 11. Echinoclathria globosa; Carpay, 1986: 24; Wieden- mayer, 1989: 63-64, p1.6, 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: QLDQMG303235. S AUST- QMG301379, SAMTS4082 (fragment NTMZ1678), SAMTS4085 (fragments QMG300472, NTMZ1599), SAMTS4057 (fragment NTMZ1664), QMG310777 (NCIQ66C-2143-U) (fragment NTMZ3524), QMG310804 (NCIQ66C2192-X) (fragment NTMZ3532). HABITAT DISTRIBUTION. Rock reef, sand and kelp bed substrata; 2-38m depth; Stradbroke I., Moreton Bay (SEQ); Byron Bay, Illawarra, (NSW); Port Phillip Bay, Westemport Bay (Vic); Bass Strait (Tas); Investigator Strait, St. Vincent Gulf, Kangaroo I., Port Noarlunga, Great Australian Bight (SA) (Fig. 257E). DESCRIPTION. Shape. Massive, subcylindrical, or lobate-digitate honeycomb reticulate sponge, 75-150mm diameter, with enlarged, flattened, 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 vanes according to amount of detritus in skeleton, ranging from flexible, soft, compressible, to distinctly brittle, arenaceous; surface reticulate, lacunose, consisting of irregularly meandering ridges forming large meshes producing characteristic 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 200p.m 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 containing foreign particles and some auxiliary megascleres scattered between fibres; mesohyl cavernous in places, with cavities up to 260p.m diameter, and elongate choanocyte chambers 80120p,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)-81pm, width 1.2-(4.1)-6p.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)261pm, width 1.5-(3.9)-6pm. 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 8411.6)15pm. Toxas absent. Larvae. Viviparous parenchymella larvae spherical or subspherical, 320-410pm diameter, with vestigial quasidiactinal megascleres scattered throughout mesohyl. REMARKS. Under Echinoclathria globosa Wiedenmayer (1989) provided a detailed dicussion of additional type specimens of each nominal species he examined (whereas only the specimens that have a direct bearing upon the 488zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 257. Holopsamma crassa Carter (lectotype BMNH1886.12.15.313). A, Subectosomal auxiliary subtylostyles/ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 489 FIG. 258. Holopsamma crassa Carter (QMG303235). 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- ^ MEMOIRS OF THE QUEENSLAND MUSEUM 490zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 external morphology (superficially resembling a faviid coral), texture (close to Xestospongia exigua; Petrosiidae), and skeletal characteristics (superficially resembling the arenaceous Teratose' sponges), but variability in these features 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 obvious components of the sponge's spiculation). SAMTS4085 had very heavy spongin fibres, heavily arenaceous fibre core, and a heavy, darkly pigmented mesohyl matrix superficially resembling a verongid sponge. Both these specimens had fine, sharp ridges forming the external honeycomb reticulation, with a bryozoan lacelike external reticulation in the preserved state. Both these specimens also had very few indigenous megascleres, and were initially confused 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. laminaefavosa, and it is conceivable that they belong to a single species. They are retained here as distinct species in order to highlight their differences and not submerge them, although additional non-skeletal characters are needed to confirm or refute the validity of these taxa. Holopsamma crassa has a brownish live colouration; the lacunae forming its external 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 is 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 (strongyles, subtylotes and sigmas). Holopsamma elegans (Lendenfeld, 1888) (Figs 259-260) Plectispa elegans Lendenfeld,1888: 226. Echinoclathria elegans; Hallmann, 1912: 203. Holopsamma elegans; Hooper & Wiedenmayer, 1994: 282. Not Echinoclathria elegans; Whitelegge, 1901: 90. Not 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 5mm 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 surface, lacunae smooth. Ectosome and subectosome. Membraneous, minutely hispid, with abundant subectosomal auxiliary subtylostyles erect on surface in plumose brushes; choanosomal fibres immediately subectosomal. Choanosome. Irregularly reticulate skeletal structure, with differentiated multispicular primary, ascending fibres, up to 200p.rn diameter, and numerous transverse, pauci- or aspicular secondary fibres, up to 70p,m diameter; echinating principal styles sparse, scattered only over ascending fibres; spongin fibres heavy; fibre meshes rounded, 30-150itm diameter; choanocyte chambers not seen (poor dry material). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 491 Megascleres. Choanosomal principal styles long, robust, straight or very slightly curved at centre, evenly rounded, entirely smooth bases, fusiform or slightly telescoped points. Length 78490.3)105p,m, width 4-(4.6)-61.m. Subectosomal auxiliary subtylostyles long, slender, slightly curved near basal end, subtylote smooth bases, tapering rounded or quasi-strongylote points. Length 90-(141.2)-255p.m, width Microscleres. Absent. REMARKS. This species is known only from the poor, dry holotype from Port Jackson. It is possibly a synonym of H. arborea, based on sitniliarities 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, p1.31, figs 4-5; Dendy, 1896: 40; Hallmann, 1912: 276-277, text-fig.61; Wiedenmayer, 1989: 59-60, p1.5, fig.10, p1.24, fig.2, textfig.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, p1.1, fig.3. Not Spongia cellulosa Esper,1797. Echinoclathria carteri Ridley & Dendy, 1886: 476; Ridley & Dendy, 1887: 162, p1.29, fig.12, p1.31, fig.3; Hamann, 1912: 284-285, text-fig.65. Holopsamma carte ri; 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 AUSTRALIA - NTMZ3211, NTMZ3226, 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, Westemport Bay (Vic); King I., 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, usually 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 substrate, without basal stalk or expanded point of attachment. Colour External surface usually red-orange or orange-brown in life (Munsell 7.5YR 7/10-12), paler orange below surface, deeper water samples honey-brown, in ethanol specimens turn yellowbrown or dark brown. Oscules. Small, no more than 2mm diameter, within meshes of surface reticulation. Texture and surface characteristics. Soft, compressible, rubbery, difficult to tear; regular, closemeshed honeycomb reticulate surface; meshes (alveoles) small, rounded or elongate formed by surface lacunae, up to 4mm diameter, more-orless 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 producing hispid surface, and paucispicular bundles or single auxiliary subtylostyles lying paratangential 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 elements; primary fibres 35-60p,m diameter, cored by pauci- or multispicular, plumose or oblique tracts of principal styles and small quantities of foreign detritus; secondary fibres up to 20p.m diameter, unispicular or aspiculose; fibres sparsely echinated by principal styles evenly dispersed throughout skeleton; fibre anastomoses form round to polygonal meshes, 20-100p.,m diameter; 492zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM A 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, 35551.1,m diameter. Megascleres. Principal styles coring and echinating fibres straight or faintly curved at centre, relatively thick, entirely smooth, with rounded, tapering or occasionally anisoxeote bases, fusiform points. Length 69-(96.4)-132p,m, width 244.8)-7 tun. Subectosomal auxiliary subtylostyles long, thin, mostly straight, with slightly constricted bases, usually hastate, slightly rounded or slighty telescoped points, sometimes subtylostrongylote with barely differentiated ends. Length 132-(l20.7)-17411m, width 1.4-(l.9)-2.111m. Microscleres. ^Palmate isochelae sparse, small, with long lateral alae entirely fused to straight shaft, front ala shorter, often with expanded, spoon-shaped alae. Length 8-( Toxas absent. REMARKS. Holopsamma favus has a red-orange external live colouration (most species of Holopsamma are pale or white), a cartilaginous, algae-like, contort lobate-digitate habit (most are globular or evenly cylindical-digitate), FIG. 259. Holopsamma elegans (Lendenfeld) (holotype BMNH1887.4.27.98). palmate isochelae (most A, Principal style (coring and echinating fibres). B, Subectosomal auxiliary lacking chelae), skeletal arsubtylostyle. C, Section through peripheral skeleton. D, Known Australian chitecture consisting of distribution. E, Holotype. slightly differentiated as- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 494zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 a 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 by Dendy, Hallmann, Whitelegge, Carpay and isochelae microscleres (Wiedenmayer, 1989), making confident identification difficult. Never- others has yet been examined their descriptions 496zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM match fairly closely those of the type specimens to 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, 1885 (Figs 263-264, Plate 11E) Holopsamma laminaefavosa Carter, 1885c: 212; Hooper & Wiedenmayer, 1994: 282. Echinoclathria laminaefavosa; Wiedenmayer, 1989: 60-61, p1.5, fig. 11, text-fig.41; Carpay, 1986: 24. Ha/me nidus-vesparum Lendenfeld, 1885c: 288, pl. 26, figs 1-2, p1.27, fig.4-5,7, p1.28, figs 8-9,11, p1.29, figs 12-13; Lendenfeld, 1888: 157; Lendenfeld, 1889a: 457, p1.11, figs 1, 7, 9-10, 15-18; Whitelegge, 1889: 184. Ha/me gigantea Lendenfeld,1886a: 847-9 [et vanr intermedia, macropora, micropora]. Aulena gigantea; Lendenfeld, 1888: 230-232 [et varr intermedia, macropora]; Lendenfeld, 1889a: 97, p1.8, figs 3,4, 7, 8, 18, 29, p1.9, figs 2-4; Whitelegge, 1889: 187; Whitelegge, 1901: 93, 118; Whitelegge, 1907: 504; Hallmann, 1912: 268, 285-287; Burton, 1938a: 20; Guiler, 1950: 10. Not Ha/me micropora Lendenfeld, 1885c: 303; Lendenfeld, 1889a: 461, p1.9, fig.12. p1.11, fig.4. Ha/me laxa Lendenfeld, 1886a: 845-7 [ et van digitata, minima]. Aulena laxa; Lendenfeld, 1888: 228-230; Lendenfeld, 1889a: 95-97, p1.8, figs 10-12, 15,20-21, p1.9, fig.1; Whitelegge, 1889: 187; Hallmann, 1912: 287; Hallmann, 1914a: 268; Burton, 1938a: 20. ? Aulena laxa; Dragnewitsch, 1906: 442. Echinoclathria laxa; Hallmann, 1912: 287. Halme irregularis Lendenfeld, 1889a: 49, 453-5, p1.9, fig.8, p1.11, fig.3 [et van micropora, lamellosa]. Echinoclathria favus var. arerufera Carter,1885f: 350. Echinoclathria arenifera; Dendy, 1896: 40-41; Hallmann, 1912: 287; Wiedenmayer, 1989: 60-61. cf. Plectispa arborea Lendenfeld, 1888: 226; de Laubenfels, 1954: 164. MATERIAL. LECTOTYPE: BMNH1886.12.15.312 (dry) (fragments BMNH1886.12.15.311, AMG2879): Port Phillip, Vic, 38°09'S, 144°52'E, coll. J.B. Wilson (dredge). PARALECTOTYPES: BMNH1886.12.15.420 (dry): same locality. BMNH1886.12.15.490 (dry): same locality. BMNH1886.12.15.49 (dry) (fragment AMG2875): same locality. BMNH1886.12.15.419 (dry): same locality. LECTOTYPE of H. nidusvesparum: BMNH 1886.8.27.73 (fragment BMNH1886.8.27.74): Port Jackson, NSW, 33°51'S, 151°16'E. PARALECTOTYPE of H. nidusvesparum: AMG8899: same locality. HOLOTYPE of H. laxa: BMNH1886.12.13.24: Port Jackson, NSW, 33°51'S, 151°16'E. LECTOTYPE of H. laxa digitata: BMNH1886.8.27.266 (fragment AMG3652): Port Jackson, NSW, 33°51'S, 151°16'E (label Psamella digitata'). PARALECTOTYPES of H. laxa digitata: AMZ121-124 (dry) (fragments AMG3759, ZMB6396): same locality. AMZ120: same locality. AMG9125 (dry): same locality. LECTOTYPE of H. laxa minima: BMNH1886.8.27.75 (fragments BMNH1886.8.27.76-77): Port Jackson, NSW, 33°51'S, 151°16'E. PARALECTOTYPES of H. laxa minima: AMZ125: same locality. AMG9157 (dry) (fragment AMG3780): same locality. HOLOTYPE of H. gigantea: BMNH1886.12. 13.19: 'E. coast of Australia'. LECTOTYPE of H. gigan tea intermedia: BMNH1886.8.27.91 (fragment BMN111886.8.27.92): Broughton I., NSW, 32°36'S, 152°19'E. PARALECTOTYPES of H. gigantea intermedia: BMNH1886.8.27.79 (dry) (fragments BMNH1886.8.27.80, AMG3762, ZMB1177): Port Jackson, NSW, 33°51'S, 151°16'E. AMA5544 (dry): same locality. AMZI 17: same locality. LECTOTYPE of H. gigan tea macropora: BMNH1886.8.27.85 (fragment BMNH1886.8.27.86): Port Jackson, NSW, 33°51'S, 151°16'E. PARALECTOTYPES of H. gigantea macropora: BMNH1886.8.27.83 (dry) (fragments BMNH1886.8.27.8 4, AMZ714, AMG3760, AMG2878): same locality. AMG9158 (dry): same locality. BMNH1886.12.15.312 (dry): same locality. HOLOTYPE of H. gigantea micropora: BMNH1886.8.27.81 (dry) (fragments BMNH 1886.8.27.82, AMG3761): Illawarra, NSW, 34 0 335, 150 ° 55'E. LECTOTYPE of H. irregularis micropora: BMNH1886.2.15.491 (fragment AMG2875): Port Phillip, Vic, 38°09'S, 144°52'E. PARALECTOTYPE of H. irregularis micropora: BMNH1886.12.15.504 (fragment AMG2874): same locality. LECTOTYPE of H. irregularis larnellosa: BMNH1886.12.15.490 (dry) (fragment AMG2876): Port Phillip, Vic, 38°09'S, 144°52'E. PARALECTOTYPES of H. irregularis lamellosa: BMNH1886.12.15.419, 420 (dry): same locality. OTHER MATERIAL: QLD- QMG303957. NSWQMG301399, QMG301441, AMZ2225. VICNCIQ66C-3402-0 (fragment NTMZ3852). HABITAT DISTRIBUTION. Rocky reefs, sand and shell grit substrata; 3-80m depth; Bass Strait, Maria I., Blackman's Bay (Tas); Port Phillip Bay, Westernport Bay (Vic); Byron Bay, Jervis Bay, Port Jackson, Port Stephens, Maroubra, Barranjoey, Illawarra, Broughton I., Sandon Bluffs (NSW); Noosa Heads, Tweed River (SEQ) (Fig. 263D). The reported localities of Torres Strait (FNQ) (Lendenfeld, 1888), and Palua Brani, Singapore (Dragnewitsch, 1906) are suspect. DESCRIPTION. zyxwvutsrqponmlkjihgfedcbaZYX Shape. Massive, lobate, cylindrical-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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 497 0 FIG. 263. Holopsamma laminaefavosa Carter (holotypc BMNH1886.12.15.312). A, Principal style (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle/ style. C, Section through peripheral skeleton. D, distribution. E, Holotype. F, QM6301441. 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 typically widely spaced, 4-13mm apart, cavities formed by meshes relatively shallow, covered by thick tympanised membrane stretched across adjacent ridges. Ectosome and subectosome. Membraneous, heavily arenaceous, without a well defined tangential spicule skeleton, but often with points of ascending principal styles protruding through surface in sparse bundles, below which are scattered individual auxiliary spicules; subectosomal MEMOIRS OF THE QUEENSLAND MUSEUM 498zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ diameter, without any consistent size differences between meshes at core or periphery; primary fibres usually cored by detritus and sparse tracts of choanosomal principal styles, and echinated by pauci- or multispicular tracts of same principal styles; coring and echinating spicules together produce plumose or oblique, vaguely ascending tracts, usually heaviest on peripheral fibres; principal styles may be absent entirely 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 usually darkly pigmented, usually with subectosomal auxiliary megascleres dispersed throughout; choanocyte chambers oval to elongate, 50-90p.m diameter, often obscured by detritus. Megascleres. Choanosomal principal styles coring and echinating fibres short, straight, relatively thick, with tapering (hastate) or rounded smooth bases, telescoped or occasionally fusiform points. Length 52-(86.4)-115p.m, width 2-(4.1)-7p.m. Subectosomal auxiliary FIG. 264. Holopsamma laminaefavosa Carter (QMG303957). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal styles. D, Ends of principal style. megascleres strongylte E, Auxiliary styles. F, Ends of auxiliary spicule. styles, long, thin, straight or flexuous, with rounded or region undifferentiated from choanosome, fibres slightly subtylote bases, rounded, telescoped or sometimes hastate points. Length 1284174.3)immediately subectosomal. Choanosome. Skeleton irregularly reticulate, 195p.m, width 1-(2.8)-4p.m. with very heavy spongin fibres forming radiating Microscleres. Absent. lamellae, fibres radiating and diverging slightly towards periphery, and more-or-less differen- REMARKS. This species is similar to H. crassa tiated into primary ascending and secondary in growth form, skeletal structure and more-ortransverse components; fibre anastomoses form less in spiculation, although both species may close or open meshed reticulations, 40-730p,m show considerable variation between regional REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 499 populations in these features; they are also sympatric although they have not yet been found on the same reefs (present study). This polymorphism has been described in detail by Hallmann (1912; as Echinoclathria gigantea) and Wiedenmayer (1989; as E. laminaefavosa), and the numerous 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 overlap 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 between 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; 1889a) 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 material examined in this study. Holopsamma macropora (Lendenfeld, 1888) (Figs 265-266) Plectispa macropora Lendenfeld,1888: 226. Wilsonella macropora; Hallmann, 1912: 240. Holopsamma macropora; Hooper & Wiedenmayer, 1994: 283. Not Echinoclathria macropora Whitelegge, 1901: 65, 89, 117; Whitelegge, 1902a: 212; Hallmann, 1912: 277. Not Echinoclathria macropora Whitlegge, 1907: 504. Not Clathria macropora; Whitelegge, 1901: 91. MATERIAL. HOLOTYPE: AMG9I59 (dry): Port Jackson, NSW, 33°51'S, 151°16'E (published locality of Torres Strait incorrect).0THER MATERIAL: S AUST- SAMTS4094 (fragments QMG300476, NTMZ1622). 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. Oscules. Small, up to 3mm in preserved state, scattered within meshes of surface lacunae. Texture and slidace characteristics. Firm, compressible, flexible; surface honeycomb reticulation close-meshed, meshes 3-6mm diameter, surface fibre bundles (lacunae) with scattered microconules on exterior edges, relatively deep meshes. Ectosome and subectosome. Surface microscopically hispid with points of principal styles protruding through surface, usually in plumose multispicular brushes; subectosomal auxiliary spicules tangential to surface in sparse bundles or individually, associated with protruding spicule brushes; choanosomal fibres immediately subectosomal. Choanosome. Skeletal architecture reticulate, increasingly plumose or plumo-reticulate near periphery, with a clearly differentiated primary and secondary fibre skeletons; primary ascending fibres very heavy, up to 120Rm 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 to 55p.rn diameter, very short, interconnecting close-set ascending fibres, usually aspicular, rarely unispicular; some fibres also cored by auxiliary spicules; all fibres abundantly echinated by principal styles, particularly at core of skeleton; spicule tracts within fibres become increasingly plumose towards periphery; fibre anastomoses form small oval or elongate meshes, 25-126Rm diameter; mesohyl matrix very lightly pigmented and auxiliary spicules scattered between fibres, particularly towards periphery; some detritus also incorporated into mesohyl but usually not into fibre skeleton; choanocyte chambers oval, 2065vim diameter. Megascleres. Choanosomal principal styles or subtylostyles coring and echinating fibres straight, moderately thin, with slightly constricted, subtylote or rounded, smooth bases, tapering to fusiforrn points. Length 49468.4)96Rm, width 3-(3.9)-61km. Subectosomal auxiliary megascleres range from strongylote subtylostyles to tornotes; usually long, thin, straight or slightly curved with blackened axial canals, sometimes setaceous, sinuous, with rounded or hastate points. Length 123-(166.1)-184Rm, width 1.5(3.2)-411,m. Microscleres. Palmate isochelae uncommon, many smaller forms twisted, with very short alae, - MEMOIRS OF THE QUEENSLAND MUSEUM 500^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 265. Holopsamma macropora (Lendenfeld) (holotype AMG9 159). 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 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 shaft, front ala usually partially, or sometimes wholly fused to lateral alae. Length 449.3)12p,m. Toxas absent. REMARKS. The two specimens are the only validated records for the species. Other records quoted as `macropora' by Lendenfeld and Whitelegge are spurious (see remarks for Plec tispa). From Lendenfeld (1888) and Whitelegge MEMOIRS OF THE QUEENSLAND MUSEUM 502zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ (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 macropora for several supposedly 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 acanthose as supposed by Lendenfeld (1888) and Hallmann (1912)), whereas all other alleged syntypes of Plectispa macropora belong to different species. In growth form H. macropora shows similarities to H. laminaefavosa, and to a lesser extent H. favus. In spiculation and skeletal construction it is reminiscent of specimens referred to H. laminaefavosa by Wiedenmayer (1989) and to a lesser extent H. ramosa (Hallmann, 1912) (i.e., those specimens with a diverging choanosomal skeleton which becomes increasingly dense towards the periphery). It is possible that H. Inacropora is simply a non-arenaceous morph of H. laminaefavosa although there are differences in spicule geometry and spicule dimensions to suggest that they are different species. In particular, the possession of long setaceous 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 (albiet uncommon in both the holotype and SA specimen), although they were not recorded in the original description of the species by Lendenfeld (1888), whereas microscleres are definitely not present in H. laminaefavosa. 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 pito-nava Pulitzer-Finali, 1982: 108109, text-figs 18-19. Holopsamma pluritoxa; Hooper & Wiedenmaycr, 1994: 283. MATERIAL. HOLOTYPE: MSNG 46938 (not seen): Heron I., Great Barrier Reef, Qld, 23°26'S, I51°55'E, 8.xii.1979, coll. A.J. Bruce (by hand).0THER MATERIAL: QLD- QMG6786 (fragments QMG300471, NTMZ1588). HABITAT DISTRIBUTION. 3-8m depth, on sand, shell grit and coral rubble substrates; Moreton Bay, Heron I. (Fig. 267E). DIAGNOSIS OF HOLOTYPE. Honeycombed reticulate growth form; irregularly reticulate skeleton with pale spongin fibres 20-100ilm diameter, fibre meshes 300-500Rm diameter, fibres sparsely cored by thin choanosomal principal subtylostyles (270-320x4-91Lm); echinating spicules absent; strongylote subectosomal auxiliary subtylostyles scattered throughout mesohyl (230-250x2-3p,m); abundant toxas wing-shaped or slightly oxhorn (10-188x14p,m); palmate isochelae unmodified (13.5p.m) (Pulitzer-Finali, 1982). DESCRIPTION. Shape. Subspherical-lobate, shaggy and irregularly honeycombed reticulate construction, 55mm wide, 40mm long. Colour. Live colouration unknown, light greybrown in ethanol. Oscules. Not seen. Texture and surface characteristics. Soft, compressible; surface reticulate fibre bundles in peripheral regions slightly flattened, with micropapillose, shaggy points. Ectosome and subectosonze. Surface microscopically hispid, with plumose brushes of choanosomal principal styles protruding, interdispersed with subectosomal auxiliary megascleres lying tangential or paratangential to surface; subectosomal skeleton plumoreticulate. Choanosome. Skeleton irregularly plumo-reticulate, with poorly developed spongin fibres, 20155Rm diameter, forming irregular ovoid to elongate meshes, 145-500Rm diameter; fibres contain pauci- to multispicular tracts of choanosomal principal styles, without any division of primary or secondary fibre components, forming plumose ascending structures which coalesce and diverge to produce fibre anastomoses ; fibres not echinated although choanosomal principal styles may protrude obliquely; mesohyl matrix relatively heavy, darkly pigmented, granular, with numerous toxas and scattered subectosomal auxiliary megascleres; choanocyte chambers large, oval, 40-90i.m. Megascleres. Choanosomal principal styles straight or slightly curved, with smooth, rounded or slightly subtylote bases, fusiform points. Length 1674217.2)-2891.1m, width 4-(5.1)-711m. Subectosomal auxiliary megascleres straight, asymmetrical, varing from quasistrongyles with REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 503zyxwvutsrqp E D. 0 0 FIG. 267. Holopsamma pluritoxa (Pulitzer-Finali) (QMG300471). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Accolada toxas. D, Section through peripheral skeleton. E, Australian distribution. F, Lateral view. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 504zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 268. Holopsamma pluritoxa (Pulitzer-Finali) (QMG300471). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal subtylostyles. D, Ends of principal subtylostyle. E, Auxiliary strongylote subtylostyle. F, Ends of auxiliary spicules. G, Accolada toxas. REVI SI ON OF MI CROCI ONI DAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 505 smooth rounded extremities, to quasitylotes with sponge, up to 165mm high, 220mm wide, with subt ylot e microspined point s. Lengt h 195small basal stalk and expanded point of attach(218.8)-26211m, width 2-(2.8)-411m. ment; branches numerous, bifurcate, occasionalMicroscleres. I sochelae absent. ly anastomosing, relatively thin, cylindrical or slightly laterally compressed, up to 75mm long, Toxas wing-shaped, variable in size, thin or 32mm diameter. thick, with rounded slight, or large central curvature, straight or slightly reflexed points, arbitrariColour. Grey-brown to dark brown in ethanol. ly divisible int o t wo size classes. Lengt h zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ I: Oscules. Moderately large, up to 5mm diameter, 8-(26.7)-30p.m, width 0.5-(0.8)-1p,m; length I I : scattered between honeycombed lacunae. 65-(177.0)-250p.m, width 1.5-(2.3)-4p.m. Texture and surface characteristics. Harsh, REMARKS. This species is unusual in its fibre slightly rubbery; fibre bundles form more-or-less characteristics and skeletal architecture, lacking regular hexagonal or elongated meshes, 1-3mm true echinating spicules, and having quasidiacdiameter. tinal auxiliary megascleres. The specimen from Ectosome and subectosome. Membraneous, with Moreton Bay also differs in some respects from point s of sparse principal st yles prot ruding the holotype from Heron I . (known from the through surface, but generally smooth, nonoriginal diagnosis; type not seen). The skeletal hispid; sparse subect osomal auxiliary architecture of the specimen is more obviously megascleres tangential to ectosome, never plumoreticulate than reticulate (reminiscent of protruding through surface. Stylinos; Halichondriidae), ectosomal structure is Choanosome. Skeletal architecture irregularly distinctly plumose (reminiscent of Lissodendoryx; Myxillidae; ectosomal structure was not reticulate, with small, heavy, evenly compressed ment ioned by Pulit zer-Finali, 1982) , and spongin fibres, 23-951.im diameter, divided into primary longitudinal and vaguely ascending, and isochelae are absent (abundant in the holotype). secondary connecting components; primary I n other respects the two specimens are obviously fibres contain uni- or paucispicular tracts of conspecific having virtually identical spicule choanosomal principal subtylostyles; secondary geometries, growth forms and surface features fibres mostly aspicular, rarely unispicular; fibre although Pulitzer-Finali's (1982) description is incomplete. meshes close-set near core, 35-85p.m diameter, more cavernous towards periphery, 112-220p,m diameter; tendency for peripheral fibres to conH olopsa m m a r a m osa (Hallmann, 1912) tain spicule tracts of styles in more-or-less(Figs 269-270) plumose brushes; fibres sparsely echinated by Echinoclathria ramosa Hallmann, 1912: 277-279, principal subtylostyles particularly on distal marp1.30, fig.3, text-fig.62. gins of peripheral fibres (cf. Hallmann, 1912); Axociella ramosa; de Laubenfels, 1936a: 119. mesohyl matrix heavy, darkly pigmented, with Holopsamma ramosa; Hooper zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA & Wiedenmayer, 1994: scattered subectosomal auxiliary megascleres; 283. choanocyte chambers small, oval, up to 50p.m Echinoclathria macropora; Whitelegge, 1901: 89, diameter. 117. Not Plectispa macropora Lendenfeld, 1888: 226. Not Echinoclathria macropora; Whitelegge, 1907: 504. MATERIAL. LECTOTYPE: AMZ949 (dry): Swansea, off Lake Macquarie, NSW, 33°11'S, 151°59'E, depth unknown, coll. FIV 'Thetis' (trawl). PARALECTOTYPE: AMZ40: 64km W. of Kingston, SA, 36°50'S, 139°05'E, 60m depth, coll. FIV 'Endeavour' (trawl).0THER 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). D ESCRI PTI ON . Shape. Profusely bushy, arborescent branching, honeycomb reticulate Megascleres. Choanosomal principal st yles straight, short, relatively thick, with tapering, constricted, smooth, slightly subtylote bases, slight ly fusiform sharp point s. Lengt h 75(100.7)-144p.m, width 4.5-(6.8)-8.5p.m. Subectosomal auxiliary subtylostyles or quasistrongyles (asymmetrical) straight or curved at centre, thin, with smooth, slightly subtylote bases and rounded points. Length 1224139.8)-157p.m, width 1.542.7)-411,m. 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)-12p.m. 506zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 269. Holopsamma ramosa (Hal!mann) (holotype AMZ949). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Palmate isochelae. D, Section through peripheral skeleton. E, Known Australian distribution. F, Holotype. G, SAMTS4105. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 507 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. MEMOIRS OF THE QUEENSLAND MUSEUM 508zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ REMARKS. Hallmann (1912) suggested that spicule dimensions were highly variable in this species, particularly choanosomal styles, but in comparison with other Holopsamma species these are in fact relatively homogeneous. Similarly, 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, differing mainly in growth form, spicule dimensions and the presence of microscleres. Indeed 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 H. macropora, H. arborea and H. ramosa is not presently supported on the basis of existing material. Holopsamma rotunda (Hallmann, 1912) (Figs 271-272) Echinoclathria rotunda Hallmann, 1912: 282-284, p1.30, fig. 1, text-fig.64. Axociella rotunda; de Laubenfels, 1936a: 119 [note]. Holopsanuna rotunda; Hooper & Wiedenmayer, 1994: 284. MATERIAL. LECTOTYPE: AMZ154: Off Swansea, Lake Macquarie, NSW, 33°11'S, 151°59'E, coll. FIV 'Endeavour' (dredge). PARALECTOTYPE: AME1272 (dry): same locality.OTHER MATERIAL. QLD- QMGL2165. VIC- QMG300275 (NCIQ66C3288-P) (fragment NTMZ3865). HABITAT DISTRIBUTION. Substrate unknown; up to 330m depth; Lake Macquarie (NSW); off Fraser I. (SEQ); Portsea (Vic) (Fig. 271E). DESCRIPTION. Shape. Massive, subspherical, globular, honeycomb reticulate sponge, up to 155mm high, 90mm maximum diameter, with short basal stalk and expanded point of attachment; sponge insubstantial, hollow internally. Colour Dark blue-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, compressible alive, harsh in dry state; open, porous, tubular external construction; honeycombed surface fibre bundles produce very thin, flattened, erect lamellae with regular meshes, up to 4mm diameter. 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. Choanosome. Skeletal architecture distinctly plumo-reticulate, with ascending multi- or paucispicular primary fibres, interconnected by pauci- or aspicular secondary fibres; fibres relatively light, 43-92p,m primary fibre diameter, 18-56p.m secondary fibre diameter, prominently flattened; fibre anastomoses form almost regular, circular or oval meshes, 38-96p,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 confined 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 dispersed subectosomal auxiliary subtylostyles; choanocyte chambers small, oval, up to 55p,m diameter. Megascleres. Choanosomal principal subtylostyles straight or slightly curved at centre, relatively long, thick, with tapering, constricted, smooth, subtylote bases, fusiform pointed or slightly telescoped points. Length 85-(107.5)-138p,m, width 4-(7.2)-9.5p,m. Subectosomal auxiliary subtylostyles thin, straight, slightly curved or srometimes sinuous, slightly subtylote bases, varying from asymmetrical strongylote with rounded points or styloid with tapering points. Length 1234152.3)196p,m, width 2.5-(2.9)-4p.m. Microscleres. Palmate isochelae abundant, unmodified, with long lateral alae completely attached to shaft, shorter front ala partially fused to lateral alae, straight shaft. Length 8410.7)12p.m. REMARKS. Holopsamma rotunda has an insubstantial, hollow morphology, paper thin, lamellate fibre bundles (lacunae), and regularly lamellate fibre reticulation with prominent flattened fibres, whereas in spicule geometry it could match most other species, particularly H. crassa, H. elegans, H. macropora and H. ramosa (i.e., with strongylote points on auxiliary spicules). So far the species is only known only from two type specimens, a poorly preserved deep water specimen from the E. continental shelf (represented only by a fragment), and a more recent REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 509 FIG. 271. Holopsamma rotunda (Hallmann) (lectotype AMZ154). A, Principal subtylostyle (coring and echinating fibres). B, Subectosomal auxiliary subtylostyle. C, Palmate isochelae. D, Section through peripheral skeleton. E, Known Australian distribution. F, Lectotype. ^ MEMOIRS OF THE QUEENSLAND MUSEUM 510zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 272. Holopsamma rotunda (Hal!mann) (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 Macquarie, NSW. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 511 Holopsamma simplex (Lendenfeld, 1885) (Figs 273-274) Halme simplex Lendenfeld, 1885c: 301-303, p1.26, fig.3, p1.27, 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 sponges, reportedly up to 20inm thick, with lobate margins and a maximum lateral spread of 100mm; surviving portion of lectotype is 30mm diameter and lOmm thick. Colour Live colouration apparently brown, beigebrown in dry state. Oscules. Undifferentiated from honeycomb reticulate trabeculae. Texture and surface characteristics. Surviving portion of lectotype brittle, partially crumbled when dry; surface minutely porous, partially arenaceous; close reticulation of rounded branches or 'lacunae' form interweaved ridges on encrusting base produced by irregularly honeycombed reticulate surface fibres. Ectosome and subectosome. Membraneous, with thick sand cortex; no specialised spicules or spongin fibres visible on FIG. 273. Holopsamma simplex (Lendenfeld) (lectotype AMG8822). A, Principal styles (coring and echinating fibres). B, Subectosomal auxiliary subtylostyles. C, surface; subectosome Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. arenaceous. 512zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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-byside, bonded together with abundant brown collagen; 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 telescoped points, sometimes slightly fusiform. Length 43-(50.8)-62p,m, width 3-(4.1)-5Rm. Subectosomal auxiliary spicules thin subtylostyles, occasionally quasi-strongyles, straight, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 513 slightly curved or sinuous, with asymmetrical ends. Length 68-(84.8)-99Rm, width 242.7)3.5 p.m. 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 accepted 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 1OR 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; surface 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 40011m diameter, partially or fully cored with detritus; mesohyl matrix very heavy; no native spicules; choanocyte chambers oval to elongate, 20-55[1,m diameter. Megascleres. Absent. Microscleres. Absent. 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. Wiedenmayer (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 architecture 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. TYPE SPECIES. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ Ophlit aspongia aust raliensis Ridley, I 884a: 442 (by subsequent designation of Hallmann, 1912: 288)). DEFINITION. Monactinal, quasi-monactinal or thin diactinal auxiliary megascleres tangential or erect on ectosome; choanosomal skeleton irregularly reticulate; fibres cored by tracts of auxiliary megascleres, identical to those in ectosomal skeleton, and echinated by principal megascleres varying from true monactinal, quasidiactinal to true diactinal forms, smooth or acanthose; microscleres may include palmate isochelae and toxas. REMARKS. Twenty six species have been included 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 & Levi, 1993a). The genus contains 2 groups: one (Echinochalina (Echinochalina)) with true monoactinal spicules, showing superficial affinities with Holopsamma (having honeycomb reticulate growth forms) and the Raspailiidae (fibre characteristics), and the other with quasi-diactinal or secondarily modified diactinal megascleres (Echinochalina (Protophlitaspongia)) superficially resembling Niphatidae (Haplosclerida). Both groups are linked by the common possession of auxiliary megascleres coring fibres and principal megascleres echinating fibres. Echinochalina (Echinochalina) Thiele, 1903 Echinochalina 1912: 288. Thiele, 1903a: 961; sensu Hallmann, MEMOIRS OF THE QUEENSLAND MUSEUM 514zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 275. Holopsamma sp. indet. (QMG300620). A, Choanosomal skeletal structure. B, Australian distribution. Tab/is de Laubenfels, 1936a: 76. [Echinoclathria]; Uriz, 1988: 89. TYPE SPECIES. Ophlitaspongia australiensis Ridley, 1884a: 442. Echinochalina anomala Hallmann, 1912: 292-294, text-fig.68; Hooper & Wiedenmayer, 1994: 276. Tab/is anomala; de Laubenfels, 1936a: 76. cf. Echinochalina anomala Burton, 1934a: 563. DEFINITION. Monactinal, quasi-monactinal or secondarily derived true diactinal auxiliary megascleres core fibres, and monactinal principal spicules echinate fibres. MATERIAL. HOLOTYPE: AMG10548 (dry): Off Woolongong, NSW, 34°25'S, 151°10'E, 110-112m depth, coll. FIV 'Thetis' (dredge). PARALECTOTYPES - AMG10549 (dry): same locality. AMG10550 (dry): same locality. AMG10551 (dry): same locality. Echinochalina (Echinochalina) anomala HABITAT DISTRIBUTION. 110-112m depth; substrate unknown; S. coast (NSW) (Fig. 276D). Hallmann, 1912 (Figs 276-277) Echinochalina glabra, in part; Whitelegge, 1907: 507. Not Echinoclathria glabra Ridley & Dendy, 1887: 163. DESCRIPTION. Shape. Massive, clathroushoneycomb 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 515 A 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 substrate. Colour Live colouration unknown, grey-brown in dry state. Oscules. Not observed. Texture and surface characteristics. Firm, harsh in dry state; surface fibres flattened, paper-like, porous reticulate with fibre-branches forming wide-meshed reticulation up to 3mm diameter. Ectosome and subectosome. Membraneous, stretched across adjacent surface fibre-branches, with tangential auxiliary megascleres sparsely dispersed within ectosome, singly or less frequently 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, 230-3601i.m diameter, irregularly interconnected by relatively thinner secondary fibres, 95-145Rm diameter, forming cavernous meshes, 43010901.i.m diameter; all fibres cored by multispicular tracts of auxiliary megascleres 516zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 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)-236p,m, width 749.3)11p,m. Auxiliary megascleres coring fibres and found interstitially long, thin, straight, entirely smooth, varying from nearly symmetrical quasi-strongyles to asymmetrical tomostyles with slightly subtylote bases and rounded points. Length 165- (215.4)-277u,m, width 2-(4.3)-6.5Rm. Microscleres. Absent. REMARKS. Burton (1934a) assigned a specimen from Eagle I. (Cairns section, Great Barrier Reef) to E. (E.) anomala but his material 518zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM 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 Echinodictyum (Raspailiidae), with very widely spaced fibremeshes, 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 organic fibres, and which have true diactinal coring spicules, the fibres of E. (E.) anomala are prominent and heavily cored by quasi-diactinal 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, differences in fibre construction and spicule dimensions. Echinochalina (Echinochalina) australiensis (Ridley, 1884) (Figs 278-279)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Ophlit aspongia aust r aliensis Ridley, 1884a: 442-443, FIG. 279. Echinochalina ( Echinochalina) aust r aliensis (Ridley) (holotype BMNH1881.10.21.299). A, Choanosomal skeleton. B, Fibre characteristics. p1.42, figs c,c'. C, Principal subtylostyle. D, Ends of principal spicule. E, Auxiliary spicule. Echinochalina aust r aliensis; Thiele, 1903a: 961-962; F, Ends of quasi-diactinal auxiliary spicule. Hooper & Wiedenmayer, 1994: 276. DESCRIPTION. Shape. Massive, branching, MATERIAL. HOLOTYPE: BMNH 1881.10. 21.299: Port Molle, Qld, 20°16'S, 148°42'E, 1.v.1881, HMS 'Alert' (dredge). OTHER MATERIAL: SMF1855 (fragment MNHNDCL2265). HABITAT DISTRIBUTION. Rock and coral rubble substrata; 24m depth; Airlie region (NEQ) (Fig. 278D); also Moluccas, Indonesia. clathrous, honeycomb reticulate growth form, 55mm diameter, fibre-branches flattened, up to 12mm long, 5mm wide, with cavernous, angular meshes up to lOmm diameter. Colour. Pale brown dry. Oscules. Small, up to 4mm diameter, between adjacent fibre bundles. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 519 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 MNHNDT3411. F, Lectotype of S. favosa. G, Holotype of E. glabra. MEMOIRS OF THE QUEENSLAND MUSEUM 520zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 281. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 521 Texture and sutface characteristics. Firm, harsh dry; conulose surface produced by anastomosing fibre bundles ('lacunae'), interconnected by translucent ectosomal membrane stretched between adjacent conules. Ectosome and subectosome. Membraneous, with thinner choanosomal, undulating fibres lying immediately below ectosome, approximately 53p,m diameter, cored by paucispicular tracts of auxiliary megascleres, and with irregular layer of auxiliary megascleres scattered tangential to surface. Choanosome. Vaguely regularly reticulate skeleton, composed of primary, distinctly ascending, multispicular fibres, 100-140p,m diameter, interconnected at irregular intervals by numerous transverse, paucispicular, secondary fibres, approximately 70p,m diameter; fibre anastomoses form rounded or elongate, narrow meshes up to 270p,m diameter; fibres cored by auxiliary megascleres, sparsely echinated by principal subtylostyles; mesohyl matrix heavy but only lightly pigmented, with sparsely dispersed auxiliary megascleres between fibres; choanocyte chambers ovoid, 20-25vm diameter. Megascleres. Principal subtylostyles (echinating fibres) straight, smooth, tapering, rounded or slightly subtylote bases, fusiform or slightly telescoped points. Length 83-(111.2)-131p,m, width 6-(7.2)-9.5p,m. Auxiliary megascleres (coring fibres and interstitial) straight, thin, subtylostyles, tornostyles or quasi-stongyles, with smooth, rounded, or very slightly subtylote bases, tapering-rounded points. Length 147-(178.2)-192pLm, width 1.5-(3.4)4.5 Rm. Microscleres. Absent. REMARKS. Ridley (1884a) reported toxas (up to 42p,m long, 2p,m 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, superficially resembling Echinodictyum cavemosum (Raspailiidae) and C. (Thalysias) vulpina in growth form, with a heavy, well developed reticulate, 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. Echinochalina (Echinochalina) barba (Lamarck, 1814) (Figs 280-281, Plate 12A) Spongia barba Lamarck,1814: 372, 354; Topsent, 1930: 15-16, p1.4,fig.6. Echinochalina barba; Hooper & Wiedenmayer, 1994: 276. Spongia favosa Lamarck, 1814: 373; Topsent, 1930: 19, p1.1, fig.5. Echinoclathria glabra Ridley & Dendy, 1886: 476; Ridley & Dendy, 1887: 163, p1.29, fig.11, p1.31, fig.2; Dendy, 1896: 40. Tab/is glabra; de Laubenfels, 1936a: 76. Echinochalina glabra; Thiele, 1903a: 962; Hallmann, 1912: 275, 290-292, text-fig.67; Burton, 1934a: 600; [?] Thomas, 1977: 115-116, text-figs 1F-G. Not Echinochalina glabra; Whitelegge, 1907: 504. MATERIAL. LECTOTYPE: MNHNDT561: unknown locality. PARALECTOTYPE: MNHNDT3411: unknown locality. LECTOTYPE of S. favosa: MNHNDT557 (fragment BMNH1954.2.20.9): Near King I., Tas, 39°50'S, 144°00'E, 1803, Peron & Lesueur collection. PARALECTOTYPE 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: QLD-QMGL711 (fragment NTMZ1531). TAS- QMG300274 (NCIQ66C-3543-R) (fragment NTMZ3849), QMG311312 (NCIQ66C3542-Q) (fragment NTMZ3853). S AUST- AMZ44. HABITAT DISTRIBUTION. Sand, shell grit, rock or coral rubble substrata, kelp beds; 0-76m depth; Moncoeur I., Port Phillip Bay (Vic); King I., Deal I., Bass Strait (Tas); Kingston SE. (SA); Flinders Is (FNQ) (Fig. 281D). Thomas' (1977) reported locality of Andaman Sea is dubious from his description. DESCRIPTION. Shape. Massive, subcylindical, honeycomb reticulate sponge, up to 155mm long, 75mm wide, consisting of thin, flattened, irregularly anastomosing, reticulate fibre bundles; fibre anastomoses form meshes 3-5mm diameter. Colour. Light grey-brown exterior, beige interior alive (Munsell 7.5Y816), 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. Ectosonze and subectosome. ^Thinly membraneous, with irregularly dispersed 522zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR A BzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFE 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^ 523 auxiliary megascleres lying tangential to surface and irregular, paucispicular, plumose brushes of principal spicules erect on peripheral fibres, barely protruding through ectosome; choanosomal fibres immediately subdermal. Choanosome. Irregularly reticulate skeletal structure, with primary, vaguely ascending, distinctly meandering paucispicular fibres, 52120p,m diameter, interconnected by aspicular or less frequently paucispicular, irregularly anastomosing, secondary fibres, 37-75p.m diameter; fibres cored by auxiliary megascleres, moderately echinated by principal subtylostyles especially on peripheral fibres; fibre anastomoses form nearly regular regular oval meshes, 165-370p,m diameter; mesohyl matrix heavy, moderately lightly pigmented, with numerous auxiliary megascleres dispersed between fibres; choanocyte chambers oval to eliptical, 38145p,m diameter. Megascleres. Principal subtylostyles (echinating fibres) short, thick, straight or very slightly curved, with smooth, tapering, distinctly subtylote bases, fusiform points. Length 62494.3)115p,m, width 5-(7.1)-8.5p,m. Auxiliary megascleres (coring fibres and interstitial) straight, quasi-diactinal subtylostrongyles, rarely strongyles, with smooth, slightly subtylote bases and rounded points. Length 147(189.2)-214pm, width 1.5-(3.3)-4.5p,m. Microscleres. Absent. REMARKS. Echinochalina barba has a honeycombed reticulate construction similar to that of E. (E.)anomala and reminiscent of Holopsamma species, but in which the surface is very smooth and fleshy (superficially resembling an Acanthella (Axinellidae)). Both E. anomala and E. barba also have asymmetrical (quasi-diactinal) 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 I., E. of Turtle Is group, Cairns section Great Barrier Reef, 14°39.8'S, 145°19.2'E, 3.ix.1994, 19m depth, coll. J. Kennedy, DPI RV "Gwendolyn May", trawl. HABITAT DISTRIBUTION. Soft substrata inter-reef region; 19m depth; Turtle Is (FNQ) (Fig. 282F). DESCRIPTION. Shape. Long, thinly cylindrical, fragile digit, 180mm long, up to 60mm diameter tapering 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 in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, compressible; 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, membraneous ectosome stretched between adjacent fibres; outer surface of peripheral fibres with small blind fibre endings and protruding spicules, both visible under low magnification; auxiliary subtylostyles scattered on membraneous ectosome, singly or in small bundles; subectosomal skeleton undifferentiated from choanosome. Choanosome. Reticulate skeletal structure; fibres more-or-less homogeneous, 40-85 p.m diameter cored by auxiliary subtylostyles in multispicular tract of up to 6 spicules abreast; fibres long, generally curved or sinuous, anastomosing, regularly bifurcate; shorter connecting fibres unior paucispicular; fibre meshes cavernous oval or eliptical, 170-490p.m 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)-278p.m, width 3-(3.3)-4.5p,m. Auxiliary subtylostyles (coring fibres and dispersed 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)-5p.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)-12pm. Toxas include both wing-shaped and slightly accolada forms, ranging from thin to raphidiform, often broken in preparations. Length 40465.8)96pm, width 0.841.7)-2.5 pm. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 524zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA z 20um 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. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 525 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, Naturhi storisches Museum Basel, for his contributions to Australian sponges. REMARKS. Echinochalina felixi is unusual having very thin megascleres and a cavernous reticulate construction, similar to E. digitata al- MEMOIRS OF THE QUEENSLAND MUSEUM ^ 526zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA FIG. 285. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 527 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°09'8, 144°52'E, 11-13m depth, coll. J.B. Wilson (dredge). HABITAT DISTRIBUTION. Encrusting of shell fragments; 11-13m depth; Port Phillip (Vic) (Fig. 284F). DESCRIPTION. Shape. Massive, lobate, semiencrusting on detritus, 130mm long, 65mm wide, up to 45mm thick, with low bulbous lobes scattered over surface. Colour Brown in ethanol. Oscules. Single, large apical oscule, up to 3.5 mm diameter, on each surface lobe. Texture and RI/face characteristics. Firm, compressible, easily torn; surface lobate-bulbous, uneven, microscopically rugose, reticulate, granular. Ectosome and subectosome. Membraneous, skinlike ectosomal skeleton, minutely hispid with points of auxiliary styles protruding through surface in pauci- or multispicular plumose brushes, some arising from ascending choanosomal skeletal 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 forming primary, more-or-less ascending, and secondary connecting tracts; primary fibres, 40-70iim diameter, cored by multi- or paucispicular tracts of auxiliary styles in plumose or disorganised arrangement; secondary connecting fibres, 1532p.,m diameter aspicular or unispicular, curved, meandering throughout core of choanosomal skeleton producing oval, elongate, cavernous meshes, 180-310p,m diameter, becoming more regular, rectangular and cavernous towards peripheral region of skeleton, with meshes 18036011,m diameter; peripheral fibres slightly more compacted; spicule tracts heavier in subectosomal 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, 62135 tan diameter. Megascleres. Principal styles echinating fibres straight, short, relatively thick, with smooth, rounded or slightly constricted, subtylote bases and nearly hastate points. Length 1024117.7)13811m, width 3.8-(5.5)-6.5p.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)-264p.m, width 1.842.7)-4.5 Microscleres. Isochelae absent. Toxas moderately uncommon, oxeote, with slight angular central curvature, straight points. Length 904116.9)-13511m, width 1.542.2)Larvae. Viviparous, oval-elongate parenchymella, 225-375x190-210Rm, dark orange, heavily collagenous, containing oxeote toxas and juvenile styles. REMARKS. De Laubenfels (1936a) referred this species to Echinochalina (sensu Hallmann, 1912) having fibres cored by megascleres geometrically equivalent to auxiliary spicules, whereas styles echinating fibres have a geometry equivalent to principal megascleres of other Microcionidae. Dendy's (1896) description differed in spicule dimensions from my observations of the type. Oxeote toxas, much smaller than those described by Dendy, are predominantly in association with 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, and 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 viminalis Lendenfeld, 1888: 225. Echinoclathriaintermedia; in part, Whitelegge, 1902a: 214-215; Dendy, 1922: 71, p1.2, fig.8. Echinochalina intermedia; Hallmann, 1912: 294-295, text-fig.69; Shaw, 1927: 427; Burton, 1934a: 563; Hooper & Levi, 1993a: 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, 153°10'E, 25m depth, 10.xii.1991. MEMOIRS OF THE QUEENSLAND MUSEUM 528zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ FIG. 286. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Echinochalina ( Echinochalina) int erm edia (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 & Levi, 1993, for QMG300834, QMG300865. INDONESIA a list of additional material used in this study): QLD- NCIOCDN-1306-F (fragment QMG303688). QMG304757, QMG304293, QMG304388, QMG303863, QM0304025, QMG303984, HABITAT DISTRIBUTION. Sand, rock and coral rubble substrata; 8-50m depth; Lizard I., Day Reef, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ ^ 529 and conules upon preservation; texture soft, compressible, mucusy; pale orange-brown alive, brown in ethanol. Ectosome membraneous, with tangential and paratangential tracts of auxiliary spicules scattered, barely protruding through surface; ectosome heavily collagenous, darker than choanosomal mesohyl; choanosomal skeleton irregularly reticulate, with long primary fibres, 90130p,m diameter, meandering through choanosome, becoming more plumoreticulate near periphery, cored by multispicular tracts of whispy auxiliary spicules, interconnected by short, aspicular ot paucispicular secondary fibres, 40-70pm diameter, producing elongate or eliptical, cavernous meshes up to 450t.im diameter; fibres sparsely echinated by principal spicules; mesohyl matrix heavy, with abundant auxiliary spicules scattered between fibres; choanocyte chambers large, oval, up to 6011m diameter. Principal styles short, thick, straight, smooth tapering or slightly subtylote bases, hastate or slightly telescoped points FIG. 287. Echinochalina (Echinochalina) intermedia (Whitelegge) (72-95x2.5-4.5 p.m). QMG300834). A, Choanosomal skeleton. B, Fibre characteristics. C, Principal Auxiliary spicules subtylosstyle. D, Ends of principal style. E, Auxiliary subtylostylc/ tornostyle. F, Ends tyles or tornostyles, long, of auxiliary spicule. slender, straight, slightly subtylote smooth bases, Direction Is, Howick Is, Snake Reef, Frankland Is, rounded points (178-206x2-3.5pm). Sudbury Reef, Stanley Reef, Gulf of Carpentaria Microscleres absent. (FNQ); Hook Reef (NEQ); Mooloolaba, Moreton Bay (SEQ); Illawarra (NSW); E. coast (Tas); Dampier Archipelago, Northwest Shelf (WA) (Fig. 286F); also New Caledonia (Hooper & Levi, I993a), Cargados Carajos, Indian Ocean (Dendy 1922); Sulawesi, Indonesia (present study). DESCRIPTION OF NEOTYPE. Massive, lobate, bulbous-digitate, 105mm high, 65mm maximum breadth; convoluted bulbous, cavernous surface in life, collapsing to form sharp ridges DESCRIPTION. (See Hooper & Levi, 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 translucent MEMOIRS OF THE QUEENSLAND MUSEUM 530zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ TABLE 44. Comparison in spicule dimensions between populations of Echinochalina (Echinochalina) intennedia (Whitelegge). Measurements in pan (N=25). Comparative data from material examined here and Hooper & Levi (1993a). Population (Latitude) Auxiliary subtylostyles (coring fibres) Principal styles (echinating fibres) Neotype (QMG 300025) (26°S) 178-(190.7)-206 x 2(2.6)-3.5 New Caledonia (22°S) I69-(175.3)-189 x I.2-(1.9)-2.5 72-(85.1)-95 x 2.5(3.1)-4.5 64-(87.1)-I13 x 2.0(2.8)-4.0 Great Barrier Reef (17-19 ° S) I84-(193.8)-205 x 2.0-(3.1)-5.0 85493.8)-110 x 4(5.2)-6 Southern Qld. (26 ° S) I 844196.2)-208 x I.5-(2.2)-3 NW Shelf WA (20 ° S) 89-(148.2)-180 x I .5-(2.5)-4.0 72-(88.3)-10I x 2.5(2.9)-4 92-(126.5)-I49 x 4(5.7)-7 Cargad9s Carajos (16 ° S) 200 x 3 110 x 7 Illawarra, NSW2 (34 ° S) 140-185 x 4 80-150 x 9 Sulawesi, Indonesia (1 ° N) 177-(183.2)-198 x I .5-( I .9)-2.5 60482.7)-113 x 2.0(3.2)-4 Source: 1. Dendy (1922). 2. Hallmann (1912). skin-like membrane covering, darker below; ectosome membraneous with tangential or occasionally paratangential sparse tracts of auxiliary spicules, lightly arenaceous, and sparse plumose brushes of auxiliary megascleres from ascending choanosomal fibres protruding short distance through surface predominantly on surface conules; choanosomal skeleton irregularly plumo-reticulate, spongin fibres well developed; primary fibres lightly cored by multispicular tracts of auxiliary megascleres becoming increasingly plumose towards periphery; secondary connecting fibres uni-, pauci- or aspicular; all fibres irregularly echinated by sparse principal styles, mainly on primary fibres; fibre anastomoses form irregular elongate, oval, eliptical or rarely rectangular meshes, meshes more cavernous in periphery than at core; mesohyl matrix heavy, numerous auxiliary spicules and detritus scattered 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 within mesohyl long, thin, mostly straight, with rounded smooth bases, varying from styles, tomostyles or less often strongyles, usually with blackened axial canals, with rounded or slightly hastate points; microscleres absent. REMARKS. This species was redescribed mainly from New Caledonian material (Hooper & Levi (1993a). 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 multispicular axis of primary spicule tracts and light pauci- or aspicular tracts in secondary fibres, and hastate choanosomal (auxiliary) styles core fibres. Hooper & Levi (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, I902a) and the Northwest Shelf of WA (Hooper & Levi, 1993a), 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 intennedia is used in the sense of Hallmann (1912) and Dendy (1922), because the synonymy of this species is still confused, and the validity of type material of Thalassodendron viminalis and Echinochalina intennedia require further clarification. Lendenfeld (1888: 225) erected T viminalis for a specimen from Illawarra, NSW, whereas Whitelegge (1901: 87) found the original description incorrect based on a misidentified 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 subsequently 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 published descriptions (see remarks for E. subhispida). I accept E. intennedia 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. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 531 FIG. 288. Echinochalina (Echinochalina) reticulata Whitelegge (holotype AMZ950). A, Principal style (echinating fibres). B, Auxiliary subtylostyle/ tornostyle (coring fibres and interstitial). C, Section through peripheral skeleton. D, Australian distribution. E, Neotype. MEMOIRS OF THE QUEENSLAND MUSEUM 532zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Echinochalina (Echinochalina) reticulata Whitelegge, 1907 (Figs 288-289) Echinochalina reticulata Whitelegge, 1907: 506, p1.45, fig.25; Hallmann, 1912: 289-290, p1.30, fig.2, text-fig.66 (et var.); Hooper& Wiedenmayer, 1994: 278. Tablis reticulata; de Laubenfels, 1936a: 76. MATERIAL. LECTOTYPE: AMZ950: Off Wollongong, NSW, 34°26'S, 150°53'E, I10-112m depth, coll. FIV 'Thetis' (dredge). PARALECTOTYPES: AMZ715: same locality. AMZ23 (dry): E. coast of Hinders 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: AMZI53, 716, 717). HABITAT DISTRIBUTION. Soft substrata; 110112m depth; Wollongong (NSW); Hinders 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 reticulation. Texture and surface characteristics. Firm, flexible, harsh in dry state; surface honeycomb reticulate, lamellae very thin, membraneous and uneven. Ectosome and subectosome. Membraneous, tangentially 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 differentiated primary and secondary spongin fibre systems; fibres relatively light, mostly flattened, meandering or sinuous; primary fibres, 65112p.,m diameter, paucimultispicular, or, running longitudinally through lamellae, intersected at regular intervals and obtuse angles by secondary fibres, 22-40iim diameter, mostly uncored, occasionally unispicular; fibres anastomoses form cavernous eliptical meshes, 175-570Rm diameter; fibres cored by auxiliary megascleres, moderately heavily echinated by principal spicules; mesohyl matrix heavy but only lightly pigmented, with numerous auxiliary megascleres scattered between fibres; choanocyte chambers large, oval, 45-751im diameter. Megascleres. Principal styles echinating fibres straight, thick, with smooth, tapering, rounded or slightly subtylote bases, fusiform points. Length 108-(139.8)-196pAn, width 6-(8.9)-11p,m. 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)-336Rm, width 2.544.1)Microscleres absent. Larvae. Viviparous, small, oval parenchymella, 80-110p,m diameter, with heavy matrix. REMARKS. In growth form, particularly its flattened spongin fibres, the holotype resembles the Caribbean Pandaros acanthifolium (Duchassaing & Michelotti), whereas Hallmann's variety crassa is more reminiscent of honeycomb reticulate 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 subtylostyles, 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) Echinodictyurn 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 Phillip Bay, Vic, 38°09'S, 144°52'E, 40m depth, coll. J.B. Wilson (dredge). PARALECTOTYPES: 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; 3840m depth; Port Phillip (Vic) (Fig. 290D). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 lamelshaped, 85-165mm long, 50-80mm maximum lae, up to 18mm wide, together producing bushy, width, with thick cylindrical stalk, 18-32mm long, 12-17mm diameter, bulbous digitate apex flattened flabelliform or claviform growth. MEMOIRS OF THE QUEENSLAND MUSEUM 534^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Echinochalina ( Echinochalina) ridleyi (Dendy) (lectotype NMVG2409). A, Principal acanthostyle FIG. 290. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA (echinating fibres). B, Auxiliary tornostyle/ oxeote (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 535 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. MEMOIRS OF THE QUEENSLAND MUSEUM 536zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Colour Reportedly pale red-brown to red-orange in life, pale brown in ethanol. Oscules. Not observed. Texture and surface characteristics. Firm, compressible, 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 immediately below ectosome. Choanosome. Plumo-reticulate skeletal structure, with incompletely differentiated, meandering, primary and secondary spongin fibres; primary fibres heavy, 52-116pm diameter, vaguely ascending, multispicular, spicules arranged in loose whispy tracts occupying most of fibre diameter, becoming increasingly plumose towards periphery; secondary, connecting fibres, 18-43p.m diameter, pauci-, uni- or aspicular, producing cavernous, oval or elongate meshes, 75-365p,m 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, 90-1201.tm diameter. Megascleres. Principal acanthostyles echinating fibres small, straight, relatively thick, with tapering rounded bases, spined apical and basal extremities, fusiform points; spines small, conical, erect. Length 87-(104.1)-116p,m, width 545.8)7p,m. Auxiliary spicules coring fibres long, thin, straight, slightly curved or sinuous, sometimes raphidiform, with variable terminations ranging from hastate oxeas, tomotes, quasi-monactinal tornostyles, or subtylostyles. Length 184(261.3)-314p.m, width 2-(4.1)-6.5p,m. Microscleres Absent. REMARKS. Dendy (1896) initially assigned this species to Echinodictyum (Raspailiidae) having diactinal coring, monactinal echinating megascleres, extra-axial styles, and a more-orless reticulate architecture. However, the spicules which actually core these fibres range from true diactinal to true monactinal forms, without obvious 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 that genus (see Hooper, 1991). Including this species in Echinochalina requires broadening the generic definition to allow for the inclusion of spined echinating styles, but these spicules appear 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 systematic levels in Microcionidae, and in other respects the species fits well with the present concept of Echinochalina. The species has closest affinities to 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. Echinochulina spongiosum; Hooper, 1991: 1348; Hooper & Wiedenmayer, 1994: 278. MATERIAL. LECTOTYPE: NMVG2452: Sorrento Jetty, Port Phillip Bay, Vic, 38°21'S, 144°42'E, 20m depth, coll. J.B. Wilson (dredge). PARALECTOTYPE: 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, compressible, spongy; surface shaggy, covered with large, mostly irregular, lobate papillae, low ridges and abundant striations. Ectosome and subectosome. Membraneous, lightly arenaceous ectosome, with irregular plumose brushes of auxiliary megascleres barely protruding through surface arising from peripheral choanosomal fibres. Choanosome. Plumo-reticulate skeletal structure, 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, 22-54p,m diameter, interconnected at irregular intervals by smaller multispicular fibres, REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 537 LO CN FIG. 292. Echinochalina (Echinochalina) spongiosa (Dendy) (lectotype NMVG2452). A, Principal acanthostyles (echinating fibres). B, Auxiliary tornostyles (coring fibres and interstitial). C, Section through peripheral skeleton. D, Known Australian distribution. E, Lectotype. MEMOIRS OF THE QUEENSLAND MUSEUM 538zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 tomostyle. F, Ends of auxiliary spicule. 8-181J.m diameter, forming irregular ovoid, rec- paucispicular towards periphery, terminating on tangular or elongate meshes, 65- 1 18p.m or near surface as single spicules; skeletal tracts diameter; fibres increasingly plumose, more in periphery occupy most of fibre diameter, rang- ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 539 FIG. 294. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Echinochalina ( Echinochalina) t ubulosa (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 12-35p,m diameter, fibre meshes cavernous, elongate, 240-66011m 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, 15-54p.m diameter. Megascleres. Principal styles echinating fibres straight, relatively thin, subtylote, with light 540zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM spines concentrated mainly on base and near points, fusiform, bare points. Length 82492.3)981,tm, width 2.5-(4.4)-5.5Rm. Auxiliary spicules coring fibres thin, mostly straight, sometimes slightly curved or sometimes sinuous, varying from tornostyles, strongyles or tomotes with rounded or subtylote extremities. Length 166-(187.6)-214Rm, width 1.543.2)4.211m. 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 skeletal 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 (versus completely smooth) principal styles echinating 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, p1.35, fig.3, text-fig.60. Echinochalina tubulosa; de Laubenfels, 1936a: 119; Hooper & Wiedenmayer, 1994: 278. Echinoclathria tubulosa; Wiedenmayer, 1989: 66, p1.6, fig.9, p1.25, 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. AMZ155 (dry): same locality.OTHER MATERIAL: QLD- QMG304737, QMGL864 (fragment NTMZ1528), QMGL2759 (fragment NTMZ1566). TAS- QMG300265 (NCIQ66C-3558-J) (fragment NTMZ3783). S AUST- SAMTS4018 (fragments NTMZ I 606, QMG300475). HABITAT DISTRIBUTION. Sand, rock and coral rubble substrata; 15-32m depth; Turtle Is, Agincourt Reefs (FNQ); Moreton I. (SEQ); IIlawarra (NSW); Westemport Bay (Vic); Kent Is, Bass Strait (Tas); Kangaroo I. (SA) (Fig. 294D). DESCRIPTION. Shape. Tubulo-digitate, massive, erect, up to 120mm high, attached directly to substrate, with erect, regularly cylindrical or irregular, stoloniferous, thin or thick, bulbous digits tapering towards their extremities, up to 45mm high, 22mm diameter. Colour. Bright orange, yellow-orange, or vermilion-red alive (Munsell lOR 6/10 - 2.5R 5/10), pale brown in ethanol. Oscules. Large, up to lOmm diameter, at apex or subapical on each digit. Texture and surface characteristics. Firm, rubbery, 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 surface, in irregular plumose formations, with or without light arenaceous ectosomal layer; choanosomal fibres immediately below ectosome. Choanosome. Regularly isodictyal to irregularly reticulate skeletal architecture, with poorly differentiated primary and secondary fibres; primary ascending fibres thin, 42-93p.m diameter, heavy, paucispicular, sometimes uni- or aspicular, occasionally meandering, sinuous, often ascending directly to periphery, interconnected at irregular intervals by thin, more-or-less transverse, uni- or aspicular secondary fibres, 18-35Rm diameter; secondary fibres branch and anastomose amongst themselves, forming oval, rectangular or triangular, relatively cavernous meshes, 124-650Rm 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, 72113p.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)-128Rm, width 4-(6.6)-8.5p.m. Auxiliary styles coring fibres and interstitial long, thin, straight or slightly curved hastate styles, occasionally modified to asymmetrical styloids, tomotes or strongyles, rarely sinuous. Length 108-(226.1)-305p.m, width 142.2)4.511m. Microscleres absent. ^ REVISION OF M1CROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 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 62-170p,m diameter, con- algal filaments within fibres, almost completely taming abundant collagen but no larval spicules. obscuring spicules within fibre skeleton; Wieden- 542zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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. styles echinating fibres indicate that it belongs to Echinochalina. Interpretation of these features is 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 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 commonly found inside fibres, whereas shorter, thicker principal styles are most frequently found REVISION OF M1CROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 543 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 thicker 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 E. (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: AB1P7B-6A. 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, p1.12, fig.1, text-fig. 60 [Barbados, West Indies]. MATERIAL. HOLOTYPE: ZMAPOR5509. Caribbean. Echinochalina (Protophlitaspongia) Burton, 1934 Protophlitaspongia Burton, 1934a: 562. [Not Protophlitaspongia; de Laubenfels, 1954: 96; Pulitzer-Final, 1986: 138]. TYPE SPECIES. Siphonochalina bispiculata 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 (coring and echinating fibres) appear to be diactinal, unlike all other microcionids, but they are interpreted here as being highly modified monactinal spicules allowing its inclusion in the Microcionidae. This interpretation is supported by the more-or-less plumose ascending, 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 coring and echinating megascleres are equated here with auxiliary and principal spicules, respectively, 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 (1934a) assigned the type to Microcionidae, for similar reasons as those outlined above, whereas de Laubenfels (1936a) referred it to the Desmacididae because, he suggested, the hastate diactinal megascleres resembled those of Guitarra and Liosina, although 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 (1936a) (P. aga, P. ada and P. antillana) are more appropriately placed in Desmacididae 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 & Levi, 1993a: 1279). Echinochalina (Protophlitaspongia) bispiculata (Dendy, 1895) (Figs 296-297, Table 45) Siphonochalina bispiculata Dendy,1895: 246. Diplodermia bispiculata; Hallmann, 1912: 255. Protophlitaspongia bispiculata; Burton, 1934a: 562. Echinochalina bispiculata; Hooper & Levi, 1993a: 1279; Hooper & Wiedenmayer, 1994: 277. MATERIAL. LECTOTYPE: NMVG2319: 5-6km from Point Lonsdale, Port Phillip Heads, Vic, 38°20.5'S, 144°35.6'E, 34-38m depth, 1894, coll. J.B. Wilson (dredge). PARALECTOTYPE: NMVG2320 (fragment BMNH1902.10.18.14.110): same locality. OTHER MATERIAL: VIC - QMG304102. HABITAT DISTRIBUTION. Rock reef; 34-38m depth; Port Phillip (Vic) (Fig. 296D). DESCRIPTION. Shape. Massive, subspherical, irregularly lobate, 90mm maximum diameter, with short bulbous surface lobes, or subcylindrical, flattened digitate sponge, 160mm long, 28mm wide, 15mm thick. Colour. Pale yellow-brown in ethanol. Oscules. Small, 2-3nun diameter, scattered over surface, particularly on apex of surface lobes. MEMOIRS OF THE QUEENSLAND MUSEUM 544zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Texture and surface characteristics. Soft, compressible, slightly rubbery; surface optically even, minutely reticulate. Ectosome and subectosome. Microscopically hispid ectosome with scattered plumose brushes of long, thin auxiliary oxeas, arising from the points of peripheral fibres and paratangential to surface; choanosomal fibres immediately below surface although 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, 60-95 p.m diameter, forming distinctly plumose tracts within core of fibre, interconnected at more-orless regular intervals by slightly thinner secondary fibres, transverse, uni-, pauci- or aspicular fibres, 20-43Rm diameter; fibres relatively light, cored by long, thin auxiliary oxeas and echinated by short, stout principal oxeas; fibre anastomoses form cavern- FIG. 297. Echinochahna (Protophlitaspongia) bispiculata (Dendy) ous ovoid meshes, 180- (QMG304102). A, Choanosomal skeleton. B, Fibre characteristics. C, 540p.m diameter; mesohyl Auxiliary oxea. D, Principal oxea. E, Ends of oxeas. matrix heavy but only lightLarvae. Incubated parenchymella larvae small ly pigmented, with moderate quantities of spherical, with heavy mesohyl matrix, 210auxiliary spicules dispersed between fibres; 240p.m diameter, no larval megascleres. choanocyte chambers large, oval to elongate, 60REMARKS. Of all E. (Protophlitaspongia) this 130p.m diameter. species is the most reminiscent of Haplosclerida Megascleres (Table 45). Principal megascleres in skeletal characteristics (e.g., three dimensional echinating fibres straight, short, thick, invariably ectosomal architecture like Hemigellius (Niphatsmooth, hastate oxeas, with abruptly pointed or idae); choanosomal fibre construction similar to Callyspongia (Callyspongiidae)). Echinochalina telescoped ends. (P) bispiculata is slightly similar to E. (P.) oxeata Auxiliary megascleres coring fibres long, thin, but that species lacks true geometric differentiastraight, hastate or rarely fusiform oxeas. tion between coring and echinating spicules and has a different growth form (Table 46). Microscleres absent. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 545 A LC) FIG. 298. Echinochalina (Protophlitaspongia) co/lain 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 sp. nov. (Figs 298-299, Table 45, Plate 12D) MATERIAL. HOLOTYPE: QMG304120: Blue Lagoon, Lizard I., Cairns section, Great Barrier Reef, Qld, 14°41.0'S, 145°27.5'E, 9m depth, 3.iv.1994, coll. J.N.A. Hooper & S.Cook (SCUBA). HABITAT DISTRIBUTION. Coral pinnacles; 9m depth; Lizard I. (FNQ) (Fig. 298F). DESCRIPTION. Shape. Thickly encrusting, up to 12mm thick, bulbous in places on surface following contours of substrate. Colour. Dark red alive (Munsell 2.5R 4/10), light brown in ethanol. 546zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ 547 TABLE 45. Comparison in spicule dimensions between species of Echinochalina (Protophlitaspongia). Measurements in micrometres (N=25); comparative data from Hooper & Levi (1993).zyxwvutsrqponmlkjihgfedcbaZYXW E.( P.) E.( P.) E. t ube rosa E. t ube rosa E.( P.) E.( P.) oxeat a zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA bargibant i E.( P.) isaaci sp.nov.zyxwvutsrqpon E. collat a sp.nov. Hooper & Hooper & favulosa (Burton) (di. gi tatezyxwvutsrqpon (t yp i c al Levi Levi sp.nov. labout ei SPI CULE (QMG300685) (QMG301270) (QMGL2166) (New (New (SE Qld) Caledonia) Caledonia) (QMG305051)(BMNH1930.8 (QMG304120) morph) morph) .13.45) (GBR,Q1d) (GBR,Q1d.) (QMG300039) (QMG300030) (GBR,Q1d) (SE. Qld.) (SE. Qld.) Auxiliary (coring) oxeas 52-(75.1)- 55-(73.3)-98 101(121.2)- 252-(265.2)- 108 x 1.8- x 0.8-(1.9)- 166 x 3.5- 287 x 3-(3.6)- (2.9)-4.0 3.0 (6.1)-7.0 Ectosomal I 15-(156.2)- 144-(216.1)- auxiliary 194 x 1.0- 278 x 1.0- styles (1.7)-2.5 (2.4)-3.5 28-(33.5)-42 32-(46.7)-58 57-(79.7)- x 2.0-(2.5)- x 1.0-(2.0)- 4.0 2.5 Chelae absent Toxas absent Principal (echinating) oxeas 4.5 128-(132)164 x 3.5-(4.2)-7.0 212-(234.0)- absent 262 x 2-(2.8)- absent 197-(235.6)326 x 6-(8.7)11 102-(123.9)158 x 344.6)-7 94-(101.7)163 x 1.5(3.8)-7.5 212-(237.2)- 114-(187.7)- 142-(181.9)- 228 x 1.5- 264 x 1.5- 215 x 1.5- 3 (2.2)-3 (2.2)-3 122 x 4.5- 118(142.7)162 x 2-(3.4)- 94-(148.0)188 x 1.5- (6.2)-7.5 4 14-(15.5)-21 9-(10.7)-14 absent absent absent absent absent absent absent absent absent 14-(73.2)118 x 1-(1.6)- 18-(44.1)-64 x 0.5-(0.9)1.5 31-(44.9)-62 x 0.541.1)1.5 absent (4.2)-6 2 Oscules. Very small, I -2mm diameter, on apex of larger surface bulbs, surrounded by radiating subectosomal drainage canals. Texture and surface characteristics. Harsh, firm, compressible; surface microconulose, minutely shaggy, conules close-set, less than 2mm high, surface silty in situ with radiating drainage canals clear of silt. Ectosome and subectosome. 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, 45-12011m diameter, forming oval or elongate meshes, 1204401..im diameter, without marked difference in mesh size between periphery and core of skeleton; ascending primary fibres cored by multispicular tracts of auxiliary spicules but occupying only about 5 0% of fibre diameter, interconnected by uni- or paucispicular secondary, 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 between fibres; choanocyte chambers large, elongate, 30-60p.m diameter. 54- ( 71.2) - 84 x 242.8)-4 (2.2)-3 67-(76.2)-88 x 1.5-(2.4)3.5 Megascleres (refer to Table 45 for dimensions). Principal (?) spicules echinating fibres short oxeas, 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 telescoped points. Microscleres (Table 45). Toxas intermediate between oxhorn 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, multispicular 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. (P.) tuberosa in having a special category of ectosomal styles and oxhorn toxas, although spicule geometry and spicule sizes are substantially different between these species. Further comparisons between species of E. (Protophlitaspongia) are given in Table 46. 548zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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 substrata; 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 MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 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. 550^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MEMOIRS OF THE QUEENSLAND MUSEUM Texture and surface characteristics. Soft, compressible; surface 'honeycomb' reticulate, highly porous, uneven, conulose. Ectosome and subectosome. Membraneous, minutely conulose surface with erect, paucispicular brushes of auxiliary megascleres arising from choanosomal spicule tracts protruding into surface conules but rarely through surface; some detritus scattered over surface; choanosomal fibres immediately below ectosomal skeleton. Choanosome. Regularly reticulate, subisodictyal skeletal architecture composed of more-or-less ascending, multi- or paucispicular, primary fibres, 22-43pm diameter, interconnected at irregular intervals by transverse, uni-, pauci- or aspicular fibres, 14-251J.m diameter; fibres cored by longer auxiliary megascleres and moderately echinated by shorter, principal spicules; fibre anastomoses form relatively wide, elongate or hexagonal meshes, 92-236p.m diameter; mesohyl matrix heavy, relatively darkly pigmented, granular, with auxiliary and principal megascleres dispersed between fibres; choanocyte chambers small ovoid to elongate, 22-46p.m diameter. Megascleres (Table 45). Principal megascleres echinating fibres relatively short, thick, straight, hastate oxeas, sometimes slightly telescoped, occasionally 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-monactinal forms, rarely true styles, longer forms generally thinner. Microscleres (Table 45). Palmate isochelae abundant, small, with approximately 15% twisted forms, lateral alae completely fused to shaft, front ala partially detached from lateral alae, shaft 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 asymmetrical intermediate geometries, and coring and echinating spicules differing only marginally in their length and thickness. The possession of isochelae microscleres in the present species sup- ports the inclusion of the genus in the Poecilosclerida, although it is still feasible that this species, and Protophlitaspongia, may be more appropriately assigned to the poecilosclerid Desmacididae (de Laubenfels (1936a) and followed by Pulitzer-Finali (1986)), but little support for its inclusion in the haplosclerid Niphatidae (as proposed for Isodictya by Hajdu et al., 1994b). Echinochalina (Protophlitaspongia) isaaci sp. nov. (Figs 302-303, Table 45, Plate 12E) MATERIAL. HOLOTYPE: QMG305051: W side Oyster Reef, Cairns section, Great Barrier Reef, Q1d, 16°38.4'S 145°54.7'E 20m depth, 21.ii.1995, coll. J.N.A. Hooper (SCUBA). PARATYPE: QMG305464: W. side Gannett Cay, Swain Reef, Mackay section, Great Barrier Reef, Qld., 21 0 58.68S, 152°28.34'E, 22m depth, 24.vii.1995, coll. J.N.A. Hooper & P. Tomkins (SCUBA).0THER MATERIAL: QLDQMG305398, QMG305430, QMG305504. HABITAT DISTRIBUTION. Fringing coral reefs, coral pinnacles, outer reef slopes, on dead coral; 2033m depth; Oyster Reef (FNQ); Swain Reefs (MEQ) (Fig. 302E). DESCRIPTION. Shape. Arborescent, digitate, branching, up to 150mm high, with conical cylindrical branches, up to 80mm long, 15mm diameter, bifurcate at tips and expanded towards apex of branches in life, collapses producing flattened branches in ethanol; basal stalk up to 25mm long, 12mm diameter, with expanded basal attachment. Colour. Pale orange-brown alive (Munsell 5R 7/8), light brown in ethanol. Oscules. Large, up to 15mm 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, spongy 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, discrete brushes of ectosomal auxiliary subtylostyles arising from ends of ascending choanosomal tracts; ectosome more heavily collagenous than choanosome, lightly pigmented; subectosomal skeleton slightly cavernous, with widely spaced peripheral fibres and skeletal tracts ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT 551zyxwvutsrqp ABCzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONM FIG. 302. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Echinochalina ( Prot ophlit aspongia) 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. 552zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 303. Echinochalina (Protophlhaspongia) 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 and distribution, 40-70Rm diameter, without any periphery.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA marked differences between primary and secondary fibres; fibres light; fibre meshes wide, oval Choanosom e. Irregularly reticulate, wisemeshed fibre skeleton, cavernous choanosomal to elongate, up to 650iLm diameter, slightly more structure; fibres relatively homogeneous in size cavernous in peripheral skeleton than at core; REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 553 ascending primary fibres cored by pauci- or multispicular tracts of auxiliary oxeas; smaller connecting 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; mesohyl matrix light, virtually unpigmented, with sparse auxiliary oxeas scattered between fibres; choanocyte chambers large, oval, up to 6011.m diameter. Megascleres (Table 45). Principal oxeas echinating fibres short, slender, straight, usually with telescoped points. Auxiliary oxeas coring fibres long, slender, straight, usually with telescoped points. Ectosomal 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 Stephen Cook, collector of many of the samples studied here. REMARKS. This species has a cavernous, irregularly reticulate skeletal structure with relatively poorly developed fibre system and mostly dominated by more-or-less plumose tracts coring major fibres. The ectosomal skeleton is particularly well developed, moreso than other species of E. (Protophlitaspongia), whereas the echinating spicule skeleton is very sparse, nearly vestigial. The species is most closely related to E. (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 E. (Protophlitaspongia) in Table 46. Echinochalina (Protophlitaspongia) oxeata (Burton, 1934) (Fig. 304, Table 45) Protophlitaspongia oxeata Burton, 1934a: 562, text- fig. 128. Echinochalina oxeata; Hooper & Levi, 1993a: 1279; Hooper & Wiedenmayer, 1994: 278. DESCRIPTION. Shape. Branching, cylindrical sponge 3-5mm 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, elastic; surface smooth, even, porous. Ectosome and subectosome. Membraneous, with sparse, more-or-less erect bundles of auxiliary oxeas from ascending choanosomal tracts protruding through surface; choanosomal fibres immediately below ectosome. Choanosome. Irregularly reticulate skeleton with differentiated primary and secondary spongin fibres; primary fibres ascending, pauci- or multispicular, 35-55p.,m diameter, interconnected by transverse, uni- or aspicular secondary fibres, up to 22iim diameter; fibre anastomoses produce large, cavernous, oval meshes in peripheral skeleton, up to 550iLm diameter, smaller, oval close-meshed reticulation at core of skeleton, up to 145Rm diameter; all spicule tracts occupy only small proportion of fibre diameter; differentiated ethinating 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; choanocyte chambers large, oval 45-951J,m diameter. Megascleres (Table 45). Principal oxeas coring fibres short, slender, straight or slightly curved at centre, with hastate or slightly telescoped points, occasionally modified to styles. Microscleres absent. REMARKS. This species has reduced spiculation, lacking either echinating (principal) spicules or ectosomal styles. In spongin fibre structure, distribution of spicules within fibres, ectosomal spiculation, it is close to E. (P.) bispiculata. Hooper & Levi's (1993a: 1279) E. (P.) oxeata from Moreton Bay is not conspecific with the holotype of E. (P.) tube rosa, 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: BMNH1930.8. 13.45: Papuan Pass, Cooktown region, Great Barrier Reef, Qld, 15°47'S, 145°48'E, 40m depth, coll. GBR Expedition (dredge). MATERIAL. HOLOTYPE: QMG300039: Inner HABITAT DISTRIBUTION. On foraminifera and coral fragments, 40m depth; Cooktown (FNQ) (Fig. 304C). Gneerings Shoals, off Mooloolaba, SE. Qld, 26°38.5'S, 153°09.5'E, 10m depth, 10.xii.1991, coll. J.N.A Hooper & S.D. Cook (SCUBA). PARATYPE: QMG300082: Shag Rock, N. Stradbroke I., Moreton MEMOIRS OF THE QUEENSLAND MUSEUM ^ 554zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 0 Echinochalina ( Prot ophlit aspongia) oxeat a (Burton) (holotype BMNH1930.13.45a). A, Auxiliary FIG. 304. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA oxeas. B, Section through peripheral skeleton. C, Australian distribution. D, Fibre characteristics. E, Choanosomal skeleton. F, Holotype. Bay, SE. Q1d, 27°25.0'S, 153°31.4'E, 15m depth, 05.ii.1992, coll. J.N.A. Hooper & J. Wilkinson (SCUBA).0THER MATERIAL: QLD- QMG304015. HABITAT DISTRIBUTION. Rock and coral reef, usually sheltered under ledges; 9-25m depth; Stradbroke I., 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, slightly hispid; surface uneven with prominent bulbous ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 555zyxwvutsrqp B C 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, Oxhom toxas. E, Section through peripheral skeleton. F, Australian distribution. G, Holotype. MEMOIRS OF THE QUEENSLAND MUSEUM 556zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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, Oxhom toxa. protruberances, up to 9mm high, 5mm diameter, auxiliary subtylostyles perched on surface, overinflated at ends. lying protruding principal oxeas from peripheral, Ectosome and subectosome. Relatively thick paratangential brushes composed of ectosomal ascending choanosomal skeletal tracts; REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP ^ 557 choanosomal fibres immediately below ectosome. Choanosome. Regularly reticulate skeletal structure, without any marked compression at core, with nearly renieroid fibre skeleton and plumoreticulate spicule tracts diverging towards periphery; spongin fibres heavy, without noticeable size differences between primary or secondary fibres, 20-60Rm diameter; all fibres cored by auxiliary oxeas and moderately heavily echinated by principal oxeas, particularly at fibre nodes; primary fibres with ascending multispicular tracts of auxiliary oxeas terminating in sparse bundles at surface; secondary fibres paucior unispicular in peripheral skeleton, uni- or bispicular at core of skeleton; fibre anastomoses produce square or round, relatively even meshes throughout skeleton, 70-15011.m diameter; mesohyl matrix moderately heavy, with few auxiliary spicules scattered between fibres; choanocyte chambers small, oval, 25-40tim diameter. Megascleres (Table 45). Principal oxeas echinating 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 curvature, points only slightly reflexed. Digitate growth form (Figs 307-308, Table 45) MATERIAL. QMG300030: Outer Gneerings Shoals, off Mooloolaba, SE. ()id, 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 bifurcate, occasionally anastomosing branches, 35mm 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, compressible, slightly rubbery; surface of branches smooth, macroscopically even, microscopically slightly hispid. Ectosome and subectosome. Thick, erect bundles of ectosomal auxiliary subtylostyles forming discrete plumose brushes on surface, usually at ends of ascending choanosomal spicule tracts, intermingled with protruding auxiliary coring oxeas; subectosomal region relatively thick, spiculose, with no fibre component. Choanosome. Regularly reticulate skeletal structure without marked compression of axis. Mega- and Microscleres. As for typical form. ETYMOLOGY. For the bulbous encrusting form. REMARKS. There are several notable differences 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 characteristics (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 protruding 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 characteristics 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) bargibanti Hooper & Levi, 1993zyxwvutsrqponmlkjihgfedcba Echinochalina ( Pr ot ophlit aspongia) bar gibant i Hooper & Levi, 1993a: 1280-1283, figs 31-32 [New Caledonia]. MATERIAL HOLOTYPE: QMG301270. SW. Pacific. Echinochalina (Protophlitaspongia) laboutei Hooper & Levi, 1993 558zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVU AB CD 0,1 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 & Levi, 1993a: 1277-1280, figs 29-30 [New Caledonia]. MATERIAL. HOLOTYPE: QMG300685. SW. Pacific. Artemisina Vosmaer, 1885 Artemisina Vosmaer, 1885a: 25; Ridley & Dendy, 1887: 112; Topsent, 1894a: 12; Lundbeck, 1905: 110; Burton, 1930a: 501, 528-531; de Laubenfels, ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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; Levi, 1960a: 61, 83; Ristau, 1978: 585; TYPE SPECIES. Artemisina suberitoides Vosmaer, Van Soest, 1984b: 122, 130. ^ 1885a: 25 (by monotypy) (junior synonym of Suberites Artenisina; Burton, 1934b: 54 Ilapsusj. ^arciger Schmidt, 1870:47 (Burton, 1930a: 528)). Qasimella Thomas, 1974: 311. MEMOIRS OF THE QUEENSLAND MUSEUM 560^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 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 moreor-less confused halichondroid reticulation of vaguely multispicular ascending and scattered transverse megascleres; echinating megascleres absent; subectosomal peripheral skeleton more radially arranged; ectosome membraneous, skinlike, with smooth styles of a single size category protruding through surface, forming paratangential 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, p1.19, 6g.10, p1.25,fig.2. A rtemisina apollinis; Lundbeck, 1905: 114-116, p1.13, fig.4; Kirkpatrick, 1908b: 34, p1.20, fig.4; Hentschel, 1914: 70; Topsent, 1917: 62, p1.4, fig.6, p1.6, fig.7; Burton, 1929a: 431; Hentschel, 1929: 876, 939; Burton, 1930a: 529; Burton, 1932a: 323; Koltun, 1964a: 73; Desqueyroux, 1975: 66, p1.4, figs 42-46; Koltun, 1976: 188; Hooper & Wiedenmayer, 1994: 256. Artemisina dianae Topsent, 1907:70-72; Topsent, 1908: 22, p1.3, fig.4, p1.5, 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 substrata; 18-380m depth; widespread boreal and antiboreal: Antarctica - MacRobertson Land, Australian Antarctic Territory (Koltun, 1976), Graham Land, William 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 (Topsent, 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, compressible, friable; surface uneven, folded, shaggy, conulose, cavernous, with skin-like ectosomal membrane stretched across adjacent conules. 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 principal spicules protruding through surface from underlying skeleton. Choanosome. Skeletal architecture irregularly reticulate or renieroid reticulate in places with rectangular and triangular meshes, up to 280i.tm diameter, made of uni- or paucispicular tracts of choanosomal principal styles; skeletal tracts not definitely differentiated into primary or secondary 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 collagen; 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)-458p.m, width 11-(14.0)-17p.m. Subectosomal auxiliary styles long, slender, straight, with slightly subtylote, lightly microspined bases, occasionally smooth, tapering to fusiform points. Length 2094244.6)293p.m, width 3-(4.4)-7pm. Microscleres. Palmate isochelae small, well silicified, unmodified. Length 8-(12.3)-16p.m. Toxas wing-shaped, variable in length and thickness, with sharply angular, deep central curvature, long straight arms on larger spicules, or slightly recurved arms on smaller spicules; largest with microspined points. Length 24(123.2)-26411m, width 0.8-(2.3)-5p.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 consequently Koltun's proposed synonymy is maintained here. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 561 FIG. 309. Artemisina 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) Artemisinajovis Dendy, 1924a: 343, p1.12, fig.6; Burton, 1930a: 530; Koltun, 1964a: 74-75; Bergquist & Fromont, 1988: 119-120, p1.56, 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. 311E); also North Cape and Three Kings Is, New Zealand (Dendy, 1924a; Bergquist & Fromont, 1988). DESCRIPTION. Shape. Flabellate, stalked, bifurcate digitate margins, 120-300mm high, 80110mm wide, 8mm thick lamellae. Colour Greyish in ethanol. Oscules. Oscules up to 6mm diameter, with slightly raised margins, clustered on osculiferous face of sponge. 562zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ 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, compressible; inhalant surface smooth, slightly undulating, with clearly visible porous ectosomal membrane stretched across pitted subectosomal region; exhalant surface osculiferous. Ectosome and subectosome. Erect plumose brushes of subectosomal auxiliary styles producing continuous palisade at surface but markedly cavernous in subectosomal region. Choanosome. Nearly regularly renieroid reticulate skeletal architecture composed of uni-, pauciand multispicular tracts of choanosomal principal styles producing triangular meshes up to 35011m diameter, enclosed within moderately light collagen; fibres absent; spicule tracts relatively homogeneous, undifferentiated into primary or secondary elements; mesohyl matrix heavy, darkly pigmented, containing abundant microscleres. 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)-448[Lm, width 12-(19.2)-26p,m. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 563 FIG. 311. Artemisina jovis Dendy (holotype BMNHI923.10.1.112). 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, Toxas intermediate accolada-wing shaped, fusiform pointed, straight or slightly curved near long, slender, with slightly rounded central curbasal end, with smooth round bases. Length 234- vature, slightly reflexed arms, microspined points. Length 984155.2)-215 p.m, width 1.5(290.6)-375p,m, width 3-(7.1)-10Rm.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB (2.4)-4p,m. Micr oscler es. Palmate isochelae unmodified, abundant, well silicified. Length 10412.5)- REMARKS. The type material revealed some 1611.m. discrepancies from published descriptions of MEMOIRS OF THE QUEENSLAND MUSEUM 564zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ both Dendy (1924a) and Bergquist & Fromont (1988), indicating variability between specimens. In particular skeletal meshes, described as being differentiated into ascending and transverse components, 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 spination of other megascleres. Artemisina plumosa Hentschel, 1914 (Fig. 312) Artemisina plurnosa Hentschel, 1914: 70-72, p1.4, fig.5, p1.6, fig.1 [et var. lipochela]; Burton, 1929a: 431 [Antarctica]; Burton, 1930a: 530; Koltun, 1964a: 73, p1.11, figs 1-8; Koltun, 1976: 190; Hooper & Wiedenmayer, 1994: 257. Artemisina strongyla Hentschel, 1914: 72-73, p1.6, fig.2. MATERIAL. HOLOTYPE: HM (fragments BMNH1933.7.20.3, ZMB4762): Off Wilhelm II Coast, Antarctica. HABITAT DISTRIBUTION. Soft substrata; 1252269m depth; Antarctica - Enderby Land, Sabrina Coast, Princess Elizabeth Land, Wilhelm II 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 choanosomal principal styles protruding, forming nearl continuous palisade on surface; subectosomal auxiliary styles tangential or paratangential to surface, sparse, dispersed between protruding bundles of choanosomal styles. Choanosome. 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 at core with diminished skeletal organisation whereby only pauci- or small multispicular tracts form skeletal reticulation at core, with square or triangular meshes, up to 3001Jim diameter; mesohyl matrix heavy, lightly 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)-472Rm, width 11413.7)18p.m. Subectosomal auxiliary styles or tornostyles short, thin, straight, with slightly subtylote bases, smooth or microspined bases, rounded, hastate or mucronate points often with small spines, or sometimes with single small terminal spine, or quasi-diactinal asymmetrical ends. Length 181(196.7)-268p.m, width 3-(6.1)-8p,m. Microscleres. Palmate isochelae very abundant, moderately large, well silicified, occasionally contort. Length 8-(13.2)-16p,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)296itm, width 0.8-(2.5)-5Rm. REMARKS. Artemisina plumosa differs from other species in having quasi-diactinal auxiliary megascleres, including some with basal spination. Its toxas are also much larger and more 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 (1964a) recorded a second category of large choanosomal style, with similar geometry to principal spicules but nearly twice their length (800-2160x21-44p,m), but these were not seen in the fragment of the holotype examined. Artemisina tubulosa Koltun, 1964 (Figs 313-314) Artemisina tubulosa Koltun, 1964a: 74, text-fig.18; Hooper & Wiedenmayer, 1994: 257. MATERIAL. HOLOTYPE: BMNH1963.7. 29.52: Mawson Coast, Australian Antarctic Territory, 100m depth. HABITAT DISTRIBUTION. Hard benthos; 100m depth; Australian Antarctic Territory (Fig. 313F). DESCRIPTION. Shape. Tubulo-digitate, hollow, cylindrical, 100mm long, 30mm diameter. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 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 multispicular 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 irregular uni-, paucispicular or smaller, transverse multispicular tracts interconnecting main tracts, together producing a nearly myxillid-like triangular reticulation, with meshes up to 8001.i.m 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, fusifonin, rounded bases, smooth, slightly curved near basal end. Length 7054778.8)874p,m, width 18-(30.2)-39p,m. Subectosomal auxiliary styles short, thin, mostly straight, with rounded or very slightly subtylote or occasionally polytylote bases, typi- MEMOIRS OF THE QUEENSLAND MUSEUM 566zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ cally microspined, and with blunt hastate microspined points. Length 384-(456.3)-494p.m, width 6-(8.4)-11p.m. Microscleres. Palmate isochelae abundant, moderately large, well silicified, unmodified. Length 12-(15.8)-19p,m. Toxas intermediate accolada-wing shaped, long, thick, angular central curvature, spined points. Length 602-(674.2)-824p,m, width 4(6.4)-91.1,m. REMARKS. Artemisina tubulosa is similar to A. plumosa in skeletal architecture and spicule geometry, and the two are probably at least sibling 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, skeletal 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 archegona Ristau, 1978b: 585-587, textfigs 2F, 3F, 6C-D [Carmel, California]. MATERIAL. HOLOTYPE: USNM24528. Province: NE. Pacific. Artemisina arciger (Schmidt, 1870) Suberites arciger Schmidt, 1870: 47, p1.5, fig.6. Artemisina suberitoides Vosmaer, 1885a: 25-26, p1.1, fig. 1 6, p1.4, figs 11-14, p1.5, figs 51-55 [Arctic]; Ridley & Dendy, 1887: 112 [off Nova Scotia]; Fristedt, 1887: 430, p1.24, figs 15-17; Whiteaves, 1901: 17; Lundbeck, 1905: 113. Artemisina arcigera; Lundbeck, 1905: 110-114, p1.1, figs 9-11, p1.13, fig.3; Arndt, 1913: 115; Topsent, 1913b: 47, p1.1, fig.6; Rezvoi, 1925: 197, [var. spiceps; Arctic]; Burton, 1930a: 528-529 [typespecies]; Hentschel, 1929: 876, 938 [Arctic]. Artemisina arciger; Burton, 1959b: 42 [Iceland]; Van Soest & Stone, 1986: 47 [note]. MATERIAL. HOLOTYPE: LMJG, fragment BMNH1870.5.3.90; holotype of suberitoides: ZMA P0R443, fragments BMNH1901.1.1604, 1351. Province: NW. Atlantic, NE. Atlantic, Arctic. Artemisina elegantula Dendy, 1924 Artemisina elegantula Dendy, 1924a: 344 lN.of New Zealand]; Burton, 1930a: 531; Bergquist & Fromont, 1988: 120; Dawson, 1993: 36 [index]; Levi, 1994: 36, fig.206, p1.7, fig.1 [S. of New Caledonia]. MATERIAL. HOLOTYPE: BMNH1923.10. 1.113. New Zealand, New Caledonia. Artemisina erecta Topsent, 1904 Artemisina erecta Topsent, 1904a: 214-215, p1.5, fig.18, p1.15, fig.10 [Azores]; Burton, 1930a: 530. MATERIAL. HOLOTYPE: MOM. NE . Atlantic. Artemisina hispanica Ferrer-Hernnadez, 1918 Artemisina hispanica Ferrer-Hernandez, 1918: 536538, fig.3, text-fig.5 [Santander]; Levi, 1960a: 83 [Santander, Mediterranean]; Burton, 1930a: 530. MATERIAL. HOLOTYPE: Madrid. Mediterranean. Artemisina indica (Thomas, 1974) Qasimella indica Thomas, 1974: 311-312, text-fig.1 [Gulf of Manaar]. MATERIAL. HOLOTYPE: CMFRIT84/1, paratype CMFR1T84/2. N. central Indian Ocean. Artemisina melana Van Soest, 1984 Artemisina melana Van Soest, 1984b: 122-124, p1.8, figs 7-8, text-fig.49 [Curacao]; Zea, 1987: 173, textfig.61, p1.11, figs 3-4 [Columbian Caribbean]. MATERIAL. HOLOTYPE: ZMAPOR4881, paratype ZMAPOR4882, 4883. Caribbean. Artemisina stipitata Koltun, 1958 Artemisina stipitata Koltun, 1958: 52-53, text-fig. I 0 [Kurile Strait]. MATERIAL. HOLOTYPE: BMNH1963.7. 29.39. NW. Pacific. Artemisina transiens Topsent, 1890 Artemisina transiens Topsent, 1890b: 66 [nomen nudum]; Topsent, 1892a: 94, p1.2, fig.5, p1.4, fig.8, p1.9, fig.18 [Atlantic coast of France]; FerrerHernandez, 1914: 3; Burton, 1930a: 529-530; Levi, 1960a: 83-84 [Asturies, Mediterranean]. MATERIAL. HOLOTYPE: MOM. NE . Atlantic, Mediterranean. Artemisina vulcani Levi, 1963 Artemisitza vulcani Levi, 1963: 64-65, p1.9F, textfig.74 [South Africa]. MATERIAL. HOLOTYPE: MNHNDCL622. 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 [possible generic synonymy]. Esperiopsis intermedia; Van Soest & Stone, 1986: 45 MATERIAL. HOLOTYPE: BMNH1910.1. 1.912. Note: referred to Desmacididae, Esperiopsis. Amphilectus pilosus Ridley & Dendy, 1886: 350; Ridley & Dendy, 1887: 126-127, p1.19, fig.5, p1.25, fig.3 [Kerguelen Is; Marion I.]. Lissodendoryx pilosus; Topsent, 1901: 20. Megaciella pilosa; Hallmann, 1920: 772; Van Soest, 1984b: 128 [possibly Myxillidae]. ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS 567 0 0 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 [Brazilian Basin]. affinities].^ MATERIAL. HOLOTYPE: BMNH1887.5.2. 124. Note: Cornulum tylota; Van Soest, 1984b: 124. - referred to Iophonidae, Megaciella. MEMOIRS OF THE QUEENSLAND MUSEUM ^ 568zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA MATERIAL. ^HOLOTYPE: MNHNDNBE943. Note: referred to Iophonidae, Comulum. Incertae sedis Pandaros Duchassaing & Michelotti, 1864zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Duchassaing & Michelotti, 1864: 88; Schmidt, 1870: 59; de Laubenfels, 1936a: 123; Wiedenmayer, 1977: 143; Van Soest, 1984b: 127. Panda ros TYPE SPECIES. Pandar os acan- Duchassaing & Michelotti, 1864: 90 (by subsequent designation ofde Laubenfels, 1936a: 123). t hifolium DEFINITION. Well-developed choanosomal skeletal reticulation of spongin fibres woven into flattened anastomosing branches, or forming continuous sheets without any regular architecture; fibres cored by auxiliary styles, subtylostyles to tylostyles, predominantly smooth, distributed within fibres in criss-cross fashion; fibres echinated by rare acanthostyles, rarely smooth styles; few anisotornotes, verging on oxeas, scattered throughout mesohyl; sparsely dispersed tangential subectosomal auxiliary spicules dispersed throughout peripheral skeleton; microscleres absent. REMARKS. No Australian species. SPECIES OF PANDAROS Pandaros acanthifolium Duchassaing & Michelotti, 1864 Duchassaing & Michelotti, 1864: 90, p1.20, fig.2 [St.Thomas, Virgin Is]; de Laubenfels, 1936a: 123 [Florida]; Hechtel, 1965: 45; Wiedenmayer, 1977: 144, p1.30, 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 Indies]; Tha, 1987: 175, p1.4, fig.4, text-fig.62 FIG. 314. Art em isina t ubulosa Koltun (holotype BMNH1963.7.27.52). A, Skeletal structure. B, Palmate isochelae and spined toxas. [Columbian Caribbean]. Pandar os acant hifolium REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU ^ 569 Pandaros arbusculum, in part, Duchassaing & Michelotti, 1864: 88, p1.18, fig.6 [St.Thomas, Virgin Is]. Thalyseutypon conulosa Hechtel, 1965:44, p1.6, fig.1, text-fig.9 [Port Royal; Jamaica]; Alcolado, 1976: 5 [Cuba]. MATERIAL. LECTOTYPE: TMPOR57 (fragments USNM31020, MNHNDNBEI309, BMNHI928.11.12.16), paralectotype: 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), 2(4); Hoshino, 1971: 24. Thalyseurypon kasumiensis; Hoshino, 1981: 159-161, text-fig.70 [Japan]. ? Pandaros kasumiensis; Hooper, 1988b: 353, 766. MATERIAL. HOLOTYPE: MMBSSIS-052. NW. Pacific. TRANSFERS List of other species described in Pandams but now transferred to another genus. Pandaros angulosa Duchassaing & Michelotti, 1864: 89, p1.19, fig.4 [St. Thomas, Virgin Is]. Mycale (Aegogropila) angulosa; Van Soest, 1984b: 16-19, p1.1, figs 5-6, text-fig.3 [plus synonymy]; Pulitzer-Finali, 1986: 130-133, figs 52-55. Not Mycale angulosa; de Laubenfels, 1936a: 116. Zygomycale parishii, in part; de Laubenfels, 1956: 3; Hechtel, 1965: 48, p1.5, fig.3; Alcolado, 1980: 4. MATERIAL. LECTOTYPE: TMPOR54 (fragments USNM31021, MNHNDNBE1310, BMNH1928.11.12.20). referred to Mycalidae, Mycale. Pandaros arbusculum Duchassaing & Michelotti, 1864: 88, p1.18, fig.6 [St. Thomas, Virgin Is]. Ptilocaulis sp. aff. spiculifer (Lamarck,1814); Wiedenmayer, 1977: 153, 251. Pidocaulis arbusculum; Van Soest et al., 1983: 204. MATERIAL. LECTOTYPE: ZMAPOR1728, paralectotype ZMAPOR1729. referred to Axinellidae, Ptilocaulis. Pandaros lugubris Duchassaing & Michelotti, 1864: 89 [St. Thomas, Virgin Is]. Echinodictyum lugubre; Wiedenmayer, 1977: 255; Pulitzer-Finali, 1986: 106-107. MATERIAL. LECTOTYPE: TMPOR55 (fragments USNM31023, MNHNDNBEI312, BMNH1928.11.12.21). referred to Raspailiidae. Pandaros pennata Duchassaing & Michelotti, 1864: 88, p1.20, fig.3 [St. Thomas, Virgin Is]. Echinodictyum pennatum; Wiedenmayer, 1977: 251, 255. MATERIAL. LECTOTYPE: TMPOR58 (fragments USNM31024, MNHNDNBE1313, BMNH1928.11.12.22), paralectotype: ZMA PORI731. referred to Raspailiidae, Echinodictyum. Pandaros walpersii Duchassaing & Michelotti, 1864: 90, p1.20, fig.4, text-fig.2D [St. Thomas, Virgin Is]. Ptilocaulis walpersi; Wiedenmayer, 1977: 255. Spongia spiculifera Lamarck, 1814: 449. Ptilocaulis spiculifera; Pulitzer-Finali, 1986: 103-104 [West Indies]. MATERIAL. LECTOTYPE: TMPOR56 (fragments USNM31025, MNHNDNBE1314, BMNH1928.11.12.17), paralectotype: IZUGCE 38.763. referred to Axinellidae, Ptilocaulis. 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 31 new species from Australia, and 3 from other localities for misidentified taxa, 18 new names for preoccupied taxa, and 162 new taxonomic combinations. Many species are also new records for Australia and 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 sampled. This revised classification is based on a reevaluation of all characters, including an analysis of their consistency within populations of several species, and places primary importance on skeletal 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 architecture (e.g., Levi, 1960a), megasclere form and distribution within the skeleton (Hallmann, 1920), ectosomal structure and composition (Van Soest, 1984a), microsclere geometry and diversity (e.g., de Laubenfels, 1936a), or combining all these characters into a a consensus matrix with a consequent proliferation of genera (e.g., Bergquist & Fromont, 1988). Re-examination of the type species of 73 nominal genera included in Microcionidae at one time or another found that many had been misdescribed, or important characters overlooked, with the consequence that previous interpretations of genera are mostly incorrect. Furthermore, many of these nominal genera are not valid in a phylogenetic sense given that their supposed apomorphic characters are unstable (inconsistent) within otherwise closely related species. These earlier classifications had extraordinary high levels of homoplasy (Hooper, 1990a), partly a consequence of relying on single morphological characters (which when used in isolation cut across a classification based on other single char- MEMOIRS OF THE QUEENSLAND MUSEUM 570^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA acters). I use combinations of characters to develop a phylogeny. At the species level several characters are prone to ecophenotypic variability (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 differences in skeletal characters are indicative of divergent, sibling species relationships, with some empirical support from biochemical and genetic data (Hooper et al., 1991). Phylogenetic analysis. Hypothesised phylogenetic 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 & Levi (1994). Determinations of plesiomorphies were largely made through outgroup comparision (Raspailiidae), but these interpretations are complicated by the prevalance of analogous characters throughout Porifera (or misinterpretation of homology by authors). Consequently some information from the fossil record is considered below to assist in determining character polarity. (1) Skeletal structure: Ordovician Saccospongia 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 & Levi, 1989), which also have a (desma-bearing) basal skeleton — a condition relatively widespread during the Ordovician (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 sistergroup relationship (Hooper, 1991), sharing a similar range of skeletal structures (including hymedesmoid, microcionid, renieroid, 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 representing convergence of a few characters. Raspailiidae was previously included in order 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 poecilosclerids. (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 plesiomorphic for the haplosclerids. However, similar features in some Microcionidae represent parallel development (e.g., quasi-diactinal ectosomal spicules in some Holopsamma, quasi-diactinal structural spicules in Echinochalina).The tangential layer of amphitylotes in the Myxillidae is unique to the group (e.g., Lissodendoryx), upon which Van Soest (1984b) based his classification, but the myxillid (tangential amphitylote) ectosomal skeleton inAcarnus is enigmatic since its microscleres are typical of Microcionidae. Thus Hooper (1988, 1990a) considered that microscleres were more easily modified, acquired independently or lost in the Poecilosclerida than was the ectosomal skeleton, which in this case defined the Myxillidae. Hajdu et al. (1994) thought that microsclere geometry was far more conservative than that of megascleres (which might be more susceptible to phenotypic influences). Less obvious is the polarity of the sequence Clathria-Dendrocia-Thalysias, varying from a virtually aspicular ectosome to a dense, continuous palisade of erect megascleres. I suggest that the latter condition is more plesiomorphic based on the `raspailiid ectosomal condition' (i.e., smaller auxiliary spicules surrounding bases of protruding larger subectosomal spicules) being common to the outgroup Raspailiidae, and diminishment of ectosomal structure or loss of spicules representing a secondary derivation. (3) Spicule axes: It could be argued that oxeote (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 oxeotes (in E. (Protophlitaspongia) for example) represent recent derivations from styloid forms (much like the derived tetractinal styloid modifications in Acamus (Myxillidae); Van Soest et al., 1991). ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP 571zyxwvutsrqpon 46. Comparison between species of Echinochaliria (Protophlitaspongia) in major morphological characters.Measurements in TABLE E. labout ei E. bargibant i zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC E. oxeat a E. favulosa E. t ube rosa E. isaaci E. collat a Hooper & Hooper & CHARACTER E. bispiculat a (Dendy) irregularly reticulate, Fibre skeleton structure detritus cavernous in (Burton) L6vi heavily reticulate, condensed distinct axis, regularly secondary reticulate fibres extra-axis rich rich poor elongate or cavernous, oval (180- oval (145- size) 450) 550) Fibre diameter 20 - 50 - - abundant mod. sparse mod, sparse well silicified well silicified periphery sp.nov. egularly irregularly primary and axis, more compact in sp.nov. 1.6vi plumoregularly reticulate throughout, cavernous meshes irregularly reticulate. reticulate, diverging wide meshes, near much detritus periphery, meshes even throughout Collagen in mesohyl Fibre mesh shape (and Spicules in skeletal elongate (50250) abundant poor Coring spicul es in form irregularly wide even reticulate, meshes even meshes, diverging fibres 2% near cored periphery very light abundant 180) (100-300) round (70150) (370-650) elongate (120-440) zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 22 55 15 90 8 40 20 50 20 60 40 70 45 120 multispicular - - very sparse poorly poorly silicified silicified on surface mod. mod. abundant abundant well silicified well silicified well silicified pauci- or bi- or uni- or paucispicular aspicular unispicular branching, cylindrical clumped cylindrical lobate- cylindrical, digitate, lobate- digit,honey- digitate, or digitate arborescent digitate combed encrusting surface digitate uni- or tubular digits !nod. paucispicular aspicular (4) Spicule ornamentation: Hypercalcified sponges ('sclerosponges') were major contributors 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 acanthostyles). Vacelet (1985) suggested that sclerosponges were polyphyletic having affinities with many demosponge groups, and if living species represent relict ancestral groups of Demospongiae, then there is argument to suggest that acanthose megascleres are ancestral. Conversely, smooth monaxons were predominant in Palaeozoic and Mesozoic sponges such as Saccospongia (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- - abundant pauci- or uni- or lobate, - multispicular aspicular massive- Growth regularly reticulate, zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONM square or oval or elongate elongate FirbrinesarY Coring i spicules n secondary fibres sp.nov square Or rounded (80- tracts Spicule silificat ion moderate sp.nov. rarely multispicular multispicular pauci- or multispicular - abundant well silicified multispicular uni- or uni- or unispicular uni- or aspicular paucispicular or aspicular paucispicular erect, digitate, erect, thickly arborescent, encrusting, cylindrical bulbous digits surface 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 interpretation 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) secondary loss (with the corollary that echinating spicules in Microcionidae is retention of an ancestral character). Similarly, smooth echinating or pseudo-echinating spicules, such as in Echinoclathria and Echinochalina, have probab- MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZY 572zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ /2 A 601 Artemisina Antho (A) Antho (P.) Antho (I.) Echinoclathria Holopsamma Echinochalina (E.) Echinochalina (P ) Pandaros Clathria (D.) Clathria (W.) Clathria (C.) Clathria (T ) Clathria (I.) Clathria (M.) Clathria (A.) 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 state 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 largely based on modifications to chelae (i.e., Microcionina with only palmate forms, Myxillina with modified bi- or tridentate modifications, Mycalina with sigmancistra derivatives). Palmate 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 archetypal palmate morphology. Similarly anchorate chelae were thought to have arisen from arcuate modifications to the primitive palmate 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 attachment between the lateral alae and shaft (i.e., a reduction from the existing fused structure). If this latter interpretation is correct (and it is energetically more probable than the acquisition of a 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 (Cl) is indicated for each character obtained from parsimony ananlysis (Swafford, 1991) ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 573zyxwvutsrqp zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIH skeleton without marked axial compression but well differenzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC tiated axial and extra-axial (radial, plumose or plumoreticu- 1. Growth form specialisation. (CI = 1.0) 1 . encrusting or bulbous-encrusting growth forms. 2.* massive, branching, lobate, flabellate, vasiform or tubular growth forms. 3. late) regions. 3.* choanosomal skeleton with noticeably specialised honeycomb-reticulate growth forms composed of compressed axis and well differentiated axial and extra-axial tightly anastomosing flattened fibre-branches (lacunae). (radial, plumose or plumoreticulate) regions. 4. choanosomal 2. Ectosomal skeleton specialisation. (CI = 0.6) 1 . mem- skeleton hymedesmoid or rnicrocionid, with basal layer of spongin braneous,skin-like exterior, with single category of (subec- lying on substrate (with or without ascending, non-anastomosing tosomal) auxiliary spicule extending from choanosomal skeleton fibre nodes), and bases of principal spicules standing perpendicular protruding through surface forming discrete paratangential or to substrate. 5. choanosomal skeleton evenly renieroid or isodictyal erect brushes. 2. with single category of (subectosomal) reticulate throughout with well developed spongin fibres cored by auxiliary spicule on surface forming tangential,paratangential smooth principal styles. or plumose tracts. 3. with two categories of auxiliary 7. Derivation of spicules coring fibres. (CI = 0.75) I .* spicules,smaller ectosomal spicules generally overlaying choanosomal fibres or skeletal tracts cored by one or more larger subectosomal spicules forming discrete bundles or category of principal spicules. 2. choanosomal fibres or continuous palisade on surface. 4• * with specialised ectosomal skeletal tracts cored by auxiliary spicules but partially or skeleton composed of smaller auxiliary spicules surrounding wholly replaced by detritus. 3. choanosomal fibres or skeletal bases of protruding larger subectosomal spicules. tracts cored by auxiliary spicules identical to those in ec- 3. Subectosomal skeleton structure. (CI = 0.5)1. radial,with tosomal and subectosomal skeletons. 4. choanosomal fibres perpendicular bundles or single spicules supporting ectosome. or skeletal tracts cored by auxiliary spicules different from 2.* plumose, plumoreticulate, or disorganised tracts of those in peripheral skeleton. spicules in peripheral skeleton. 4. Differentiation within choanosomal skeleton. (Cl = 8. Differentiation of primary and secondary skeletal components. (Cl = 1.0) I • * primary and secondary skeletons zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA undifferentiated. 2. primary skeleton renieroid cored by axial- 0.667) 1 . choanosomal skeleton more-or-less undifferentiated, unstructured. 2. choanosomal skeleton well structured, hy- ly or basally compressed tracts of acanthostyles,secondary medesmoid to reticulate, but lacking any differentiated com- skeleton cored by smooth principal styles in plumose, sub- ponents. 3.* choanosomal skeleton well structured, predominantly isodictyal or plumoreticulate tracts. 3. primary skeleton reticulate, differentiated into two distinct components. renieroid cored by axially or basally compressed tracts of 5. Fibre development and skeletal architecture. (CI = 0.714) acanthostrongyles,secondary skeleton cored by smooth prin- 1. choanosomal skeleton without spongin fibres (or indefinite cipal styles in plumose, subisodictyal or plumoreticulate fibres), with spicules forming more-or-less disorganised hali- tracts. 4 . primary skeleton axially compressed spongin fibres chondroid, vaguely ascending, longitudinal reticulate tracts. 2. cored by renieroid tracts of sparsely spined principal styles choanosomal skeleton with poorly developed spongin fibres, intermingled with plumose or plumoreticulate tracts of smooth primary (basal or axial) renieroid component cored by acanthose principal styles, overlaid by secondary extra-axial plumose principal spicules,secondary plumose, subisodictyal or plumo- skeleton cored by larger smooth principal styles. 5. primary reticulate component cored by smooth principal spicules. 3. renieroid reticulate skeleton cored by smooth principal styles choanosomal skeleton with well developed spongin fibres, and echinated by identical spicules, with secondary radial primary skeleton compressed, renieroid, cored smaller smooth extra-axial skeleton on exterior edge of skeleton only cored principal styles, secondary skeleton vestigial or greatly reduced, by larger smooth principal styles. consisting only of larger smooth principal styles embedded in 9. Derivation of echinating spicules. (CI = 0.444)1. echinat- peripheral fibres forming sparse radial tracts. 4 . choanosomal ing spicules absent. 2.* special category of (acantho)styles skeleton with well developed spongin fibres forming more-or- present echinating fibres, differentiated from principal less evenly reticulate or plumo-reticulate meshes throughout in spicules. 3. echinating spicules styles or acanthostyles undif- massive forms, or thick basal layer of spongin and hymedesmoid ferentiated from principal spicules coring spongin fibres. 4. structure in encrusting forms. 5. choanosomal skeleton with well echinating spicules styles or acanthostyles representing prin- developed spongin fibres woven into flattened reticulate cipal spicules, but different from those coring fibres. 5. branches or forming continuous sheets without any regular echinating spicules oxeas or anisoxeas representing principal architecture, cored by criss-cross of auxiliary styles. 6.* spicules, but different from those coring fibres. choanosomal skeleton with well developed spongin fibres form- 10. Modifications to chelae microscleres. (Cl = 0.5) / . ing compressed reticulate axis and well differentiated radial, chelae absent. 2.* isochelae palmate. 3. isochelae arcuate-like. plumose or plumo-reticulate extra-axis. choanosomal skeleton without any marked axial compression or 4 . isochelae anchorate-like. 1 1 . Ornamentation of toxa microscleres. (Cl = 0.667) 1. toxas absent. 2.* toxas with predominantly smooth points. 3. differentiated axial and extra-axial regions. 2. choanosomal toxas with predominantly spined points. 6. Compression of choanosomal skeleton. (Cl = 0.8) I . MEMOIRS OF THE QUEENSLAND MUSEUM 574^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA preceed arcuate chelae in the series from palmate to bidentate- or tridentate-derived chelae. However, the practical problem with the Hajdu et al. (1994a) proposal is that there is no sharp distinction 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 'teeth', 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' become a matter of degree rather than an absolute descriptor. Several Microcionidae genera have been established solely on the basis of bidentate- or tridentate-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., 1994a), 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 different 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.e., with thickened curved shaft and partially detached lateral alae, verging on 'arcuate') to anchorate-like chelae seen in two species (i.e. in which there are completely 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 discounted by Hajdu et al. (1994a), who suggested that anchorate and arcuate modifications to chelae were homologous within a single phylogeny containing bidentate-derived taxa). Interpretation of this transformation continuum in Clathria (Dendrocia) from 'typical' palmate to detached `anchorate' chelae suggests that detachment of the lateral alae from the shaft of the spicule, leaving the residual 'scar' or lateral ridge along the shaft, may be a simple reduction process occurring more than once in the group, and that `anchorateness' may not be homologous throughout the order. Other examples, such as the strongly unguiferous sigmoid chelae of Bipocillopsis and Damoseni, of indeterminable arcuate or anchorate derivation, are less easily accountable 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' (1936a) solution to this problem (in which species with modified isochelae have skeletons and spiculation otherwise structurally and geometrically identical to those containing 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' (1936a) classification had extraordinary high levels of homoplasy and the inferred relationships based on isochelae geometry cut across classifications based on both skeletal architecture (e.g., Hallmann, 1912, 1920) and ectosomal characteristics (e.g., Van Soest, 1984b). This proposal was rejected by most contemporary authors. Hooper (1990a) provided an alternative proposal that considered modified isochelae to be homoplastic, and a classification based primarily on this feature was both unparsimonious and incongruent with other structural and geometric features within the Poecilosclerida. Hajdu et al. (1994a) correctly noted that many cases of `arcuateness' 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 turned out to be merely modified palmate forms), and this is also true for most recorded instances within the Microcionidae. However, from evidence presented here it is likely that `anchorateness' and `arcuateness' may be a homoplasy for the Poecilosclerida, and consequently Hajdu et al. (1994a) major reorganisation of the Poecilosclerida requires further refinement, as to family composition. (7) Toxa morphology: Van Soest et al. (1991) set a precedent for interpretation of toxa geometry in which oxhorn-like toxas (including wing-shaped toxas) were considered to be an- REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 575 cestral because they are shared by outgroups such Myxillidae, whereas accolada-like toxas (including raphidiform and oxeote toxas) are more derived. (8) Growth form: Microcionids in areas of environmental 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 is considered here that in most cases an encrusting habit and a 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 Holopsamma, with may have some systematic value, whereas other growth forms occur throughout Porifera and are more difficult to interpret phylogenetically. Phylogenetic analysis (Fig. 315) shows two fundamental groups of genera (A, B) differentiated primarily by their skeletal structure (character 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 suggested myxillid-like features, most possessing a secondary renieroid skeleton. The other group (A) containing Clathria, Echinochalina, Holopsamma and Pandaros has suggested similarities to raspailiids, including skeletal structure of typical genera (e.g., compare Clathria (Axociella) and Raspailia) and atypical genera (e.g. compare Echinochalina (Echinochalina) and Echinodictyum). It is suggested that the `raspailoid' group (A) retains more ancestral features common to the outgroup, whereas the `myxilloid' group (B) is more derived. Within group (A) there are three taxa indicated as possibly polyphyletic (C. (Wilsonella), C. (Dendrocia), Pandaros). The first two may be simply more derived than other Clathria at the base of the tree, or they may represent full genera. The latter explanation is rejected given that the characters inferring polyphyly (partial substitution of coring megascleres by detritus in C. (Wilsonella), and loss of principal spicules completely in C. (Dendrocia)) are homoplasious representing convergences via functional acquisition 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, not 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 classification, following the precedent adopted for the Raspailiidae (Hooper, 1991), is admittedly partially artificial given the existence of these homoplasies, but no alternative is presently available 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 that 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 subgeneric 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); encrusting habit and plumose 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 E. (Echinochalina) and E. (Protophlitaspongia); presence or absence of ectosomal specialisation in C. (Thalysias) and C. (Clathria)). Within Clathria there are many species-groups that could be used to subdivide these taxa further (e.g., `spicata', `procera"phorbasiformis' groups; see also Hooper et al., 1991; Hooper & Levi, 1994). Many of these groups contain species that span across several subgenera and as such are of limited usefulness in phylogenetic reconstruction, but they are most useful in interpretation of sister-group relationships in a biogeographical context (Hooper & Levi, 1994), and these species groups will be considered further in a more detailed study Indo-west Pacific microcionids, 576^ MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 48. List of species included in Microcionidae with their current taxonomic assignments. SPECIES abietina Lamarck ORIGINAL GENUS Spongia abrolhosensis sp.nov. abyssorum Carter acanthifolium Duchassaing & Michelotti acanthodes Hentschel CURRENT ASSIGNMENT antarctica Topsent Anchinoe C. (Microciona) antarcticus Koltun [preocc.] Stylotellopsis synonym of C. (Thalysias) koltuni Hooper synonym of A.(Antho) dichotoma (Esper) anthoides Levi Clathria C. (Clathria) antyaja Burton & Rao Dendrocia C. (Clathria) Pandaros aphylla sp.nov. C. (Thalysias) C. (Wilsonella) Dictyocylindrus Pandaros Clathria synonym of C. (Thalysias) cactiformis (Lamarck) C. (Thalysias) apollinis Ridley & Dendy Amphilectus Artemisina appendiculata Lamarck Spongia synonym of C. (Thalysias) cactiformis (Lamarck) acantho styli Hoshino Thalysias C. (Clathria) acanthotoxa Stephens C. (Clathria) araiosa Hooper & Levi arborea Tanita Clathria (Thalysias) C. (Thalysias) Eurypon Litaspongia Echinoclathria acanthotoxa Levi & Levi [preocc.] Microciona see C (Microciona) claudei nom.nov. arborea Lendenfeld Plectispa Holopsamma aceratoobtusa Carter Microciona C. (Microciona) arborescens Ridley Ritaphidophlus C. (Thalysias) arbuscula Row Ophlitaspongia C. (Clathria) Clathria synonym of C. (Thalysias) abietina (Lamarck) arbusculum Duchassaing & Michelotti Pandaros Ptilocaulis (Axinellidae) aculeata Ridley adioristica de Laubenfels Dictyociona C. (Microciona) archegona Ristau Artemisina Artemisina affinis Carter Microciona C. (Microciona) arcifera Schmidt Tenacia affinis Topsent [preocc.] Hymeraphia see C. (Microciona) campecheae nom.nov. arciger Schmidt Sube rites Echinoclathria Artemisina C. (Clathria) Microciona C. (Microciona) arcuophora White legge Clathria africana Levi alata Dendy Clathria synonym of C. (Dendrocia) pyramida Lendenfeld arenifera Carter Echinoclathria synonym of Holopsamma laminaefavosa Carter amabilis Thiele Stylotellopsis C. (Thalysias) Microciona C. (Microciona) ambigua Bowerbank Microciona Plocamionida (Anchinoidae) artnata Bowerbank arteria de Laubenfels Axociella C. (Thalysias) amiranteiensis nom.nov. [for Colloclathria ramosa Dendy] C. (Thalysias) Hymeraphia C. (Thalysias) anchorata Carter Dictyocylindrus C. (Clathria) Microciona C. (Microciona) anchoratum Carter Echinonema synonym of C. (Thalysias) cactiformis (Lamarck) aruensis Hentschel ascendens Cabioch asodes de Laubenfels Eurypon C. (Clathria) anplaris Sara & Sinbelli Microciona C. (Microciona) angulifera Dendy Clathria C. (Clathria) angulosa Duchassaing & Michelotti atoxa Bergquist C. (Clathria) Dictyociona & Fromont atlantica SarazyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO Echinoclathria ^Echinoclathria Panda ros Mycale (Mycalidae) atrasan guinea Bowerbank Microciona C. (Microciona) Wilsonella C. (Wilsonella) Ophlitaspongia E. (Echinochalina) - assimilis Topsent Clathria anomala Burton Rhaphidophlus C. (Thalysias) anomala Hallmann australiensis Carter Echinochalina E. (Echinochalina) anonyma Burton Microciona C. (Microciona) australiensis Ridley C. (Microciona) REVISION OF MICROCIONIDAE ^ 577 CURRENT SPECIESzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ORI GI NAL GENUS brondst edi ASSI GNMENTzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONML Ant ho A.( Ant ho) Bergquist & synonym of Fromont aust ralis Lendenfeld aust ralis Whitelegge axinelloides Clat hria Plum ohalichondria Crella incrust ans var. arenacea Carter Micr ociona C. ( Microciona) synonym of bulbosa Crella incrust ans Hooper & Levi Clat hria C. ( Clat hria) (Carter) (Crellidae) bulbot oxa Clat hria ( Microciona) C. ( Microciona) burt oni nom.nov. [for Clat hria prolifera Burton] C. ( Clat hria) burt oni Levi Plocam illa Ant ho ( Plocam ia) Spongia C. ( Thalysias) Dendy Ophlit aspongia Echinoclat hria axociona Levi Clat hria C. ( Clat hria) barba Lamarck barba E. ( Echinochalina) barbadensis Plocam illa Ant ho ( Plocam ia) Bowerbank I sodict ya C. ( Clat hria) bargibant i Hooper & Levi E. ( Prot ophlit aspongia) E. ( Prow phlit aspongia) Van Soest barleei basiarenacea bulboret ort a (Carter) (Crellidae) Van Soest cact iforrnis Lamarck caelat a Hallmann Clat hria C. ( Clat hria) caespes Ehlers Scopalina unrecognisable caespit osa Carter calla Boury-Esnault Rhaphidophlus C. ( Thalysias) de Laubenfels basifixa Topsent Ophlit aspongia C. ( Microciona) calochela basilana Levi Clat hria C. ( Clat hria) basispinosa Burton beanii Bowerbank Microciona I sodict ya biclat hrat a nom.nov. biham igera Waller bispiculat a Dendy bispinosus Whitelegge Whitelegge calypso Siphonochalina Hym eraphia C. ( Thalysias) Clat hria C. ( Clat hria) C. ( Clat hria) synonym of A. cam pecheae [for Hym eraphia affinis Topsent] C. ( Microciona) Esperiopsis C. ( Axociella) Spongia C. ( Thalysias) ( Ant ho) involvens nom.nov. (Schmidt) canaliculat a Echinoclat hria C. ( Clat hria) Whitelegge] Microciona C. ( Microciona) Clat hria [for Microciona clat hrat a (Anchinoidae) Axociella Boury-Esnault cancellaria Lamarck Phakellia Plurnohalichondria (Topsent) Echinoclat hria sp. nov. Hentschel calopora Whitelegge berg quist ae beringensis synonym of C. ( Microciona) ant arct ica Hentschel Echinonem a Pronax cant abrica Orueta Rhaphidophlus C. ( Clat hria) carbonaria Spongia Haliclona Rhaphidophlus C. ( Thalysias) Microciona C. ( Microciona) Lamarck (Chalinidae) coriocrassus (Anchinoidae) Bergquist & Fromont E. ( Prot ophlit aspongia) carnosa Bowerbank synonym ofzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGF synonym of Rhaphidophlus C. ( Thalysias) lendenfeldi cart eri Topsent Clat hria Ridley & Dendy C. ( Uat hria) foliacea Topsent synonym of cart eri bit oxa BurtonzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA C. ( Microciona) ^Hym ant ho Echinoclat hria Holopsam m a Ridley & Dendy favus (Carter) bit oxifera Koltun Axociella C. ( Thalysias) cercidochela [for Clat hria C. ( Clat hria) borealis nom.nov. C. ( Wilsonella) Vacelet & Clat hriopsam m a robust a Koltun] Vasseur brat t egardi Van Soest & Stone brepha de Laubenfels Ant ho Aaat a A.( Ant ho) C. ( Microciona) synonym of brevispina Lendenfeld Thalassodendron C. ( Thalysias) cact iform is (Lamarck) brondst edi nom.nov. [for Hym edesm ia pennat a Brondsted] C. ( Microciona) cervicornis Thiele Rhaphidophlus C. ( Thalysias) chalinoides Cart er Axinella Echinoclat hria chart acea Clat hria Ant ho ( I sopenect ya) Spanioplon C. ( Clat hria) [preocc.] Micr ociona see C. ( Microciona) t ut ust ae nom.nov. circonflexa Levi Plocam illa A nt ho ( Plocam ia) (Whitelegge) chehfera Hentschel chelifera Levi 578^ SPECIES cladoflagellata Carter MEMOIRS OF THE QUEENSLAND MUSEUM ORIGINAL GENUS Axinella clathrata Schmidt Tenacia claudei nom.nov. synonym of Echinoclathria chalinoides (Carter) synonym of C. (Thalysias) virgultosa (Lamarck) corticata var. elegans Lendenfeld Sigmatella synonym of C. (Wilsonella) australiensis (Carter) clathrata Whitelegge [preocc.] Microciona see C. (Clathria) biclathrata nom.nov. clavtforrnis Hentschel Clathria C. (Wilsonella) synonym of Antho (Plocamia) cliftoni [for Microciona Hymeniacidon frondifera Bowerbank ^C. (Microciona) acanthotoxazyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA (Lamarck) Levi & Levi] coralliophilus Rhaphidophlus C. (Thalysias) C. (Microciona) Clathria Thiele cleistochela Topsent coccinea Berquist Microciona coccinea Duchassaing & Michelotti CURRENT ASSIGNMENT Thalysias collata sp.nov. C. (Microciona) costifera Hallmann Spirastrella (Spirastrellidae) craspedia sp. nov. crassa Lendenfeld Antherochalina C. (Thalysias) E. (Protophlitaspongia) crassa Carter^Holopsamma cratitia Esper^Spongia Holopsamma ctenichela Alander Microciona C. (Microciona) curvichela Hallmann Wilsonella C. (Dendrocia) Microciona see C (Microciona) vacelettia nom.nov. conectens Hallmann Wilsonella C. (Clathria) compressa Schmidt Clathria C. (Clathria) concentrica Lendenfeld Antherochalina Cymbastela (Axinellidae) Clathria C. (Thalysias) C. (Clathria) C. (Thalysias) confragosa Hallmann Ophlitaspongia Echinoclathria curvichela Vacelet & Vasseur [preocc.] conica Levi Clathria C. (Clathria) cullingworthi Burton Clathria C. (Thalysias) contexta Sara contorta Bergquist & Fromont Echinoclathria Echinoclathria curvispiculifera Carter Microciona ? C. (Clathria) virtually unrecognisable cylindrica Ridley & Dendy Esperiopsis C. (Axociella) cylindrica sensu Sim & Byeon Axociella see C (Microciona) simae sp.nov. Rhaphidophlus synonym of C. (Thalysias) kieschnicki Hooper Dictyociona C. (Clathria) copiosa Topsent Clathria synonym of C. (Thalysias) virgultosa (Lamarck) copiosa var. curacaoensis^Clathria Arndt synonym of C. (Thalysias) schoenus (dc Laubenfels) cylindricus Kieschnick [preocc.] coppingeri Ridley Clathria C. (Thalysias) darwinensis sp.nov. coppingeri var. aculeata synonym of C. (Thalysias) lendenfeldi Ridley & Dendy dayi Levi corallitincta Dendy Clathria Clathria coralloides Olivi Spongia synonym of C. (Thalysias) vulpina (Lamarck) C. (Clathria) synonym of Lissodendolyx cotnplicata (Lundbeck) (Myxillidae) C. (Thalysias) Clathria C. (Clathria) decumbens Ridley Clathria C. (Clathria) delaubenfelsi Levi Rhaphidophlus C. (Thalysias) delaubenfelsi Little Holoplocamia Antho (Plocamia) delicata Lambe Clathria synonym of C. (Ctathria) prolifera (Ellis & Solander) dendyi Berquist & Fromont Microciona C. (Microciona) densa Burton Microciona C. (Microciona) corallorhizoides Fristedt Clathria coriacea Bowerbank Isodictya^Antho (Plocamia) corneolia Hooper & Levi Clathria (Thalysias) C. (Thalysias) dentata Topsent Clathria corona Lieberkiihn Halichondria synonym of C. (aathria) coralloides synonym of C. (Thalysias) fascicularis Topsent depressa Sara & Melone Clathria C. (Clathria) REVISION OF MICROCIONIDAE SPECIES ORIGINAL GENUS CURRENT ASSIGNMENT ^ erectus Thiele^Rhaphidophlus C. (Thalysias) eutypa de Laubenfels^Dictyociona C. (77talysias) 579 dianae Schmidt Sube rites C. (Microciona) C. (Microciona) Artemisina synonym of Arterntsina apollinis (Ridley & Dendy) fallax Bowerbank Microciona dianae Topsent fascicularis Topsent Clathria C. (Thalysias) dichotoma Esper Spongia A.(Antho) fasciculata Wilson Clathria C. (Thalysias) dichotoma Levi Ophlitaspongia Echinoclathria fascispiculifera Carter Microciona C. (Microciona) Clathria synonym of C. (Thalysias) lendenfeldi Ridley & Dendy fauroti Topsent Axosube rites diechinata Hallman digitata Lendenfeld favosa Whitelegge Clathria favosa Lamarck Spongia C. (Axociella) synonym of C. (Thalysias) cacttformis (Lamarck) synonym of E. (Echinochalina) barba (Lamarck) Thalyssodendron Echinoclathria digitiformis Row Ophlitaspongia Echinoclathria discreta Thiele Microciona C. (Clathria) distincta Thiele Hymeraphia C. (Thalysias) ditoxa Stephens Eurypon C. (Microciona) favulosa sp.nov. dives Topsent Microciona ? Plumohalichondria (Anchinoidae) favus Carter Echinoclathria Holopsamma favus var. arerufera Carter Echinoclathria synonym of Holopsamma laminaefavosa Carter dubia Kirkpatrick Microciona duplex Sara dura Whitelegge dura var. mollis Hentschel C. (Thalysias) Clathria Clathria C. (Microciona) C. (Dendrocia) Clathria synonym of C. (Clathria) squalorum Wiedenmayer eccentrica Burton Ophlitaspongia C. (Isociella) echinata Alcolado Axociella echinonematissima wilsonella Carter C. (Microciona) egena Wiedenmayer Echinoclathria Echinoclathria elegantula Ridley & Dendy Clathria C. (Dendrocia) C. (Clathria) E. (Protophlitaspongia) E. (Echinochalina) felixi sp.nov. ferrea de Laubenfels Fisherispongia C. (Wilsonella) fictitia Bowerbank Microciona Phorbas (Anchinoidae) filifer Ridley & Dendy Rhaphidophlus C. (Thalysias) filifer var. cantabrica Orueta Rhaphidophlus C. (Clathria) cantabrica (Orueta) C. (Thalysias) filifer var. mutabilts mutabilis Rhaphidophlus (Topsent) Topsent C. (Clathria) elastica LevizyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^Clathria see C. (Thalysias) filifer var. see C. (Clathria) elastica Clathria spinifera spinifera Rhaphidophlus sarai nom.nov. Sara [preocci (Lindgren) Lindgren ? C. (Clathria) elegans synonym of Clathria virtually Vosmaer flabellata C. (Axociella) unrecognisable Ophlitaspongia nichficata Topsent Raspailia elegans (Kirkpatrick) Antherochalina (Raspailiidae) Lendenfeld synonym of elegans C. (Isociella) flabellata Holopsamma Plectispa Phakellia Lendenfeld macropora Riley & Dendy Lendenfeld elegans Antho (Plocamia) Plocamia Ridley & Dendy flabellata Burton Rhaphidophlus^C. (Clathria) flabellifera Hooper & Levi Clathria (Thalysias) C. (Thalysias) flabelliformis Carter Echinonema synonym of C. (Thalysias) cactiformis (Lamarck) foliacea Topsent Clathria C. (Clathria) Clathria ^C. (Clathria) Aulenella elegantula Dendy Artemisina Artemisina elliptichela Alander Microciona C. (Microciona) encrusta Kumar ensiae sp.nov. Clathria C. (Thalysias) erecta FerrerHernandez Plocamia Antho (Plocamia) foliascens Vacelet & Vasseur erecta Topsent Artemisina Artemisina foraminifera Burton & Rao C. (Wilsonella) C. (Wilsonella) 580^ SPECI ES MEMOIRS OF THE QUEENSLAND MUSEUM ORI GI NAL GENUS hartmeyeri Hentschel Clathria synonym of A.(Antho) tuberosa (Hentschel) CURRENT ASSI GNMENT {3-adaa ovv u erbank Microciona C. (Microciona) frogeti Vacelet Microciona C. (Microciona) [for Microciona microchela Hechtel] C. (Thalysias) fromontae nom.nov. [for Axociella toxitenuis Bergquist & Fromont] hechteli nom.nov. C. (Axociella) hentscheli nom.nov. [for Hymeraphia lendenfeldi Hentschel] C. (Microciona) frondiculata Schmidt Reniera C. (Clathria) hesperia sp.nov. - C. (Thalysias) frond ifera Lamarck Spongia Antho (Plocamia) heterospiculata Brondsted Microciona A.(Antho) frondifera Bowerbank synonym of C. (Microciona) C. (Thalysias) vulpina (Lamarck) heterotoxa Hentschel Microciona Halichondria hexagonopora Levi Clathria C. (Clathria) Clathria synonym of C. (Thalysias) vulpina (Lamarck) Thalysias unrecognisable frondifera var. dichela Hentschel Clathria synonym of C. (Thalysias) vulpina (Lamarck) hians Duchassaing & Michelotti hispanica FerrerHernandez Artemisina Artemisina frondifera var. major Hentschel Clathria see C. (Thalysias) major Hentschel hirsuta Hooper & Levi Clathria (Thalysias) C. (Thalysias) fucoides Bowerbank Ophlitaspongia Terpiosella (Suberitidae) hispidula Ridley Amphilectus C. (Clathria) hjorti Arnesen Echinoclathria Echinoclathria C. (Thalysias) horrida Row Ophlitaspongia C. (Clathria) hymedesmioides Clathria Van Soest (Microciona) C. (Microciona) ignis Duchassaing & Michelotti Thalysias Tedania (Tedaniidae) Plocamilla Antho (Plocamia) - frond ifera var. setotubulosa Wilson fusterna sp.nov. gabrieli Dendy Ophlitaspongia E. (Echinochalina) georgiaensis nom.nov. [for Ophlitaspongta thielei Burton] C. (Axociella) HaIme synonym of Holopsamma laminaefavosa Carter gigantea Lendenfeld glabra Ridley & synonym of E. globosa Lendenfeld synonym of Holopsamma crassa Carter illawarrae sp.nov. illgi Bakus C. (Microciona) impetfecta Dendy Clathria (Echinochalina) ^Echinoclathria DendyzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA inanchorata barba (Lamarck) HaIme gorgonoides Dendy Echinodictyum C. (Clathria) gracea Bakus Anthoarcuata A.(Antho) gracilis Ridley Echinonema synonym of C. (Thalysias) procera (Ridley) Echinoclathria synonym of Echinoclathria subhispida Carter gracilis Carter gradalis Topsent Clathria C. (Microciona) grisea Hentschel Leptosia C. (Microciona) r Plocamia Antho (Plocamia) haematodes de Laubenfels Microciona nags a cimm C. (Dendrocia) Ridley & Dendy Clathria C. (Clathria) incrustans Bergquist Isociella C. (Isociella) incrustans Carter Echinonema Crella (Crellidae) incrustans Svarcevskij Raspailia A.(Antho) involvens (Schmidt) ? synonym of indica Dendy^Clathria C. (Clathria) indica Thomas^Qasimella A rtemisina indistincta Bowerbank Hymedesmia synonym of Plocamionida ambigua (Bowerbank) (Anchinoidae) indurata Hallmann Clathria inhacensis Thomas synonym of C. (Thalysias) cactiformis (Lamarck) ^ C. (Clathria) Clathria C. (Microciona) inornata Hallmann Ophlitispongia Echinoclathria C. (Thalysias) intermedia Kirk Clathria C. (Clathria) C. (Microciona) hallezi Topsent^Heteroclathria A.(Antho) hallmanni sp.nov. - C. (Thalysias) haplotoxa Topsent Leptoclathria hartmani Simpson Axocielita • ^ REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR 581 SPECIES intermedia Whitelegge ORIGINAL GENUS CURRENT ASSIGNMENT laevis Bowerbank Microciona C. (Microciona) laevissima Dendy Hymedesmia C. (Microciona) Anaata C. (Microciona) C. (Thalysias) Echinoclathria E. (Echinochalina) lajorei de Laubenfels Paresperia Esperiopsis (Desmacididae lambda Levi Leptoclathria Rhabderemia (Rhabderemiidae) lambei Koltun Microciona C. (Axociella) lambei Burton Heteroclathria Antho (Plocamia) involvens Schmidt Myxilla A.(Antho) Holopsamma C. (Clathria) laminaefavosa Carter Holopsamma irregularis Burton Marleyia irregularis Lendenfeld synonym of Holopsamma laminaefavosa Carter /axa Lendenfeld Halme synonym of Holopsatnma laminaefavosa Carter intermedia Burton intexta Carter Microciona Halme [for Microciona rubens Bergquist] C. (Microciona) [for Microciona lematolae placenta sensu de C. (Thalysias) sp.nov. Laubenfels] leporina Lamarck Spongia Echinoclathria leighensis nom.nov. isochelifera Uriz Echinoclathria E. (Prowphlitaspongia) E. (Echinochalina) isodictyoides Van Soest Rhaphidophlus C. (Thalysias) ixauda Levi Microciona C. (Microciona) jacksoniana Dendy Phakellia synonym of C. (lsociella) macropora Lendenfeld jecusculum Bowerbank C. (Microciona) levis Lendenfeld Echinonema Hymeniacidon synonym of Crella incrustans (Carter) (Crellidae) johnsoni de Laubenfels Comulum Cornu/um (Coelosphaeridae) lendenfeldi Ridley & Dendy Clathria C. (77ia/ysias) jolicoeuri Topsent Rhaphidophlus C. (Thalysias) Hymeraphia jovis Dendy Artemisina Artemisina lendenfeldi Hentschel [preocc.] see C. (Microciona) hentscheli nom.nov. i a .vf Wilson .i . neTnlialry s?ifa s) virgultosa (Lamarck) linda de Laubenfels Axocielita C. (Thalysias) Clathria lindgreni nom.nov. [for Clathria ramosa Lindgren] C. (Wilsonella) juncea Burton Clathria C. (Clathria) juniperina (var. A) Lamarck Spongia C. (Thalysias) lipochela Burton Clathria lissocladus Burton Rhaphidophlus C. (Clathria) C. (Thalysias) juniperina (var. B) Lamarck Spongia synonym of C. (Thalysias) coppingeri Ridley lissosclera Bergquist & Fromont Clathria C. (Clathria) kasumiensis Tanita ? Pandaros uncertain placement lit hophoenix de Laubenfels Plocamia A.(Antho) Raspailia kentii Bowerbank Microciona C. (Microciona) /itos Hooper & Levi Clathria C. (Wilsonella) (Clathriopsamma) kieschnicki Hooper Clathria (Thalysias) C. (Thalysias) isaaci sp.nov. levii Sara & Siribelli Microciona Echinoclathria levii sp. nov. C. (Microciona) lizardensis sp.nov. kilauea de Laubenfels Axocielita C. (Thalysias) koltuni Hooper Clathria (Thalysias) C. (Thalysias) kylista Hooper & Levi Clathria C. (Clathria) laboutei Hooper & Levi E. (Protophlitaspongia) E. (Protophlitaspongia) labyrinthica Schmidt Reniera C. (Clathria) laciniosa Bowerbank & Norman lsodictya synonym of C. (Clathria) barleei (Bowerbank) laevigata Lambe Clathria C. (Clathria) C. (Microciona) Clathria C. (Clathria) lobosa Lendenfeld Clathriopsamma synonym of C. (Wilsonel la) australiensis (Carter) longichela Topsent Clathria synonym of C. (Clathria) anchorata (Carter) longispiculum Carter longistyla Burton Microciona C. (Microciona) Microciona C. (Microciona) longitoxa Hentschel Hymeraphia C. (Thalysias) loveni Fristedt Clathria Mycale (Mycalidae) lobata Vosmaer 582^ MEMOIRS OF THE QUEENSLAND MUSEUM CURRENT ORI GI NAL GENUS SPECIESzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA microxea Vacelet Paratenaciella ASSI GNMENT & Vasseur lugubris Echinodictyum mima de Pandaros Duchassaing & Ophlitaspongia (Raspailiidae) Laubenfels Michelotti C. (Microciona) C. (Microciona) Ophlitaspongia minor Burton minuta Van Soest Rhaphidophlus Echinoclathria minutula Carter Microciona Rhabderemia (Rhabderemiidae) mixta Hentschel macrochela Levi Microciona C. (Microciona) macroisochela Levi Clathria C. (Clathria) macropora Lendenfeld Clathria C. (Isociella) Clathria C. (Wilsonella) macropora Lendenfeld Plectispa Holopsamma mo//is Kirkpatrick Clathria Pronax (Anchinoidae) monticularis Ridley & Dendy Axinella Aulospongus (Raspailiidae) morisca Schmidt Clathria synonym of A. (Antho) involvens (Schmidt) mortensii Brondsted Clathria C. (Clathria) macrotoxa Bergquist & Fromont Arociella C. (Axociella) madrepora Dendy Clathria synonym of C. (Gathria) spongodes Dendy maeandrina Ridley Clathria C. (Clathria) major Hentschel Clathria C. (Thalysias) manaarensis Carter Dictyocylindrus C. (Thalysias) Antho (Plocamia) mosulpia Clathria Sim & Byeon mutabilis Topsent Rhaphidophlus C. (Thalysias) Ceraochalina synonym of Echinoclathria subhispida Carter C. (Clathria) marissuperi Pulitzer-Finali Clathria C. (Clathria) multiformis Whitelegge massalis Carter Thalysias Reniera (Chalinidae) multipes Hallmann Clathria (Plectispa) C. (Clathria) maunaloa de Laubenfels Microciona C. (Thalysias) multipora Whitelegge Clathria mediterranea Babic synonym of C. (Thalysias) rubra (Lendenfeld) Artemisina synonym of A.(Antho)involvens (Schmidt) Axociella C. (Axociella) melana Van Soest & Stentoff Echinochalina E. (Echinochalina) multitoxaformis Bergquist & Fromont C. (Clathria) murphyi sp. nov. melana Van Soest Artemisina Artemisina mutans Sara Ophlitaspongia Echinoclathria membranacea Thiele Ophlitaspongia C. (Thalysias) Halichondria menoui Hooper & Levi mutula Bowerbank Clathria C. (Clathria) synonym of C. (Clathria) barleei (Bowerbank) meyeri Bowerbank Ophlitaspongia C. (Clathria) michaelseni Hentschel Hymeraphia C. (Thalysias) microchela Stephens Eurypon C. (Clathria) Microciona see C. (77talystas) hechteli nom.nov. microcionides Carter Plumohalichondria Plocamionida (Anchinoidae) microjoanna de Laubenfels Microciona C. (Microciona) micronesia de Laubenfels Microciona C. (Microciona) microchela Hechtel [preocc.] micropora Lendenfeld micropunctata Burton & Rao microxa Desqueyroux Halme Tenacia Clathria myxilloides Dendy Clathria C. (Dendrocia) ^Eurypon naikaiensiszyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO C. (Thalysias) Hoshino^ C. (Microciona) namibiensis Uriz Microciona nervosa Levi Axociella C. (Thalysias) nexus Koltun Bipocillopsis C. (Clathria) nidijicata Kirkpatrick Ophlitaspongia C. (Axociella) nidus-vesparum Lendenfeld Ha/me synonym of Holopsatnma laminaefavosa Carter Echinoclathria noarlungae sp.nov. C. (Clathria) synonym of Holopsamma crassa Carter nodosa Carter Echinoclathria nonnani Burton Hymantho C. (Thalysias) now Tanita Ophlitaspongia Echinoclathria C. (Clathria) novaezealandiae Brondsted Microciona C. (Microciona) C. (Microciona) Echinoclathria notialis sp. nov. REVISION OF MICROCIONIDAE ^ 583 CURRENT ^ORIGINAL GENUS SPECIESzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA Antho A.(Antho) paucispina Sara ASSIGNMENT & Siribelli novizelanica Ridley Dirrhopalum Antho (Plocamia) nuda Hentschel Clathria C. (Thalysias) obliqua George & Wilson oculata Burton Esperiopsis C. (Clathria) Clathria C. (Clathria) ongulensis Hoshino opuntioides Lamarck Axociella C. (Thalysias) Alcyonium A. (Antho) orientalis Brondsted Rhaphidophlus C. (Thalysias) originalis de Laubenfels Esperiopsis C. (Thalysias) ornata Dendy oroides Schmidt Bubaris Antho (Plocamia) Clathria Agelas (Agelasidae) osismica Cabioch Microciona C. (Microciona) ostacina Rafinesque Spongia oxeata Bergquist & Fromont oxeata Burton pauper Brondsted Clathria C. (Clathria) pectiniformis Carter Echinonema synonym of C. (Thalysias) cactiformis (Lamarck) pellicula VVhitelegge Clathria C. (Clathria) pelligera Schmidt Clathria Dictyonella (Dictyonellidae) pennata Lambe Desmacella C. (Microciona) pennata Brondsted [preocc.] Hymedesmia see C (Microciona) brondstedi nom.nov. pennata Duchassaing & Michelotti Pandaros Echinodictywn (Raspailiidae) penneyi de Laubenfels Holoplocamia Antho (Plocamia) synonym of C. (Clathria) prolifera (Ellis & Solander) petforata Lendenfeld Antherochalina synonym of Antho (Isopenectya) chartacea (Whitelegge) Ophlitaspongia Echinoclathria perforata in part Lendenfeld Antherochalina C. (Clathria) Protophlitaspongia E. (Protophlitaspongia) Amphilectus Megaciella (Iophonidae) A.(Antho) phorbasiformis sp.nov. pilosus Ridley & Dendy oxeotus Van Soest Rhaphidophlus C. (Thalysias) placenta Lamarck Spongia C. (Thalysias) oxitoxa Levi oxneri Topsent Clathria C. (Thalysias) Hymedesmia C. (Clathria) placenta de Laubenfels [preocc.1 Microciona see C. (Thalysias) lematolae sp.nov. oxyphila Hallmann Wilsonella C. (Clathria) planum Carter Microciona pachyaxia Levi Axociella C. (Thalysias) plena Sollas Plocamia C. (Microciona) Antho (Plocamia) pachystyla Levi Clathria C. (Clathria) plinthina de Laubenfels Microciona C. (Microciona) papilla iBowerbank Ophlitaspongia synonym of C. (M wrociona) sertata (Grant) plumosa Montagu Spongia Pronax (Anchinoidae) papillosa Thiele Clathria C. (Clathria) Artemisina Artemisina papyracea Carter Phakellia synonym of Echinoclathria leporina (Lamarck) plumosa Hentschel pluritoxa Pulitzer-Finali Echinoclathria Holopsamma paradoxa Babic ? synonym of A. (Antho) involvens (Schmidt) Clathria C. (Clathria) Artemisina oxeifera FerrerHernandez Clathria parkeri sp. nov. Echinoclathria plurityla Pulitzer-Finali piniformis Carter plana Carter C. (Thalysias) Dictyocylindrus C. (Clathria) Microciona synonym of C. (Thalysias) virgultosa (Lamarck) parthena de Laubenfels Microciona C. (Microciona) partita Hal!mann Clathria C. (Clathria) poecilosclera Sara & Siribelli Microciona C. (Microciona) parva Levi Clathria C. (Axociella) polita Ridley C. (Axociella) prima Brondsted Hymedesmia Lissoplocamia Antho (Plocamia) C. (Clathria) primitiva Koltun Microciona C. (Microciona) primitiva Burton Clathriella Ant/so (lsopenectya) procera Ridley Rhaphidophlus C. (Thalysias) patula sp.nov. paucispicula Burton Rhapidophlus paucispina Lendenfeld Thalassodendron synonym of C. (Thalysias) rubra (Lendenfeld) C. (Microciona) 584^ SPECIES MEMOIRS OF THE QUEENSLAND MUSEUM ORIGINAL GENUS CURRENT ASSIGNMENT synonym of C. (Mtcrociona) calla (de Laubenfels) rarispinosa Hechtel Microciona reinwardti Vosmaer rein wardti var. palnuaa Ridley Clathria C. (Thalysias) Clathria synonym of C. (Thalysias) vulpina (Lamarck) reinwardti var. subcylindrica Ridley Clathria synonym of C. (Thalysias) reinwar&i Vosmaer renieroides Lendenfeld Antherochalina synonym of Phalcelliaflabellata (Carter) (Axinellidx) repens Duchassaing & Michelotti Thalysias synonym of Xestospongia subtriangularis (Duchassaing) (Petrosiidae) reticulata Lendenfeld Clathriopsamma C. (Wilsonella) reticulata Bergquist & Fromont Ophlitaspongia Echinoclathria reticulata Whitelegge Echinochalina E. (Echinochalina) synonym of Ascena procumbens ^Clathria procumbenszyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA rectangulosa Lendenfeld sensu Brondsted C. (Clathria) Clathria Schmidt (Calcarea) productitoxa Hoshino Thalysias C. (Clathria) prolifera Ellis & So'ander Spongia C. (Clathria) prolifera Burton jpreocc.] Clathria see C. (Clathria) burtoni nom.nov. f,r° unVa eck Hymedesmia C. (Microciona) proxima Duchassaing & Michelotti Thalysias Xestospongia (Petrosiidae) pseudonapya de Laubenfels Clathriopsamma C. (Wilsonella) pugio Lundbeck Hymedesmia ^C. (Microciona) punicea sp.nov. Antho (Isopenectya) synonym of Rhabderemia minuttda (Carter) (Rhabderemiidae) pusilla Carter Microciona pustulosa Carter Halichondria^C. (Microciona) pyramida Lendenfeld pyramidalis Brondsted Clathria C. (Dendrocia) rhaphidotoxa Stephens Clathria C. (Clathria) Microciona C. (Clathria) rhopalophora Hentschel Hymeraphia C. (Microciona) Microciona C. (Microciona) riddlei sp. nov. Rhaphidophlus C. (Thalysias) ridleyi Hentschel Plocamia Antho (Plocamia) E. (Echinochalina) quadriradiata Carter Echinoclathria quercifolia Keller Antherochalina Phakellia (Axinellidae) ridleyi Lindgren quinqueradiata Carter Microciona Cyamon (Raspailiidae) ridleyi Dendy Echinodiciyum synonym of C. (Axociella) nidificata (Kirkpatrick) robusta Dendy Microciona C. (Thalysias) robusta Koltun [preocc.] Clathria see C. (Clathria) borealis nom.nov. rameus Koltun Axociella ramosus Kieschnick ramosa Lindgren [preocc.] Rhaphidophlus ^C. (Thalysias) Clathria see C. (Wilsonella) lindgreni nom.nov. ramosa Dendy fpreocc.] Colloclathria see C. (Thalysias) amiranteiensis nom.nov. ramosa Hallmann Echinoclathria^Holopsamma ramsayii Lendenfeld Tho recta synonym of C. (Wilsonella) australiensis (Carter) rotunda Hallmann Echinoclathria Holopsamma rubens Lendenfeld rubens Bergquist [preocc.] C. (Clathria) rubens var. dura Lendenfeld Thalassodendron rubens var. lamella Lendenfeld Thalassodendron synonym of C. (Thalysias) rubra (Lendenfeld) C. (Thalysias) Thalassodendron Microciona see C. (Microciona) leighensis nom.nov. synonym of C. (Thalysias) rubra (Lendenfeld) rubra Lendenfeld Echinonema Clathria (Clathriopsamma) C. (Wilsonella) Thalysias raphanus Lamarck Spongia C. (Clathria) rugosa Hooper & Levi raphida sensu Hechtel Clathria synonym of Cliona rhaphida Boury-Esnault (Clionidae) rugosa Duchassaing & Michelotti raraechelae Van Soest Rhaphidophlus synonym of C. (Thalysias) venosa (Alcolado) saintviticenti sp.nov. synonym of Xestospongta subtriangularis (Duchassaing) (Petrosiidae) A ntho (Isopenectya) REVISION OF MICROCIONIDAE SPECIES ORIGINAL GENUS CURRENT ASSIGNMENT ^ 585 spinifera Sara [preocc.] Clathria see C. (Clathria) saraspinifera nom.nov. C. (Clathria) sarai nom.nov. [for Clathria elastica Sara] C. (Clathria) [for Clathria spinifera Sara] spinispicula Tanita Clathria saraspinifera nom.nov. C. (Clathria) spinosa Wilson Microciona C. (Microciona) Microciona lophon (Iophonidae) spinulenta sartaginula Lamarck Spongia C. (Clathria) saxicava Duchassaing & Michelotti Thalysias unrecognisable scabida Carter Halichondria C. (Dendrocia) schoenus de Laubenfels Clathria C. (Thalysias) spongiosa Dendy Echinodictyum E. (Echinochalina) scotti Dendy Clathria C. (Microciona) spongodes Dendy Clathria C. (Clathria) C. (Isociella) squalorum Wiedenmayer C. (Clathria) selachia sp.nov. seriata Grant Spongia C. (Microciona) seriatus Thiele Rhaphidophlus synonym of C. (Thalysias) vulpina (Lamarck) sessilis Carter Dictyocylindrus unrecognisable shirahama Tanita Clathria C. (Clathria) sigmoidea Cuartas Microciona C. (Microciona) signata Topsent Plocamiopsis Antho (Plocamia) simae sp.nov. [for Axaciella cylindrica sensu Sim & Byeonl C. (Microciona) simi/is Thiele Hymeraphia C. (Microciona) simi/is Stephens [preocc.] Microciona simi/is sensu Uriz Eurypon see C (Microciona) stephensae nom.nov. see C (Microciona) urizae nom.nov. simplex Lendenfeld Halme Holopsamma simplicissima Norman Microciona Bubaris (Axinellidae) simpsoni Van Soest C. (Microciona) synonym of C. (Microciona) echinata (Alcolado) skia sp.nov. C. (Isociella) ssallicamtaann Clathria synonym of C. (Thalysias) lendenfeldi Ridley & Dendy spiculosus Dendy Rhaphidophlus Bowerbank spongigartina de Laubenfels spongiosa Burton Aaata C. (Microciona) Clathria synonym of C. (Clathria) spongodes Dendy Clathria stipitata KoltunzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR ^Artemisina Artemisina striata Whitelegge Clathria C. (Clathria) strongyla Hentschel Artemisina synonym of Artemisina plumosa Hentschel stephensae nom.nov. [for Microciona similis Stephens] C. (Microciona) strepsitoxa Hope strepsitoxa var. robusta Dendy Microciona C. (Microciona) Microciona C. (Thalysias) robusta (Dendy) styloprothesis sp.nov. C. (Thalysias) suberitoides Vosmaer Artemisina subhispida Carter synonym of Artetnisina arciger (Schmidt) Echinoclathria Echinoclathria subtriangularis Duchassaing Thalysias Xestospongia (Petrosiidae) surculosa Esper Spongia virtually unrecognisable ? C. (Clathria) svarchevskyi de^mtcrociona — •^• Laubenfels synonym of C. (Mwrociona) armata (Bowerbank) tenebratus Whitelegge Rhaphidophlus synonym of C. (Clathria) striata Whitelegge tener Carter Thalysias ? C. (ThaNsias) imperfectly known tenuifibra Whitelegge Clathria synonym of C. (Clathria) rubens (Lendenfeld) Microciona C. (Microciona) Hentschel C. (Thalysias) synonym of C. (Thalvsias) reinwardti Vosmaer spiculosus var. ramosa Hentschel Clathria synonym of C. (Thalysias) procera (Ridley) tenuis Stephens spinarcus Carter & Hope Microciona C. (Microciona) tenuis Carter Echinoclathria synonym of Echinoclathria leporina (Lamarck) spinatoxa Hoshino Microciona C. (Microciona) tenuispina Lendenfeld Antherochalina synonym of Echinoclathria leporina (Lamarck) spiculosa var. macilenta spinifera Lindgren Clathria Rhaphidophlus C. (Thalysias) 586^ SPECIES t enuissim a Stephens MEMOIRS OF THE QUEENSLAND MUSEUM ORIGINAL GENUS Eur w on t erranovae Dendy Clat hria t et rast yla CURRENT ASSIGNMENTzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHG Art em isina Art em isina t ubulosa Koltun C. ( Microciona) t unisiae C. ( Clat hria) ty/ota Boury-Esnault Hym eraphia C. ( Microciona) t ext ile Carter Cornulum C. ( Clat hria) t het idis Hallmann Ophlit aspongia C. ( Axociella) t hielei Hentschel Hym eraphia C. ( Microciona) Hentschel t hielei Burton [preocc.] see C. .( Axociella) Ophlit aspongia georgraenst s nom.nov. t ingens sp.nov. C. ( Thalysias) t opsent i Thiele Rhaphidophlus C. ( Thalysias) t ort uosa Uriz Clat hria C. ( Clat hria) nom.nov. [for Microciona chelifera Levi] C. ( Microciona) Art em isina Cornu/um (Iophonidae) t ypica Kirkpatrick Clat hria C. ( Clat hria) i'ypicum synonym of C. ( Thalysias) cact iform is Carter Echinonem a (Lamarck) synonym of t ypica var. porrect a Hent schel Clat hria u/mus Vosmaer Clat hria unica Cuartas Clat hria C. ( Thalysias) reinwardt i Vosmaer C. ( Clat hria) C. ( Clat hria) synonym of C. ( Ciat hria) prolifera (Ellis & t oxifera Hentschel Hym eraphia C. ( Thalysias) urceolat a see C. ( Microciona) ant arct ica (Topsent) Desor Spongia t oxiferum C. ( Microciona) urizae nom.nov. [ for Eurypon C. ( Microciona) sim ilis sensu Uriz] vacelet t ia [for Microciona curvichela Topsent [preocc.] St ylost ichon t oxim aj or Topsent Clat hria t oxipraedit a Topsent t oxirect a Sara & Siribelli t oxist rict a Topsent t oxist yla Sara Clat hria C. ( Clat hria) Micr ociona C. ( Microciona) Clat hria C. ( Clat hria) Micr ociona C. ( Clat hria) t oxit enuis Topsent Clat hria nom.nov. Duchassaing & Michelotti vasiforrnis de Carter synonym of Echinodict yum m esent erinum venosa Alcolado Microciona vasiplicat a t oxivaria Sara Microciona C. ( Clat hria) t oxot es Schmidt Scopalina C. ( Clat hria) Hallmann Clat hria C. ( Clat hria) t ransiens Topsent Art em isina Art em isina Ophlit aspongia Echinoclat hria t ricurvat ifera Carter t uberculat a Burton t ube rosa (Spirastrellidae) Echinonem a see C. ( Axociella) from ont ae nom.nov. Pulitzer-Finali Ant hosigm ella C. ( Clat hria) Bergquist & Axociella Fromont [preocc.] t ranslat a Thalysias C. ( Microciona) Thalyseurypon Laubenfels t ransiens Vacelet & Vasseur] varians C. ( Microciona) t oxit enuis Solander) (Lamarck) (Raspailiidae) C. ( Thalysias) synonym of vent ilabrum var . ^Phakellia aust raliensis C. ( Thalysias) cact 'form is vespariut n Spheciospongia (Lamarck) ? C. ( Thalysias) Thalysias imperfectly known Lamarck vit ninalis Lendenfeld Alcyonium synonym of Thalassodendron synonym of St ylost ichon C. ( Mt crociona) ant arct ica (Topsent) virgula Sara & Siribelli (Spirastrellidae) Echinoclat hria subhispida Cart er synonym of A. Microciona ( Ant ho) involvens (Schmidt) Bowerbank Microciona t ube rosa Lissodendoryx zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^A.( Ant ho) vulcani Levi Art em isina A rt em isina Hentschel E. ( Prot ophlit aspongia) t uberosa sp.nov. t uberosocapit at a Topsent t ubulat um Bowerbank t ubulosa Hallmann t um ulosa Bowerbank C. ( Wilsonella) Hym eraphia Haliphysem a Discorhabdella vt rgult osa Lamarck Spongia vulpina Lamarck Spongia C. ( Thalysias) Echinoclat hria Echinoclat hria waldoschm it t i de Laubenfels (Hymedesmiidae) ? Aulospongus (Raspailiidae) synonym of walpersii Duchassaing & Michelotti Ophlit aspongia ^E. ( Echinochalina) wesselensis Microciona ^C. ( Microciona) C. ( Thalysias) sp.nov. Pandaros Pt ilocaulis spiculifera (Lamarck) (Axinellidae) C. ( Thalysias) REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 587 incorporating the Indonesian and Micronesian faunas (in prep.). My phylogeny, based primarily on structural features of the skeleton, conflicts with that of Hajdu et al., 1994. Their hypothesis is based on the premise that microgeometry is less likely to be influenced by modification during the course of evolution than structural features. I emphasise skeletal structure and skeletal differentiation as primary characters in the evolution of Microcionidae. My classification allows inclusion of arcuate and anchorate modified chelae (which would otherwise be included in Myxillina). Identical structural features, primarily, and megasclere geometry, secondarily, in species with arcuate or anchorate chelae may indicate evolutionary similarity, whereas it is debatable whether or not observed modifications to chelae are homologous or merely convergent. Biogeography. Many early attempts to analyse biogeographic patterns of marine sponges were unsuccessful because authors attempted too broad a taxonomic coverage without detailed taxonomic revisions and the belated recognition that many socalled 'widely distributed' species actually consist of allopatric, cryptic sibling species, thus masking potentially informative patterns 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 conprovincial species; grey hatch = % of species endemic to each province. temporary biogeographic MEMOIRS OF THE QUEENSLAND MUSEUM 588zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ species endemism throughout world marine provinces, comparing regional faunas within the Indo-west Pacific region (targeting the New Caledonian fauna in particular), and contrasting these broad distribution patterns for Microcionidae with those of the families Raspailiidae and Axinellidae. These analyses are taken further in this study, focussing in particular on the Australian fauna, and incorporating additional data derived from the present revision. Hooper & Levi (1994) also provided an areacladistic analysis of one species group (Clathria `procera' group), comparing sibling species' distributions and postulating historical biogeographic patterns and relationships throughout the world's seas. Although it is clear from this study and others that historical biogeography can provide many more facets to the questions posed by biogeographers, who attempt to understand relationships between both FIG. 317. A,B, Biogeographic relationships between Australian microcionid Provincial species and the species within Indo-west Pacific marine provinces, showing numbers of shared Provinces themselves, it is species between provinces (provincial endemic species circled), also clear that analysis of many species groups are reanalyses have had greater success based on rela- quired to gain a meaningful tively well-revised, restricted taxa (genera, interpretation of data and to resolve a general area families), (e.g., Van Soest et al., 1991; Hooper, statement for the marine biome (Van Soest et al., 1991; Van Soest & Hooper, 1993; Hooper & Levi, 1991). It is inappropriate to undertake such a 1994; Bergquist & Kelly-Borges, 1995). Yet detailed areacladistic analyses in this present these studies too have not reached any consensus work restricted to the Australian fauna, whereas concerning general area statements for shallow it will be much more useful to include a revised water marine sponges, and they fail to distinguish Indonesian and Micronesian microcionid fauna between vicariant events or subsequent dispersals into analyses (in prep.). The present analysis folto explain species' distributions (Hooper & Levi, lows the format used for Raspailiidae (Hooper, 1994). 1991). From present data and the earlier analysis Only one (Hooper & Levi, 1994) included species of Microcionidae, providing a prelimi- (Hooper & Levi, 1994) there is little evidence for nary analysis of biogeographic patterns amongst cosmopolitan microcionids. A possible exception Indo-west Pacific species, comparing levels of is Clathria atrasanguinea which has contiguous REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 589 populations (i.e. substantiated by comparison of voucher specimens) within the W and E Indian Ocean system, Mediterranean-E Atlantic system, and W Atlantic 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 distributions (possibly explained by local extinctions within intermediate provinces; e.g., C. cactiformis). It 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 between populations based solely on skeletal characters. Similarly, most species of Microcionidae are restricted to single ocean systems, and only about 5% of species range widely within any particular system (whereas most other species are much more restricted in their ranges). Worldwide 72% of species are restricted to a single biogeographic province although levels of regional endemism range from 3080% of species (Fig. 316). Microcionids are relatively diverse and prevalent, with most species found predominantly in shallow-waters but some ranging down to 2500m depth. They comprise 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 ^ MEMOIRS OF THE QUEENSLAND MUSEUM 590zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA diversity indicated: Indo-Australia region and NE Atlantic (Hooper & Levi 1994: fig.3). Within the Indo-west Pacific, extending from the Andaman Sea to islands of the W Pacific rim, there are 196 species of which 115 (or about 60%) are endemic to the region. This level of endemism is 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 contain 148 species of microcionids, comprising about 11% of 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 microcionid distribution data do support the concept of a differentiated southern Gondwanan fauna and northern Tethys fauna within coastal Australia (Hooper & Levi, 1994) based on a different data set. Temperate Australian marine provinces contain a higher diversity of microcionids (90 species) and greater endemism (81%) 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 raspailiid fauna was less diverse but had greater endemism than did the tropical fauna. This observation is further supported considering the Australian fauna in more detail (Fig. 317). Peronian (Fig. 317A). Of all Australian continental provinces the SE Australian temperate (Peronian) province contains the greatest diversity of species (60) and also the highest proportion of regional endemism (28 species or 48%). Not surprisingly greatest similarities in species composition (i.e., numbers of shared species) are with the adjacent temperate provinces (Flindersian, 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 contains 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 (Dampierian) province (no doubt reflecting the relative importance of the Leeuwin current to the marine biogeography of the west coast (Pearce & Walker, 1991; Hooper, 1994)). Maugean (Fig. 317C). The cool 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. Solanderian (Fig. 317D). The NE Australian tropical (Solanderian) province contains 45 species (18 or 40% endemic), with similarities to the adjacent temperate Peronian, tropical Dampierian 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 40% endemic). From these comparisons between adjacent provincial faunas it is clear that the two tropical provinces (Dampierian and Solanderian) share the highest number of sympatric species and also share a large number of species with both Indonesia 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, temperate) 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 and tropical Australian microcionid species. Only two species of microcionids have confirmed trans-Tasman Sea distributions (Bergquist & Fromont, 1988), and only four are found in both the Solanderian and New Caledonian provinces (Hooper & Levi, 1993a), with similar observations reported for Raspailiidae (Hooper, 1991) in which both New Zealand and New Caledonian faunas have exceptionally high numbers of endemic microcionid species (nearly 70% and 67% endemism, respectively). The Antarctic and austral islands faunas have a low diversity of microcionids (18) and few endemic species (28%), 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 observed for the Raspailiidae (Hooper, 1991). REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 591 LITERATURE CITED ACKERS, R.G., MOSS, D. & PICTON, B.E. 1992. Sponges of the British Isles. A colour guide and working document. 5th Edn. (Marine Conservation Society: Ross on Wye, Herefordshire). AHO, S., TURAKAINEN, H., ONNELA, M.L. & BOEDTKER, H. 1993. Characterization of an intonless collagen gene family in the marine sponge Microciona prolifera. Proceedings of the National Academy of Sciences of the United States of America 90(15): 7288-7292. AKIYAMA, S.K. &JOHNSON, M.D. 1983. Fibronectin in evolution: presence in invertebrates and isolation from Microcionaprolifera. Comparative Biochemistry and Physiology (B) 76(4): 687-694. ALANDER, H. 1942. Sponges from the Swedish West coast and adjacent waters. (H. Struves: Goteborg). ALCOLADO, P.M. 1976. Lista de nuevos registros de Poriferos para Cuba. Academia de Ciencias de Cuba, Instituto de Oceanologia (36): 1-11. 1980. Esponjas de Cuba: Nuevos registros. Poeyana (197): 1-10. 1984. Nuevas especies de esponjas encontrades en Cuba. Poeyana (271): 1-22. ALLEE, W.C. 1923. Studies in marine ecology. 1. The distribution of common litoral invertebrates of the Woods Hole region. Biological Bulletin of Woods Hole 44: 167-191. ANDERSON, E.S. 1973. The association of the nudibranch Rostanga pukhra MacFarland 1905 with the sponges Ophlitaspongia pennata, Esperiopsis originalis and Plocamia katykina. Dissertation Abstracts International (B) 33(12): 5668. ANONYMOUS 1985. International Code of Zoological Nomenclature. Third edition. (International Trust for Zoological Nomenclature, H. Charlesworth and Co.: Huddersfield). ARNDT, W. 1913. Zoologische ergebnisse der ersten Lehr-Expedition der Dr. P. Schottlanderschen Jubilaums-Stiftung. Jahresbericht Schles. Gesellschaft Vaterl. Cultur 1912: 110-136. 1927. Kalk- und Kieselshwamme von Curacao. Bijdragen tot der Dierkunde, Amsterdam 25: 133-158. 1935. Porifera. Pp. 1-140. In Grimpe Die Tierwelt der Nord u Ostsee. Volume 3a(27) (Leipzig). ARNESEN, E. 1903. Spongien von der norvegischen Ktiste. II. Monaxonida: Halichondrina. Bergens Museums Aarbog (1): 1-30. 1920. Brutknospenbildung bei Polymastia mammilaris (0.F. Milli.) Bow. (Rinaldia arctica Merej.). Norske Selsk. Skr. 1920(1): 1-24. AYANOGLU, E., RIZZOLIO, M., BEAULIEU, S., OZ, O. & DJERASSI, C. 1990. Covalently bound fatty acids in membrane proteins of some sponges. Comparative Biochemistry and Physiology (B, Comparative Biochemistry) 96(3): 597-603. AYLING, A.L., STONE, S. & SMITH, B.J. 1982. Catalogue of types of sponge species from Southern Australia described by Arthur Dendy. Reports of the National Museum of Victoria 1: 87-109. BABIC, K. 1921. Monactinellida und Tetractinellida der Adria. Glasnik Societas Scientiarum Naturalium Croatica 33: 77-93. 1922. Monactinellida und Tetractinellida des Adriatischen Meeres. Zoologische Jahrbucher Jena Abteilung fur Systematik, Okologie und Geographie de Tiere 46: 217-302. BAGBY, R.M. 1966. The fine structure of myocytes in the sponges Microciona prohlera (Ellis and Solander) and Tedania ignis (Duchaissaing and Michelotti). Journal of Morphology 118: 167-181 . 1970. The fine structure of pinacocytes in the marine sponge Microciona prohfera (Ellis and Solander). Z. Zellforsch 105: 579-594. 1972. Formation and differentiation of the upper pinacoderm in reaggregation masses of the sponge Microciona prolifera (Ellis and Solander). Journal of Experimental Zoology 180(2): 217-244. BAKUS, G.J. 1966. Marine poeciloscleridan sponges of the San Juan Archipelago, Washington. Journal of Zoology, London 149: 415-531. BAKUS, G.J. & GREEN, K.D. 1987. The distribution of marine sponges collected from the 1976-1978 Bureau of Land Management Southern California Bight Program. Bulletin of the Southern California Academy of Sciences 86(2): 57-88. BARNARD, F. 1879. Notes on sponge from Northern Territory. Quarterly Journal of the Microscopical Society of Victoria 1: 14-15. BERGMANN, W., SCHEDL, H.P. & LOW, E.M. 1945. Contributions to the study of marine products. XVII. Spongosterol. XVIII. Microcionasterol and other sterols of sponges. Journal of Organic Chemistry 10(6): 570-579, 580-586. BERGQUIST, P.R. 1961a. A collection of Porifera from Northern New Zealand with Descriptions of seventeen new species. Pacific Science 25(1): 33-48. 1961b. Demospongiae (Porifera) of the Chatham Islands and Chatham Rise, collected by the Chatham Islands 1954 Expedition. New Zealand Department of Scientific and Industrial Research Bulletin. Biological Results of the Chatham Islands 1954 Expedition 5(139): 169-206. 1965. The Sponges of Micronesia, Part 1. The Palau Archipelago. Pacific Science 19(2): 123-204. 1967. Additions to the Sponge fauna of the Hawaiian Islands. Micronesica 3: 159-173. 1977. Porifera. Bernice P. Bishop Museum Special Publication 64(1): 53-69.. 1978. Sponges. (Hutchinson: London). 1980a. The ordinal and subclass classification of the Demospongiae (Porifera); appraisal of the present arrangement, and proposal of a new order. New Zealand Journal of Zoology 7: 1-6. 19806. A revision of the supraspecific classification of the orders Dictyoceratida, Dendroceratida and MEMOIRS OF THE QUEENSLAND MUSEUM 592zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ Verongida (class Demospongiae). New Zealand Journal of Zoology 7:443-503. BERGQUIST, P.R. & FROMONT, J. 1988. The marine fauna of New Zealand: Porifera, Demospongiae, Part 4 (Poecilosclerida). New Zealand Oceanographic Institute Memoir 96: 1-197. BERGQUIST, P.R. & GREEN, C.R. 1977a. A method for preserving larva substrate relationships during preparation for electron microscopy. Biologic Cellulaire 28(1): 85-86. 1977b. An ultrastructural study of settlement and metamorphosis in sponge larvae. Cahiers de Biologie Marine 18(3): 289-302. BERGQUIST, P.R. & HARTMAN, W.D. 1969. Free amino acid patterns and the classification of the Demospongiae. Marine Biology 3(3): 247-268. BERGQUIST, P.R. & HOGG, J.J. 1969. Free amino acid pattern in Demospongiae: a biochemical approach to sponge classification. Cahiers de Biologic Marine 10: 205-220. BERGQUIST, P.R. & KELLY-BORGES, M. 1995. Systematics and biogeography of the genus lanthella (Demospongiae: Verongida: Ianthellidae) in the south-west Pacific. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 12: 151-176. BERGQUIST, P.R. & SINCLAIR, M.E. 1968. The morphology and behaviour of larvae of some intertidal sponges. New Zealand Journal of Marine and Freshwater Research 2: 426-437. 1973. Seasonal variation in settlement and spiculation of sponge larvae. Marine Biology 20: 35-44. BERGQUIST, P.R. & TIZARD, C.A. 1967. Australian intertidal sponges from the Darwin area. Micronesica 3: 175-202. BERGQUIST, P.R., MORTON, J.E. & TIZARD, C.A. 1971. Some Demospongiae from the Solomon Islands with descriptive notes on the major sponge habitats. Micronesica 7(1-2): 99-121. BERGQUIST, P.R., SINCLAIR, M.E. & HOGG, J.J. 1970. Adaptation to intertidal existence: Reproductive cycles and larval behaviour in Demospongiae. Zoological Society of London Symposia 25: 247-271. BERTOLONI, A. 1819. Amoenitales italicae sistentes opuscula ad rem herbarian et zoologiam Italiae spectantia. Bononiae Typis Annesii de Nobilibus. BIBLIONI, M.A. 1993. Some new or poorly known sponges of the Balearic Islands (western Mediterranean). Scientia Marina 57(4): 307-318. BIBLIONI, M.A. & GILI, J.M. 1982. Primera aportacion al conocimiento de las cuevas submarinas de la isla de Mallorca. Oecologia Aquatica (6): 227-234. BIERNBAUM, C.K. 1981. Seasonal changes in the amphipod fauna of Microciona prolifera (Ellis and Solander) (Porifera: Demospongiae) and associated sponges in a shallow salt-marsh creek. Estuaries 4(2): 85-96. BITO, L.Z. 1972. Comparative study of concentrative prostaglandin accumulation by various tissues of mammals and marine vertebrates and invertebrates. Comparative Biochemistry and Physiology (A) 43(1): 65-82. BLAINVILLE, M.H.D. DE. 1819. Eponge. Dictionaire Sciences Naturelles 15: 93-133. BOROJEVIC, R., CABIOCH, L. & LEVI, C. 1968. Inventaire de la faune marine de Roscoff. Spongiaires. Pp. 1-44 (Editions de la Station Biologic Roscoff: Roscoff). BOSC, L.A.G. 1802. Histoire naturelle de vers. Volume 3 (Paris). BOSE, A.K. 1974. Accelerating the quest for bioactive compounds from the sea. Food-Drugs of the Sea Conference Proceedings 1974: 476-490. BOSE, A.K., KRYSCHUCK, J. & NIGRELLI, R.F. 1972. Chemical examination of the red beard sponge (Microciona prolifera). Food-Drugs Sea Conference Proceedings 1972/1973: 217-222. BOURY-ESNAULT, N. 1971. Spongiaires de la zone rocheuse de Banyuls-sur-mer. II. - Systematique. Vie Milieu (B) 22(2): 287-350. 1973. Campagne de la 'Calypso' au large des cOtes Atlantiques de l'Amerique du sud (1961-1962). 1. 29. Spongiaires. Annales de l'Institut Oceanographique, Paris 49 Supplement: 263295. BOURY-ESNAULT, N. & LOPES, M.T. 1985. Les Demosponges littorales de l'Archipel des Acores. Annales de l'Institut Oceanographique, Paris 61(2): 149-225. BOURY-ESNAULT, N. & BEVEREN, M. VAN. 1982. Les Démosponges du plateau continental de Kerguelen-Heard. Territoire des Terres Australes et Antarctiques Francaises Comite National Francaise des Recherches Antarctiques (52): 1132. BOURY-ESNAULT, N., PANSINI, M. & URIZ, M.J. 1992. A new Discorhabdella (Porifera, Demospongiae), a new Tethyan relict of pre-Messinian biota ? Journal of Natural History 26: 1-7. BOWERBANK, J.S. 1861. List of the British marine invertebrate fauna. Sponges. Report of the 30th Meeting of the British Association (Oxford) 1861: 217-236. 1862a. On the anatomy and physiology of the Spongiadae. Part II. Philosophical Transactions of the Royal Society, London 152: 747-829. 1862b. On the anatomy and physiology of the Spongiadae. Part III: On the generic characters, the specific characters and the method of examination. Philosophical Transactions of the Royal Society, London 152: 1087-1135. 1864. A monograph of the British Spongiadae. Volume 1. (Ray Society: London). 1866. A monograph of the British Spongiadae. Volume 2 (Ray Society: London). 1872a. Contributions to a general history of the Spongiadae. Part 1. Proceedings of the Zoological Society of London 1872: 115-129. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 593 1872b. Contributions to a general history of the Spongiadae. Part 2. Proceedings of the Zoological Society of London 1872: 196-202. 1872c. Contributions to a general history of the Spongiadae. Part 3. Proceedings of the Zoological Society of London 1872: 626-635. 1873a. Contributions to a general history of the Spongiadae. Part 4. Proceedings of the Zoological Society of London 1873: 3-25. 1873b. Contributions to a general history of the Spongiadae. Part 5. Proceedings of the Zoological Society of London 1873: 319-333. 1873c. Report on a collection of Sponges found at Ceylon by E.W.H. Holdworth Esq. Proceedings of the Zoological Society of London 1873: 25-31. 1874. A monograph of the British Spongiadae. Volume 3 (Ray Society: London). 1875. Contributions to a general history of the Spongiadae. Part 7. Proceedings of the Zoological Society of London 1875: 281-296. 1876. Contributions to a general history of the Spongiadae. Part 8. Proceedings of the Zoological Society of London 1876: 768-775. 1877. Description of five new species of sponges discovered by A.B. Meyer on the Philippine Islands and New Guinea. Proceedings of the Zoological Society of London 1877: 456-464. 1882. A monograph of the British Spongiadae. Volume 4 (Ray Society: London). BOWERBANK, J.S. & NORMAN, A.M. 1869. A monograph of the siliceo-fibrous sponges. Parts I, II. Proceedings of the Zoological Society of London 1869: 66-100, 323-351. BRIGGS, J.C. 1987. Biogeography and plate tectonics. Developments in palaentology and stratigraphy. (Amsterdam: Elsevier). BRONDSTED, H.V. 1923. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. 15. Sponges from the Auckland and Campbell Islands. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 75: 117-167. 1924. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. 23. Sponges from New Zealand. Part I. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 77: 435-483. 1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. 35. Sponges from New Zealand. Part 2. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 81: 295-331. 1927. Antarctic and subantarctic sponges collected by S. Wallin 1924. Arkiv fOrZoologi A19(6): 1-6. 1929. Neue Schwamme aus Amoy an der Formosa - Strasse. Zoologischer Anzeiger, Leipzig 81: 224-227. 1932. Marine spongia. Pp. 1-32. In 'Zoology of the Faroes'. (Copenhagen). 1934. Resultats scientifiques du voyage aux Indies Orientales Neerlandaises. Sponges. Brusseles Institut royal des sciences naturelles de Belgique. Memoires 2(15): 3-26. BURGER, M.M. 1977. Mechanism of cell-cell recognition: some comparisions between lower organisms and vertebrates. Pp. 357-376. In Karkinen-Jaaskelãinen, M., Saxen, L. & Weiss, L. (eds) 'Cell interactions in differentiation' (Academic Press: London). BURKART, W., JUMBLATT, J., SIMPSON, T.L. & BURGER, M.M. 1979. Macromolecules which mediate cell-cell recognition in zyxwvutsrqponmlk Microciona prolifera. Colloques international du Centre national de la Recherche Scientifique (291): 239246. BURTON, M. 1928. Report on some deep-sea sponges from the Indian Museum collected by the R.I.M.S. 'Investigator'. Part II. Tetraxonida (concluded) and Euceratosa. Records of the Indian Museum, Calcutta 30(1): 109-138. 1929a. Porifera. Part II - Antarctic sponges. British Antarctic (Terra Nova) Expedition. 1910-1913. Natural History Report, Zoology, 6(4): 393-458. 1929b. Mission Saharienne Augieras-Draper, 19271928. Porifera. Bulletin du Museum National d'Histoire Naturelle (2) 1: 157-158. 1930a. Norwegian sponges from the Norman Collection. Proceedings of the Zoological Society of London 2: 487-546. 1930b. Report on a collection of sponges from South Georgia and from Campbell Island, South Pacific, obtained by Dr. Kohl-Larsen. Senckenbergiana 12(6): 331-335. 1930c. Additions to the sponge fauna of the Gulf of Mannar. Annals and Magazine of Natural History (10) 5: 665-676. 1931a. On a collection of marine sponges mostly from the Natal coast. Annals of the Natal Museum 6(3): 337-358. 193 lb. Studies on Norwegian sponges. I. Det Kongelige Norske Videnskabers Selskab 4(39): 136143. 1932a. Sponges. Discovery Reports 6: 237-392 (Cambridge University Press: Cambridge). 1932b. Report on a collection of sponges made in South Saghalin by Mr. Tomoe Urita. Science Reports of the TOhoku Imperial University (4 (Biology)) 7(2): 195-206. 1933. Four new marine sponges from Natal. Annals of the Natal Museum 7(2): 249-254. 1934a. Sponges. Scientific Reports of the Great Barrier Reef Expedition 1928-29 4(14): 513-621 (British Museum (Natural History): London). 1934b. Sponges. Further Zoological Results of the Swedish Antarctic Expedition 1901-1903 3(2): 1-58 (Norstedt and SOner: Stockholm). 1934c. Report on the sponges of the Norwegian expeditions to East-Greenland (1930, 1931, and 1932). Skrifter om Svalbard og Ishavet. Zoological Results of the Norwegian Scientific Expeditions to East-Greenland (III) (61): 1-33. 1935a. The Family Plocamiidae with descriptions of four new genera of sponges. Annals and Magazine of Natural History (15) 87: 399-404. MEMOIRS OF THE QUEENSLAND MUSEUM 594^ zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA 1935b. Notes on British sponges with a description of a new genus and species. Annals and Magazine of Natural History (15) 90: 651-653. I935c. Some sponges from the Okhotsk Sea and the Sea of Japan. Issledovaniya Morei SSSR (Explorations des Mers d'U.R.S.S., Leningrad) 22: 61-79. 1936. Notes on sponges from South Africa, with descriptions of new species. Annals and Magazine of Natural History 17: 141-147. 1938a. Supplement to the littoral fauna of Krusadai Island in the Gulf of Manaar. Porifera. Bulletin of the Madras Government Museum (n.s.), Natural History Section 1(2): 1-58. 1938b. Non-calcareous sponges. Australian Antarctic Expedition 1911-14, Scientific Reports (C, Zoology and Botany): 9(5): 5-23. 1940. Las Esponjas Marinas del Museo Argentino de Ciencias Naturales. Anales Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" 40:95-121. 1950. The synonymy and distribution of Myxilla fimbriata (Bowerbank), with notes on other related species of Sponges. Annals and Magazine of Natural History (12) 3(34): 888-892. 1956. The sponges of west Africa. Atlantide Report. Scientific Results of the Danish Expedition to the coasts of Tropical West Africa 1945-46 4: 1 1 1 147. (Danish Science Press: Copenhagen). 1959a. Sponges. In Scientific Reports of the John Murray Expedition 1933-34. 10(5 ): 151-281 (British Museum (Natural History): London). 1959b. Spongia. The zoology of Iceland 2(3-4): 1-69. BURTON, M. & RAO, H.S. 1932. Report on the shallow-water marine sponges in the collection of the Indian Museum. Records of the Indian Museum 34(3): 299-356. CABEROY, R.A. 1981. The survey of class Demospongiae in Tayabas Bay. Zoological Papers, National Museum of the Philippines (7): 1-56. CABIOCH, L. 1968a. Contribution a la connaissance de la faune des Spongiaires de la Manche occidentale. Demosponges de la region de Roscoff. (Travaux de la Station Biologique de Roscoff. Nouvelle Serie 18.) Cahiers de Biologie Marine 9: 211-246. 1968b. Contribution a la connaissance des peuplements benthiques de la Manche Occidentale. (Travaux de la Station Biologique de Roscoff. Nouvelle Serie 18). Cahiers de Biologie Marine 9: 493-720. CAPON, R.J. & MACLEOD, J.K. 1987. 5-Thio-DMannose from the marine sponge Clathria pyramida (Lendenfeld). The first example of a naturally occurring 5-Thiosugar. Journal of the Chemical Society, Chemical Communication 1987: 1200-1201. CARBALLEIRA, N.M., SHALABI, F. & MALDONADO, M.E. 1990. Identification of the new I8-hexacosenoic acid in the sponge Thalysias juniperina. Lipids 25(4): 235-237. CARPAY, M. 1986. The marine sponges of Tasmania. A checklist of a number of marine sponges, occurring along the Tasmanian coast. Pp. 1-77 (M.Sc. thesis, Department Bijz,ondere Dierkunde, Institute for Taxonomic Zoology: Amsterdam). CARTER, H.J. 1870a. On the ultimate structure of marine sponges. Annals and Magazine of Natural History (4) 6: 329-341. 18706. Notes on the sponges Grayella, Osculina, and Cliona. Annals and Magazine of Natural History (4) 5: 73-83. 1871a. On two undescribed sponges and two Esperiadae from the West Indies; also on the nomenclature of the calcisponge Clathrina Gray. Annals and Magazine of Natural History (4) 7: 268-283. 1871b. A descriptive account of three pachytragous sponges growing on the rocks of the south coast of Devon. Annals and Magazine of Natural History (4) 7: 1-15. 1872a. Description of two new sponges from the Philippine Islands. Annals and Magazine of Natural History (4) 10: 100-113. 18726. One two new sponges from the Antarctic Sea, and on a new species of Tethya from Shetland; together with observations on the reproduction of sponges commencing from zygosis of the sponge-animal. Annals and Magazine of Natural History (4) 9: 409-435. 1874a. Descriptions and figures of deep-sea sponges and their spicules from the Atlantic Ocean, dredged up on board H.M.S. 'Porcupine', chiefly in 1869; with figures and descriptions of some remarkable spicules from the Agulhos Shoal and Colon, Panama. Annals and Magazine of Natural History (4) 14: 207-221, 245-257. 1874b. Development of the marine sponges from the earliest recognizable appearance of the ovum to the perfected individual. Annals and Magazine of Natural History (4) 14: 321-337, 389-406. 1874c. Further instances of the sponge-spicule in its mother cell. Annals and Magazine of Natural History (4) 14: 456-458. 1875. Notes introductory to the study and classification of the Spongida. Annals and Magazine of Natural History (4) 16: 1-40, 126-145, 177-200. 1876. Descriptions and figures of deep-sea sponges and their spicules, from the Atlantic Ocean, dredged up on board H.M.S. 'Porcupine', chiefly in 1862 (concluded). Annals and Magazine of Natural History (4) 18: 226-240, 307-324, 388410, 458-473. 1878. Parasites of the Spongida. Annals and Magazine of Natural History (5) 2: 157-172. 1879. Contributions to our knowledge of the Spongida. Annals and Magazine of Natural History (5) 3: 284-304, 343-360. 1880a. Report on specimens dredged up from the Gulf of Manaar and presented to the Liverpool REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS ^ 595 Free Museum by Capt. W.H. Cawne Warren. Annals and Magazine of Natural History (5) 6: 34-61, 129-156. 1880b. Sponges. (Including list of sponges dredged by the Birmingham Natural History and Microscopical Society, Falmouth Excursion, 1879. Depth 15-20 Fathoms). Midland Naturalist 1880: 55-60, 190-195. 1880c. Note on the so-called `Farringdon' (Coral Rag) sponges. Annals and Magazine of Natural History (5) 4: 431-437. 1881a. Supplementary report on specimens dredged up from the Gulf Manaar, together with others from the sea in the vicinity of the Basse Rocks and from Bass's Straits respectively, presented to the Liverpool Free Museum by Capt. H. Cawne Warren. Annals and Magazine of Natural History (5) 7: 361-385. 1881b. Contributions to our knowledge of the Spongida. Order II. Ceratina. Annals and Magazine of Natural History (5) 8: 101-121. 1882a. Some sponges from the West Indies and Acapulco in the Liverpool Free Museum described, with general and classificatory remarks. Annals and Magazine of Natural History (5) 9: 266-301, 346-368. 1882b. New Sponges, Observations on old ones, and a proposed new group. Annals and Magazine of Natural History (5) 10: 106-125. 1883a. Contributions to our Knowledge of the Spongida. - Pachytragida. Annals and Magazine of Natural History (5) 11: 344-369, pls 14-15. 1883b. Contributions to our knowledge of the Spongida. Annals and Magazine of Natural History (5) 12: 308-329. 1884a. Catalogue of marine sponges, collected by Mr. Jos. Willcox, on the west coast of Florida. Proceedings of the Academy of Philadelphia 1884: 202-209. 1884b. Generic characters of the sponges described in Carter's 'Contribution to our knowledge of the Spongida.' Annals and Magazine of Natural History (5) 13: 129-130. 1885a. New sponges from South Australia. Annals and Magazine of Natural History (5)13: 465-512. 1885b. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History (5) 15: 107-117. 1885c. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 15: 196-222. 1885d. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 15: 301-321. 1885e. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 16: 277-294. 1885f. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 16: 347-368. 1886a. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 17: 40-53. 1886b. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 17: 112-127. 1886c. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 17: 431-441. 1886d. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 17: 502-516. 1886e. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 18: 34-55. 1886f. Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History (5) 18: 126-149. 1886g. Supplement to the Descriptions of Mr. J. Bracebridge Wilson's Australian Sponges. Annals and Magazine of Natural History (5) 18: 271-290, 369-379, 445466. 1887a. Report on the marine sponges, chiefly from King Island, in the Mergui Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S., Superintendent of the Museum. Journal of the Linnean Society of London, Zoology 21: 61-84. I 887b. On the reproductive elements of the Spongida. Annals and Magazine of Natural History(S) 19: 350-360. 1889a. Sketch of the history of known fossil sponges in relation to those of the present day. Annals and Magazine of Natural History (6) 4: 280-290. I 889b. A correction in British spongology. Annals and Magazine of Natural History (6) 4: 249-250. CARTER, H.J. & HOPE, R. 1889. On a new British species of zyxwvutsrqponmlkjihgfedcbaZYXWVUTS Micr ociona Bk., in which the ends of the Tricurvate are Spiniferous & c. Annals and Magazine of Natural History (6) 3: 99-106. CAULDWELL, C.B., HENKART, P. & HUMPHREYS, T. 1973. Physical properties of sponge aggregation factor. A unique proteoglycan complex. Biochemistry, Pa 12(16): 3051-3055. CHATIN, J. 1890. Contribution a l'etude du moyan chez les Spongiaires. Comptes Rendus, Paris I 1 1 : 889890. CHEN, Y.-H. & MOK, H.-K. 1993. First record of the poecilosclerid sponge Rhapidophlus schoenus (de Laubenfels, 1936) (Poecilosclerida: Clathridae) MEMOIRS OF THE QUEENSLAND MUSEUM 596zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ from Taiwan. Bulletin of the Institute of Zoology Academia Sinica (Taipei) 32(4): 278-280. CHERNOFF, H. 1987. Factors affecting mortality of the scallop Chlamys asperrima (Lamarck) and its epizooic sponges in South Australian waters. Journal of Experimental Marine Biology and Ecology 109(2): 155-172. CHONG, A.S.F., PARISH, C.R. & COOMBE, D.R. 1987. Evidence that the cytoskeleton plays a key role in cell adhesion. Immunology and Cell Biology 65(1): 85-95. CIMINO, G., DE STEFANO, S., MINALE, L., ROCCIO, R., HIROTSU, K. & CLARDY, J. 1979. Two novel sesterterpene hydroxyquinols from the sponge Microciona toxistyla. Tetrahedron Letters (38): 3619-3622. CLAUS, C.F.W. 1868. Ueber Euplectella aspergillum (R. Owen). Pp. 1-28. Ein Beitrag zur Naturgeschichte der Kieselschwamme. (N.G. Elwertische Universitats-Buchhandlung: Marburg). COLLIER, J.R. 1983. The molecular weight of ribosomal ribonucleic acids among the Protostomia. Biological Bulletin of the marine biological Laboratory, Woods Hole 164(3): 428-432. COOMBE, D.R., JAKOBSEN, K.B. & PARISH, C.R. 1987. A role for sulfated polysaccharide recognition in sponge cell aggregation. Experimental Cell Research 170(2): 381-401. COTTE, J. 1903. Contribution a l'etude de la nutrition chez les Spongiaires. Bulletin des Sciences de France et Belgique 38: 420-573. COUES, E. & YARROW, H.C. 1879. Notes on the Natural History of Fort Macon, N.C., and vicinity. Proceedings of the Academy of Natural Sciences of Philadelphia 1878: 297-315. CRIVELLI, G.B. 1863. Di alcuni Spongiari del Golfo di Napoli. Memoria del Professore Crivelli. Atti della Societa Italia 5: 284-302. CUARTAS, E.I. 1992. Poriferos de la provincia biogeografica Argentina. III. Poecilosclerida (Demospongiae), del litoral Marplatense. Physis (Buenos Aires) (A) 47(113): 73-88. CUARTAS, E.I. 1993. Poriferos intermareales de San Antonio Oeste provincia de Rio Negro, Argentina (Porifera: Demospongiae). Neotropica (La Plata) 38(100): 111-118. CUENOT, L. 1903. Contribution a la faune du Bassin D'Arcachon. 3. Doridiens. Travaux du Laboratoire Arcachon 1903: 4. CZERNIAWSKY, V. 1880. Spongiae littorales Portis Euxini et Maris Caspii. Bulletin de la Societe Naturelle du Moscou 54: 88-128, 228-320. DALWITZ, M.J. & PAINE, T.A. 1986. User's guide to the DELTA system. A general system for processing taxonomic descriptions. CSIRO, Australia, Division of Entomology Report (13): 1-106 DAWSON, E.W. 1993. Index to the fauna: 2. Porifera New Zealand Oceanographic Institute Memoir (100): 1-98. WEERDT, W.H. DE. 1985. A systematic revision of the north eastern Atlantic shallow-water Haplosclerida (Porifera, Demospongiae), Part 1: Introduction, Oceanapiidae and Petrosiidae. Beaufortia 35(5): 61-91. DENDY, A. 1887. Report on a zoological collection made by the officers of H.M.S. 'Flying-Fish' at Christmas Island Indian Ocean. IX. Porifera. Proceedings of the Zoological Society of London 3: 524-526. 1889a. An alphabetical list of the genera and species of sponges described by H.J. Carter, Esq., F.R.S., together with a number of his more important references to those of other authors, with an introductory notice. Proceedings of the Royal Society of Victoria (n.s.) 1: 34-59. 1889b. Report on a Second Collection of sponges from the Gulf of Manaar. Annals and Magazine of Natural History (6) 3: 73-99. 1889c. Some old and new questions concerning sponges. Zoologischer Anzeiger 13(325): 14-18. 1895. Catalogue of non-calcareous sponges collected by J. Bracebridge Wilson, Esq., M.A., in the neighbourhood of Port Phillip Heads. Part 1. Proceedings of the Royal Society of Victoria ser. 2, 7: 232-260. 1896. Catalogue of non-calcareous sponges collected by J. Bracebridge Wilson, Esq., M.A., in the neighbourhood of Port Phillip Heads. Part 2. Proceedings of the Royal Society of Victoria ser.2, 8: 14-51. 1897. Catalogue of non-calcareous sponges collected by J. Bracebridge Wilson, Esq., M.A., in the neighbourhood of Port Phillip Heads. Part 3. Proceedings of the Royal Society of Victoria ser.2, 9: 230-259. 1905. Report on the sponges collected by Professor Herdman, at Ceylon, in 1902. Pp 57-246 in Herdman, W.A. Report to the Government of Ceylon on the pearl oyster Fisheries of the Gulf of Manaar. 3(18):57-246 (Royal Society: London). 1916a. Report on the non-calcareous sponges collected by Mr. James Hornell at Okhamandal in Kattiawar in 1905-1906. In Report to the Government of Baroda on the Marine Zoology of Okhamandal. 2(17): 96-146. 1916b. On the occurrence of gelatinous spicules, and their mode of origin, in a new genus of siliceous sponges. Proceedings of the Royal Society of London 89: 315-322. 1921. The tetraxonid sponge spicule: - a study in evolution. Acta Zoologica (Stockholm) 2: 95152. 1922. Report on the Sigmatotetraxonida collected by H.M.S. `Sealark' in the Indian Ocean. Transactions of the Linnean Society of London, Zoology 18: 1-164 in Reports of the Percy Sladen Trust Expedition to the Indian Ocean in 1905, Vol. 7. 1924a. Porifera. Part I. Non-Antarctic sponges. British Antarctic (Terra Nova') Expedition, 1910. Natural history report. 6(3): 269-392 REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 597 (British Museum (Natural History), Zoology: London). 1924b. On an orthogenetic series of growth-forms in certain tetraxonid sponge spicules. Proceedings of the Royal Society of London (B) 97: 243-250, pls 10-11. DENDY, A. & FREDERICK, L.M. 1924. On a collection of sponges from the Abrolhos Islands, Western Australia. Journal of the Linnean Society of London, Zoology 35: 477-519. DESCATOIRE, A. 1969. Les peuplements sessiles de l'Archipel de Glenan de l' infralittoral rocheux. II. - Notes Systernatiques a propos de l'inventaire des Spongiaires. Vie et Milieu (A, Biologic Marine) 20: 9-30. DESOR, E. 1851. Description of two new sponges: Spongia urceolata and Spongia sulphurea. Proceedings of the Boston Society of Natural History 3: 67-68. DESQUEYROUX, R.P. 1972. Demospongiae (Porifera) de la costa de Chile. Gayana (Zoologia) (20): 1-71. 1975. Esponjas (Porifera) de la region Antarctica Chilena. Cahiers de Biologic Marine 16(1): 4782. DESQUEYROUX-FAUNDEZ, R. 1981. Revision de la collection d'eponges d'Amboine (Moluques, Indonesie) constituee par Bedot and Pictet et conservee au Museum d'histoire naturelle de Geneve. Revue Suisse de Zoologie 88(3): 723764. DESQUEYROUX-FAUNDEZ, R. & MOYANO, H. 1987. Zoogeografia de demospongias Chilenas. Boletin de la Sociedad de Biologia de Concepcion 58: 39-66. DESQUEYROUX-FAUNDEZ, R. & STONE, S.M. 1992. O. Schmidt Sponge Catalogue. An illustrated guide to the Graz Museum collection, with notes on additional material. Pp. 1-190 (Museum d'Histoire naturelle: Geneva). DICKINSON, M.G. 1945. Sponges of the Gulf of California. Reports on the collections obtained by Allan Hancock Pacific Expeditions of Velcro III off the coast of Mexico, Central America, South America, and Galapagos Islands in 1932-40. 11(1): 1-5. DRAGNEWITSCH, P. 1905. Spongien von Singapore. Pp. 1-36. (Inaugural Dissertation: Bern). 1906. Spongien von Singapore. Zoologische Jahbucher Jena Abteilung fiir Systematik, Okologie, Geographic der Tiere 23: 439-448. DUCHASSAING, P. DE FONBRESSIN 1850. Animaux radiares des Antilles. Pp. 1-35 (Typographic Plon Freres: Paris). DUCHASSAING, DE FONBRESSIN, P. & MICHELOTTI, G. 1864. Spongiaires de la mer CaraIbe. Naturk. Verh. Holland Maatsch. Wetensch Haarlem (2) 21(3): 1-124. DUNHAM, P., ANDERSON, C., RICH, A.M. & WEISSMANN, G. 1983. Stimulus-response coupling in sponge cell aggregations: evidence for calcium as an intracellular messenger. Proceedings of the National Academy of Science, USA (Biological Sciences) 80(15): 4756-4760. DUNHAM, P.B., VOSSHALL, L.B., BAYER, C.A., RICH, A.M. & WEISSMANN, G. 1985. From Beaumont to Poison ivy: marine sponge cell aggregation and the secretory basis of inflammation. Federation Proceedings 44(14): 2914-2924. DYBOWSKY, W. 1880. Studien iiber die Spongien des russischen Reiches, mit besonderer Berticksichtigung der Spongien-Fauna des Baikal-Sees. Memoires de l'Academia Imperiale des Sciences de St. Petersbourg 27: 45-50. EHLERS, F. 1870. Die Esper'schen Spongien. Pp. 1-36. In Zoologischen Sammlung der K. Universitat Erlangen. Programm zum Eintritt in der Senat der KOniglichen Friedrich-Alexanders-Universitat in Erlangen: Erlangen. ELLIS, J. & SOLANDER, D. 1786. The Natural History of many curious and uncommon zoophytes, collected from various parts of the globe. Systematically arranged and described by the late Daniel Solander. Pp. 1-206 (London). ESPER, E.J.C. 1794. Die Pflanzenthiere in Abilungen nach der Natur mit Fabben erleuchtet nebst Beschreibungen. Zweyter Theil. Pp. 1-303 (Niirnberg). 1797. Fortsetzungen der Pflanzenthiere in Abbildungen nach der Natur mit Farben erleuchtet nebst Beschreibungen. (Erster Theil: Niirnberg). EVANS, C.W. & BERGQUIST, P.R. 1977 Are-evaluation of the relevance of acid mucopoly-saccharides in sponge taxonomy. Cahiers de Biologic Marine 28: 191-199. FELL, H.B. 1950. The Kirk collection of sponges (Porifera) in the Zoology Museum, Victoria University College. Zoological Publications of the Victoria University College, New Zealand 4: 1-12. 1984. Porifera. in Adyodi, K.G. and Adyodi, R.G. (eds), 'Reproductive Biology of Invertebrates, Vol. IV: Fertilization and Larval Development'. (John Wiley and Sons: Chichester, Sussex). 1990. 1. Porifera. Pp. 1-44. In Adiyodi, K.G. and Adiyodi, R.G. (eds) 'Reproductive Biology of Invertebrates'. Volume VI, Part A. Asexual propagation and reproductive strategies. (Oxford & IBH Publishing Co.: New Delhi, Bombay, Calcutta). 1990. Tolerances of the dormant forms of some estuarine sponges, notably Microciona prolifera. Pp. 497-503. In Riitzler, K. (ed.) 'New perspectives in sponge biology'. (Smithsonian Institution Press: Washington D.C.). FELL, P.E., KNIGHT, P.-A. & RIEDERS, W. 1989. Low-salinity tolerance of and salinity-induced dormancy in the estuarine sponge Microciona prolifera (Ellis & Solander) under long-term laboratory culture. Journal of Experimental Marine Biology and Ecology 133(3): 195-211. FERRER HERNANDEZ, F. 1914. Esponjas del Cantabrico. Parte Segunda. III: Myxospongida; MEMOIRS OF THE QUEENSLAND MUSEUM 598zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ IV: Tetraxonida; V: Triaxonida. Trabajos Museo Nacional de Ciencias Naturales Madrid (ZoolOgica) (17): 1-46. 1918. DescripciOn de tres esponjas nuevas del litoral espaiiol. Revista de la Real Academia de Ciencias Exactas, Fisicas y Naturales de Madrid 16: 532540. 1921. Esponjas recogidas en la campana preliminar del `Giralda'. Boletin de Pescas Madrid 6: 161-177. 1923. Mas datas para el conocimiento de las esponjas de las costas espaiiolas. Boletin de Pescas Madrid 7: 247-272. FINKS, R.M. 1967. The structure of Saccospongia lazata Bassler (Ordovician) and the phylogeny of the Demospongea. Journal of Paleontology 41(5): 1137-1149. 1970. The evolution and ecological history of sponges during Palaeozoic times. Symposium of the Zoological Society of London 25: 3-22. FRISTEDT, K. 1885. Bidrag till Kannedomen, om de vid sveriges vestra Kust lefvande Spongiae. Kaniglichen Svenska Vetenskap Akademie Handlinger 21: 1-56. 1887. Sponges from the Atlantic and Arctic Oceans and the Behring Sea. Vega-Expeditionens Vetenskap Iakttagelser (Nordenskiiild) 4: 401471. FROMONT, J. 1989. Aspects of the reproductive biology of Xestospongia testudinaria (Great Barrier Reef). Proceedings of the 6th International Coral Reef Symposium, Australia, 1988, 2: 685-691 (James Cook University of North Queensland: Townsville). FROMONT, J.P. & BERGQUIST, P.R. 1990. Structural characters and their use in sponge taxonomy: when is a sigma not a sigma? Pp. 273-278. In Raizler, K. (ed.) 'New perspectives in sponge biology. (Smithsonian Institution Press: Washington D.C.). FRY, W.G. 1970. The sponge as a population: a biometric approach. Zoological Society of London Symposia 25: 135-161. 1971. The biology of larvae of Ophlitaspongia seriata from two North Wales populations. Pp. 155-178. In Crisp, D.J. (ed.) 'Fourth European Marine Biology Symposium'. (Cambridge University Press: Cambridge). 1973. The role of larval migration in maintenance of an encrusting sponge population. Netherlands Journal of Sea Research 7: 159-170. GALTSOFF, P.S. & PERTZOFF, V. 1926. Some physicochemical properties of dissociated sponge cells. Journal of General Physiology 1926: 239255. GEORGE, W.C. & WILSON, H.V. 1919. Sponges of Beaufort (N.C.) Harbor and vicinity. Fishery Bulletin, United States National Fisheries Service (36): 130-179. GRAEFFE, E. 1882. Uebersicht der Seethierfauna des Golfes von Triest nebst Notizen iiber Vorkommen, Lebensweise, Erscheinungs - und Fortpflan- zungszeit der einzelnen Arten, 2, Coelenteraten, Spongiariae. Arbeiten aus dem Zoologischen Institute der Universitat Wein 4: 313-321. GRANT, R.E. 1826. Observations on the Structure and Function of the Sponge. Edinburgh New Philosophical Journal 1826: 121-141. 1861. Tabular view of the primary division of the Animal Kingdom. (London). GRAY, J.E. 1848. List of the specimens of British sponges in the collection of the British Museum. Pp. 1-24. (British Museum: London). 1858. Description of a new genus of sponge (Xenospongia) from Torres Strait. Procedings of the Zoological Society of London 1858:229-230. 1867. Notes on the arrangement of sponges, with description of some new genera. Proceedings of the Zoological Society of London 1867:492-558. 1868. Observations on sponges and on their arrangement and nomenclature. Annals and Magazine of natural History (4) 1868: 161-173. 1869. Note on Ianthella, a new genus of keratose sponges. Proceedings of the Zoological Society of London 4: 49-51. 1870. Note on a new genus of sponge from West Australia. Annals and Magazine of natural History (4) 1870: 272. GREENBERG, M.J., REED, C. & PIERCE, S.K.Jr. 1977. Dissociated cells of Microciona prolifera (Porifera) are inhibited from reaggregation by cytochalasins, A, B, and E. Comparative Biochemistry and Physiology (C) 56(2): 95-102. GUILER, E.R. 1950. Notes on Tasmanian marine sponges. Records of the Queen Victoria Museum2: 5-14. HAJDU, E., WEERDT, W.H. DE & SOEST, R.W.M.VAN. I994b. Affinities of the 'Mermaid's Glove sponge lsodictya palmata, with a discussion on the synapomorphic value of chelae microscleres. Pp. 141-150. In Soest, R.W.M. van, Kempen, T.M.G. van and Braekman, J.C. (eds) 'Sponges in time and space'. (Balkema: Rotterdam). HAJDU, E., SOEST, R.W.M. VAN & HOOPER, J.N.A. 1994. Proposal of a phylogenetic subordinal classification of poecilosclerid sponges (Demospongiae, Porifera). Pp. 123-140. In Soest, R.W.M. van, Kempen, T.M.G. van and Braekman, J.-C. (eds) 'Sponges in time and space'. (Balkema: Rotterdam). HALLMANN, E.F. 1912. Report on the sponges obtained by the F.I.S. 'Endeavour' on the Coasts of New South Wales, South Australia, Queensland, and Tasmania, 1909-10. Part 1. Zoological Results of the Fishing Experiments carried out by the F.I.S. 'Endeavour' 1909-10 (2): 117-300, pls 21-36 1914a. A revision of the monaxonid species described as new in Lendenfeld's 'Catalogue of the Sponges in the Australian Museum'. Part 1. Proceedings of the Linnean Society of New South Wales 29: 263-315. REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT ^ 599 1914b. A revision of the monaxonid species described as new in Lendenfeld's 'Catalogue of the Sponges in the Australian Museum'. Part 2. Proceedings of the Linnean Society of New South Wales 29: 327-376. 1914c. A revision of the monaxonid species described as new in Lendenfeld's 'Catalogue of the Sponges in the Australian Museum'. Part 3. Proceedings of the Linnean Society of New South Wales 29: 398-446. 1916a. A revision of the genera with microscleres included, or provisionally included, in the Family Axinellidae, with descriptions of some Australian Species. Part 1. (Porifera). Proceedings of the Linnean Society of New South Wales 41: 453491. 1916b. A revision of the genera with microscleres included, or provisionally included, in the Family Axinellidae, with descriptions of some Australian Species. Part 2. (Porifera). Proceedings of the Linnean Society of New South Wales 41: 495552. 1916c. A revision of the genera with microscleres included, or provisionally included, in the Family Axinellidae, with descriptions of some Australian Species. Part 3. (Porifera). Proceedings of the Linnean Society of New South Wales 41: 634675. 1917. On the genera Echinaxia and Rhabdosigma [Porifera]. Proceedings of the Linnean Society of New South Wales 42: 391-405. 1920. New genera of monaxonid sponges related to the genus Clathria. Proceedings of the Linnean Society of New South Wales 44: 767-792. HANITSCH, R. 1889. Second Report on the Porifera of the L.M.B.C. District. Proceedings of the Biological Society of Liverpool 3: 155-173. 1890. Third report on the Porifera of the L.M.B.C. District. Transactions of the Biological Society of Liverpool 4: 192-238. 1894. Revision of the Generic Nomenclature and Classification in Bowerbank's 'British Spongiadae'. Transactions of the Liverpool Biological Society 8: 173-206. 1895. Notes on a collection of Sponges from the West Coast of Portugal. Transactions from the Liverpool Biological Society 9: 205-219. HARTMAN, W.D. 1955. A collection of sponges from the West Coast of the Yucatan Peninsula with descriptions of two new species. Bulletin of Marine Science of the Gulf and Caribbean 5(3): 161-189. 1958. Natural history of the marine sponges of southern New England. Bulletin of the Peabody Museum of Natural History 12: 1-155. 1979 . A new sclerosponge from the Bahamas and its relationships to Mesozoic stromatoporoids. Pp. 467-474. In Levi, C. and Boury-Esnault, N. (ecls) 'Biologic des spongiaires. Sponge biology'. (Colloques Intemationaux du Centre National de la Recherche Scientifique : Paris). 1981. Form and distribution of silica in sponges. Pp. 453-493. In Simpson, T.L. & Volcani, B.E. (eds) 'Silicon and siliceous structures in biological systems'. (Spinger-Verlag: New York). 1982. Porifera. Pp. 640-666. In Parker, S.P. (ed.) 'Synopsis and classification of living organisms'. Vol. 1 (McGraw-Hill: New York). HARTMAN, W.D. & GOREAU, T.F. 1970. Jamaican coralline sponges: their morphology, ecology and fossil relatives. Symposium of the Zoological Society of London 25: 205-243. 1975. A Pacific tabulate sponge, living representative of a new order of sclerosponges. Postilla 167: 1-21. HECHTEL, G. 1965. A systematic study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History 20: 1-103. 1976. Zoogeography of Brazilian marine Demospongiae. Pp. 237-260. In Harrison, F.W. & Cowden, R.R. (eds) 'Aspects of Sponge Biology'. (Academic Press: New York). HEIDER, A. VON. 1895. Liste der Schrnideschen Spongien in der Zoologischen Abtheilung des steiermarlcischen Landes-Museums. Mitteilungen des naturwissenschaftlichen Vereines fuer Steiermark 1894: 276-285. HELLER, C. 1864. Horae Dalmatinae. Verhandlungen der Zoologisch-botanischen Gesellschaft in Wien 14: 17-64. HENKART, P., HUMPHREYS, S. & HUMPHREYS, T. 1973. Characterization of sponge aggregation factor. A unique proleoglycan complex. Biochemistry, Pa 12(16): 3045-3050. HENTSCHEL, E. 1909. Tetraxonida. 1. Teil. 2(21): 347-402, pls 22-23 in Michaelsen, W. and Hartmeyer, R. (eds) Die Fauna SildwestAustraliens. (G. Fischer: Jena). 1911. Tetraxonida. 2. Teil. 3(10): 279-393 in Michaelsen, W. and Hartmeyer, R. (eds) Die Fauna Sadwest-Australiens (G. Fischer: Jena). 1912. Kiesel- und Homschwamme der Aru und Kei-Inseln. Abhandlungen Senckenbergiana naturforschende Gessellschaft 1912: 295-448. 1914. Monaxone Kieselschwamme und Homschwamme der Deutschen Siidpolar-Expedition 1901-1903. Deutsche Sudpolar-Expedition 1901-1903 15 (Zoologie 7): 37-141. 1923. Erste Unterabteilung der Metazoa. Parazoa. Einziger Stamm und einzige Klasse der ersten Unterabteilung: Porifera = Schwamme. Handbuch der Zoologie 1: 307417. 1929. Die Kiesel- und Hornschwamme der NOrdlichen meeres. Fauna Arctica, Jena (4)5(4): 859-1042, pls 12-14. HEWATT, W.G. 1946. Marine ecological studies on Santa Cruz Island, California. Ecological Monographs, Durham, N.C. 16(3): 1-197. HIEMSTRA, F. & HOOPER, J.N.A. 1991. Additions to the Indo-Australian representatives of Acarnus Gray (Porifera: Demospongiae: Poecilosclerida), MEMOIRS OF THE QUEENSLAND MUSEUM 600zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA ^ with description of a new species. Memoirs of the Queensland Museum 30(3): 433-442. HOOPER, J.N.A. 1984a. A new genus and two new species of haplosclerid sponges (Porifera: Demospongiae) from the Timor Sea, northwest Australia. Proceedings of the Royal Society of Victoria 96(2): 55-60. 1984b. Sigmaxinella soelae and Desmacella ithystela, two new desmacellid sponges (Porifera, Axinellida, Desmacellidae) from the Northwest Shelf of Western Australia, with a revision of the Family Desmacellidae. Northern Territory Museum of Arts and Sciences. Monograph Series (2): 1-58. 1986a. Revision of the marine sponge genus Axos Gray (Demospongiae: Axinellida) from northwest Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 3(1): 167-189. 1986b. A new species of Timea Gray (Porifera: Hadromerida) from northern Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 3(1): 191194. 1987. New records of Acarnus Gray (Porifera: Demospongiae: Poecilosclerida) from Australia, with a synopsis of the genus. Memoirs of the Queensland Museum 25(1): 71-105. 1988a. Structural features of the benthic community of East Point Reef Fish Reserve. A comparative study between oceanic, near-shore and inshore reefs of northwest Australia. Australian National University North Australian Research Unit Mangrove Monograph 4: 214-225. 1988b. Character Stability, Systematics and Affinities Between Microcionidae (Poecilosclerida) and Axinellida (Porifera: Demospongiae). A Revision of the Australasian Microcionidae. PhD Thesis, University of Queensland, Department of Zoology: 1-1208. 1990a. Character stability systematics and affinities between Microcionidae (Poecilosclerida) and Axinellida. Pp. 284-294. In Riitzler, K. (ed.) 'New perspectives in sponge biology'. (Smithsonian Institution Press: Washington D.C.). 1990b. A new species of Rhabderemia Topsent (Porifera: Demospongiae) from the Great Barrier Reef. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 7(1): 65-78. 1991. Revision of the Family Raspailiidae (Porifera: Demospongiae), with description of Australian species. Invertebrate Taxonomy 5(6): 1179-1415. 1994. Coral reef sponges of the Sahul Shelf - a case for habitat preservation. Memoirs of the Queensland Museum 36(1): 93-106. HOOPER, J.N.A. &BERGQUIST, P.R. 1992. Cymbastela, a new genus of lamellate coral reef sponges. Memoirs of the Queensland Museum 32(1): 99137. HOOPER, J.N.A. & KRASOCHIN, V.B. 1989. Redescription of the burrowing sponge Zyzzya massalis (Dendy) from the Seychelles and Houtman Abrolhos Islands. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 6(1): 133-140. , HOOPER, J.N.A. & LEVI, C. 1989. Esperiopsis desmophora n.sp. (Porifera: Demospongiae): a desma-bearing Poecilosclerida. Memoirs of the Queensland Museum 27(2): 437-441. 1993a. Poecilosclerida from the New Caledonia lagoon (Porifera: Demospongiae). Invertebrate Taxonomy 7(5): 1221-1302. 1993b. Axinellida from the New Caledonia lagoon (Porifera: Demospongiae). Invertebrate Taxonomy 7(6): 1395-1472. 1994. Biogeography of Indo-west Pacific sponges: Microcionidae, Raspailiidae, Axinellidae. Pp. 191-212. In Soest, R.W.M. van, Kempen, T.M.G. van and Brockman, J.-C. (eds) 'Sponges in Time and Space'. (Balkema: Rotterdam). HOOPER, J.N.A., CAPON, R.J. & HODDER, R.A. 1991. A new species of toxic marine sponge (Porifera: Demospongiae: Poecilosclerida) from northwest Australia. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 8(1): 27-36. HOOPER, J.N.A., CAPON, R.J., KEENAN, C.P. & PARRY, D.L. 1990. Biochemical and morphometric differentiation of two sympatric sibling species of Clathria (Porifera: Demospongiae) from northern Australia. Invertebrate Taxonomy 4: 123-148. 1991. Morphometric and biochemical differences between sympatric populations of Clathria `spicata' species complex (Demospongiae: Poecilosclerida: Microcionidae) from northern Australia. Pp. 271-288. In Keupp, H. & Reitner, J. (eds) 'Fossil and Recent sponges. (SpringerVerlag: Berlin, Heidelberg). HOOPER, J.N.A., CAPON, R.J., KEENAN, C.P., PARRY, D.L. & SMIT, N. 1992. Chemotaxonomy of marine sponges: families Microcionidae, Raspailiidae, and Axinellidae, and their relationships with other families in the orders Poecilosclerida and Axinellida (Porifera: Demospongiae). Invertebrate Taxonomy 6: 261301. HOOPER, J.N.A., KELLY-BORGES, M. & RIDDLE, M. 1993. Oceanapia sagittaria from the Gulf of Thailand. Memoirs of the Queensland Museum 33(1): 61-72. HOOPER, J.N.A. & WIEDENMAYER, F. 1994. Porifera. Zoological Catalogue of Australia. 12: 1-621. HOPE, R. 1889. On two new British Species of Sponges, with short notices of an ovigerous specimen of Hymeniacidon Dujardinii, Bowk., and of a Fossil Toxite. Annals and Magazine of Natural History (6) 4: 333-342. HOPKINS, S.H. 1956. Notes on the boring sponges in gulf coast estuaries and their reaction to salinity.

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