Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae

John N . A . Hooper; Belinda Glasby

Fifteen species in six genera of the family Halichondriidae, including two new species, Halichondria (Halichondria) carotenoidea sp. nov. and Halichondria (Halichondria) microbiana sp. nov., are recorded for northern Australia as part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. Descriptions and discussion of those species are presented here. Eight new combinations within the family Halichondriidae are here established, i.e. Amorphinopsis fenestrata (Ridley, 1884, as Leucophloeus), Amorphinopsis maculosa (Pulitzer-Finali, 1996, as Topsentia), Axinyssa bergquistae (Hooper et al., 1997, as Halichondria), Axinyssa mertoni (Hentschel, 1912, as Ciocalypta), Axinyssa gracilis (Hentschel, 1912, as Ciocalypta rutila gracilis), Axinyssa terpnis (De Laubenfels, 1954, as Phycopsis), Ciocalypta vansoesti (Hooper et al., 1997, as Halichondria) and Topsentia ridleyi (Hooper et al., 1997 as Halichondria) and one species is relocated into the family Dictyonellidae, i.e, Stylissa vernonensis Hooper et al., 1997, as Hymeniacidon). A lectotype is designated for Ciocalypta stalagmites Hentschel, 1912.

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 27: 55–84 Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae BELINDA ALVAREZ1 and JOHN N. A. HOOPER2 1 Museum and Art Gallery of the Northern Territory. GPO Box 4646, Darwin NT 0801, AUSTRALIA belinda.glasby@nt.gov.au 2 Queensland Museum. PO Box 3300, South Brisbane QLD 4101, AUSTRALIA john.hooper@qm.qld.gov.au ABSTRACT Fifteen species in six genera of the family Halichondriidae, including two new species, Halichondria (Halichondria) carotenoidea sp. nov. and Halichondria (Halichondria) microbiana sp. nov., are recorded for northern Australia as part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. Descriptions and discussion of those species are presented here. Eight new combinations within the family Halichondriidae are here established, i.e. Amorphinopsis fenestrata (Ridley, 1884, as Leucophloeus), Amorphinopsis maculosa (Pulitzer-Finali, 1996, as Topsentia), Axinyssa bergquistae (Hooper et al., 1997, as Halichondria), Axinyssa mertoni (Hentschel, 1912, as Ciocalypta), Axinyssa gracilis (Hentschel, 1912, as Ciocalypta rutila gracilis), Axinyssa terpnis (De Laubenfels, 1954, as Phycopsis), Ciocalypta vansoesti (Hooper et al., 1997, as Halichondria) and Topsentia ridleyi (Hooper et al., 1997 as Halichondria) and one species is relocated into the family Dictyonellidae, i.e, Stylissa vernonensis (Hooper et al., 1997, as Hymeniacidon). A lectotype is designated for Ciocalypta stalagmites Hentschel, 1912. Keywords: sponge, Porifera, Halichondrida, Halichondriidae, northern Australia, new species, taxonomy. INTRODUCTION The family Halichondriidae was revised by Erpenbeck & Van Soest (2002). Its deinition is based entirely on a few skeletal characters, i.e. presence of an ectosome with specialised skeleton, a disorganised choanosomal skeleton, dimensions of oxeas and styles, absence of microscleres. Other morphological characters (e.g. general shape of megascleres, presence or size categories among the megascleres, spicule density and its relation to consistency, orientation of spicules at the ectosomal level) are used in combination with the diagnostic characters to separate species and genera within this family. All these characteristics are very simplistic, and often displayed as a gradient of variation which makes separation of species very subjective as relected by the large number of synonyms (see Van Soest et al. 2008). The family includes 14 genera and at least 296 valid species (Van Soest et al. 2008); however, the status and generic allocation of many of the species listed on this database on the Internet needs to be revised and validated against the current concept of the genera. The Australian Faunal Directory (Hooper 2005) currently lists a total of 42 species of Halichondriidae. Twenty species were reported and described for the Beagle Gulf (northern Australia) by Hooper et al. (1997). No other revision of species of Halichondriidae in Australia or adjacent areas is presently available. The present paper represents the third part of a revision of the order Halichondrida in the northern Australian region and includes the family Halichondriidae. Alvarez & Hooper (2009, 2010) provided details and presented an introduction to the revision of the order and the families Axinellidae and Dictyonellidae. MATERIALS AND METHODS 55 This revision includes material of the family Halichondriidae recorded for the tropical northern Australian waters of the Northern Territory and Queensland coast (from Admiralty Gulf in the west to Torres Strait in the east, approx. between the 125º E and 142º E). Complete locality and collection data for non-type voucher material deposited at the Queensland Museum and the Museum and Art Gallery of the Northern Territory are available in Appendix 1. The distribution of species is given according to the marine provinces deined by Spalding et al. (2007). Spicule measurements are in micrometres and are based on 25 spicules (unless indicated in square brackets) and denoted as range (and mean ± 1 S.E.) of spicule length and width. All other methods as discussed in Alvarez & Hooper (2009, 2010). B. Alvarez and J. N. A. Hooper A B C D E G F H I Fig. 1. A, Amorphinopsis excavans, specimen in situ at South Shell I., Darwin Harbour, N.T; B, Amorphinopsis fenestrata, specimen in situ at Weed Reef, Darwin Harbour, N.T. C, Amorphinopsis maculosa: Specimen (QM G313577) dredged at Groote Eylandt, Gulf of Carpentaria, Queensland; D, Axinyssa bergquistae, specimen (NTM Z. 5976) in situ at Stevens Rock, Darwin Harbour, N.T.; E, Axinyssa mertoni, specimen in situ at South Shell I., Darwin Harbour, N.T.; F, Ciocalypta heterostyla, specimen in situ at Channel Rock, Darwin Harbour, N.T.; G,H, C. stalagmites, specimens with different colouration in situ at Channel Rock and Stevens Rock, respectively; I, C. vansoesti, specimen in situ at Channel Rock, Darwin, N.T. Photographs: A,B – M. Browne; D,G–I – H. Nuguyen, H,E,F – B. Alvarez. 56 Halichondriidae from northern Australia ABBREVIATIONS Abbreviations used in the manuscript are: AIMS, Australian Institute of Marine Sciences, Townsville; BMNH, Natural History Museum, London (formerly British Museum (Natural History)); CRRF, Coral Reef Research Foundation, Palau; GBR, Great Barrier Reef, MAGNT, Museum and Art Gallery Northern Territory; MNHN, Musée National d’Historie Naturelle, Paris, France; MSNG, Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy; MAGNT/NTM, Museum and Art Gallery Northern Territory (formerly Northern Territory Museum), Darwin, Northern Territory, Australia; NHMB – Naturhistorisches Museum, Basel, Switzerland; SMF, Senckenberg Research Institute and Natural History Museum, Frankfurt; QLD, Queensland, Australia; QM, Queensland Museum, Brisbane; WA, Western Australia; ZMA, Zoologisch Museum, University of Amsterdam, ZMB, Museum für Naturkunde and der Universität Humboldt zu Berlin, Berlin, Germany. Numbers preixed with Q666C, 0CDN, 0M9H are the cross-reference sample number collected for the United States National Cancer Institute, under the ‘Collection of shallow-water organisms’ programme, by the Australian Institute of Marine Sciences, CRRF and MAGNT (subcontracted through CRRF) respectively. TAXONOMY Family Halichondriidae Gray, 1867 Genus Amorphinopsis Carter, 1887 Gender feminine. Type species, by monotypy, Amorphinopsis excavans Carter, 1887. Recent, Indian Ocean. Amorphinopsis excavans Carter, 1887 (Figs 1A, 2) Amorphinopsis excavans Carter, 1887: 77; Hooper & Wiedenmayer 1994: 205; Hooper et al. 1997: 25; Erpenbeck & Van Soest 2002: 791; Lim, de Voogd & Tan 2008: 115. Amorphinopsis sacciformis.– Hooper et al. 1997: 27. Material examined. Darwin Harbour: NTM Z.2215, Z.4093, Z.4125 (0CDN -8016-W), Z.5213 (0M9H2184-Q); Z.5222 (0M9H2251-O), Z.5736. Vernon Is: G303658. Description Shape (Fig. 1A). Thinly to thickly encrusting (up to 50 mm thick), massive to lobate, or developing short projections and small lumps, generally growing in patches and following substrate, semi-buried in substrate. Colour. Olive green, yellow inside. Oscula. Round to ovate, inconspicuous, 10 mm diameter. Surface. Hispid, bumpy. Skeleton. Ectosomal skeleton (Fig. 2A) thin, detachable tangential layer, composed by a disorganised criss-cross reticulation of paucispicular-multispicular tracts of oxeas, 57 up to 100 µm thick, with small styles tangentially to paratangentially oriented, sometimes in disorganised tufts. Choanosomal skeleton (Fig. 2B) halichondroid, with large oxeas oriented in all directions, sometimes grouped in directionless multispicular tracts; slightly cavernous at subectosomal area with short multispicular tracts supporting the ectosomal skeleton. Spicules (Fig 2C). Oxeas, hastate, in a large range of sizes, 213.4–945.3 µm (598±221) x 5.9–25.1 µm (16.3±5.5). Smaller ectosomal styles 140.8–264.9 µm (193.8±41.3) x 4.1–7.5 µm (5.5±0.9). Remarks. The material examined here agrees with the description of the syntype by Erpenbeck & Van Soest (2002: 790). The specimen described under Amorphinopsis sacciformis by Hooper et al. (1997) is better allocated to A. excavans. It does not agree with the syntype of Ciocalypta sacciformis Thiele which, as mentioned below (see under remarks of A. carpentariensis), should be interpreted as Halichondria. The material described by Hooper et al. (1997) is a thin crust covering a bivalve shell with skeleton similar to the rest of the material assigned here to A. excavans. Species considered synonyms of Amorphinopsis excavans by Hooper & Wiedenmayer 1994, following Burton (1959), were excluded by Hooper et al. (1997) and Erpenbeck & Van Soest (2002). Distribution. The species was recorded originally from the Mergui Archipelago (Andaman province). Records from Singapore and northern Australia extend the distribution of the species to the Sunda and Sahul Shelf provinces. The northern Australian and Singaporean populations seem to be common at the intertidal region associated with piers and wharfs. The species is found also subtidally between 9 and 16 m. Amorphinopsis fenestrata (Ridley, 1884) comb. nov. (Figs 1B, 3) Leucophloeus fenestratus Ridley, 1884: 464; Dendy 1922: 124; Burton 1928: 127. Leucophloeus fenestratus unnamed variety. – Ridley 1884: 464. Suberites oculatus Kieschnick, 1896: 534. Hymeniacidon fenestratus. – Lindgren 1897: 483; Lindgren 1898: 312 [?]. Ciocalypta oculata. – Thiele 1900: 75. Ciocalypta oculata maxima Hentschel, 1912: 428. Axinyssa fenestratus.–Van Soest, Díaz & Pomponi 1990: 27; Hooper & Bergquist 1992: 102. Ciocalypta confossa Hooper et al., 1997: 23. Ciocalypta fenestrata.– Hooper et al. 1997: 17. Ciocalypta oscitans Hooper et al., 1997: 20. Amorphinopsis foetida. – Hooper et al. 1997: 28. Material examined. Type maTerial – Leucophloeus fenestratus, HoloType, BMNH 1882.2.23.255, Darwin Harbour, N.T., 16–24 m depth, October 1881, HMS Alert. Leucophloeus fenestratus unnamed variety, BMNH B. Alvarez and J. N. A. Hooper C A B Fig. 2. Amorphinopsis excavans (NTM Z.5213): A, light microphotograph of tangential section of ectosomal skeleton, showing oxeas organised in bundles and small styles, paratangentially oriented in disorganised brushes; B, light microphotograph of section perpendicular to surface, showing choanosomal skeleton; C, diagram of spicules. Scale bars: A,C, 100 µm; B, 500 µm. 1882.2.23.195, Arafura Sea, 64–72 m depth, 18 October 1881, coll. HMS Alert. Suberites oculatus, syntype SMF 680, Ternate, Maluku Sea, coll. Kükenthal, W, 1894. Ciocalypta oculata maxima, SMF 971, Aru Is, between Meriri and Leer, 6–10 m depth, 31 March 1908, coll. Merton, H. Ciocalypta oscitans, HoloType, QM G303560, Bynoe Harbour, E Fish Reef, N.T., 12°24.1334’ S, 130°28.16’ E, 17 m depth, 6 October 1993, coll. CCNT Ocean Rescue 2000 Program, dredge. Ciocalypta confossa, NTM Z.3106, Parry Shoals, Arafura Sea, N.T., 11°12.5167’ S, 129°42.07’ E., 20 m depth, 15 August 1987, coll. Mussig, AM and NCI team. addiTional specimens – Melville Is: QM G313543. Shoal Bay, Vernon Is, Cape Hotham: QM G303558, G303677, G303541. Bynoe Harbour: NTM Z.5207 (0M9H2319-N), Z.5208 (0M9H2543-H). Darwin Harbour: QM G303287, G310170; NTM, Z.2018, Z.4085, Z.4122, Z.4123, Z.5215 (0M9H2137-P), Z.5206 (0M9H2192-Y). Cobourg Peninsula, NTM Z.1391. Gulf of Carpentaria: QM G314246, G314247, G315205. Description Shape (Fig. 1B). Massive to subspherical, with tapering, hollow, rudimentary, subconical or volcano-shaped istules, up to 18 mm long and 30 mm diameter; basal portion buried beneath sediment and istules protruding through substrate. 58 Halichondriidae from northern Australia A C B Fig. 3. Amorphinopsis fenestrata: A, light microphotograph of tangential section of ectosomal skeleton (NTM Z.5206), showing spicule tracts criss-crossing and forming a reticulation of polygonal meshes; B, light microphotograph of section perpendicular to surface (NTM Z.5215) showing choanosomal skeleton; C, diagram of spicules. Scale bars: A-B, 500 µm; C, 100 µm. oriented and cavernous at subectosomal region. In istules choanosomal tracts more compressed in central region (Fig. 3B), cavernous towards periphery. Single spicules, including smaller styles scattered through choanosome. Spicules (Fig. 3C, Table 1). Choanosomal styles and styloids (thicker in apical third and with basal ends narrower than the centre), slender, straight or slightly curved at centre, fusiform, in a great size range (163–895 x 3.3–17.4 µm). Smaller styles can be transitional to subtylostyles. Relative proportions of styles, styloids and subtylostyles vary among populations. Remarks. The species was originally described by Ridley (1884) under Leucophloeus, a junior synonym of Ciocalypta (Van Soest et al. 1990; Erpenbeck & Van Soest 2002) and related to species of Ciocalypta by Hooper et al. (1997) based on the characteristics of the ectosomal skeleton, and the presence of styles. As described above, the growth form of Amorphinopsis fenestrata is characterised by the presence of istule-like projections of several shapes Colour. Yellow, brown or pale mauve, alive. Oscula. Large, up to 10 mm in diameter or, grouped on a terminal sieve-plate, on apex of istules. Texture and consistency. Compressible, harsh, easily torn. Surface. Irregular, rugose, translucid, hispid, marked in some specimens with longitudinal channels. Skeleton. Ectosomal skeleton (Fig. 3A) detachable, supported by subectosomal multispicular tracts. Formed by multispicular tracts of larger choanosomal styles, up to 3 spicules abreast, lying tangential to surface, directionless or criss-crossing, forming a nearly regular reticulation of polygonal meshes; and, irregular bundles of smaller ectosomal styles arranged mostly paratangential to surface as plumose brushes or tufts. Choanosomal skeleton disorganised, halichondroid criss-cross of both unispicular and multispicular tracts, containing 5–20 spicules abreast, with larger choanosomal styles mainly conined to central region; becoming more wide-meshed, paratangentially 59 B. Alvarez and J. N. A. Hooper Table 1. Comparison of spicule dimensions among specimens of Amorphinopsis fenestrata. Specimen Locality Styles BMNH 1882.2.23.255 (Holotype) Darwin Harbour 193.6–611.8µm (390.4±136.4) x 5.3–12.7µm (8.6±2) BMNH 1882.2.23.195 Arafura Sea 252.9–895.3µm (617.7±217.3) (Ridley’s unnamed variety specimen) x 5.9–15.4µm (9.6±2.6) Ternate, Indonesia 210.2–761.6µm (431.1±160) SMF 680 (Syntype of Suberites oculatus) x 4.7–17.4µm (8.1±2.8) Aru Is, Indonesia 306.4–892.8µm (563.3±176) SMF 971 (Syntype of Ciocalypta oculata maxima) x 3.3–11.6µm (6.6±1.8) Parry Shoals 217.8–649µm (448.9±149.7) NTM Z.3106 (Holotype of Ciocalypta confossa) x 6.4–12.6µm (9.2±1.9) Bynoe Harbour 175.1–690.4µm (502.7±155.8) QM G303560 (Holotype of Ciocalypta oscitans) x 5.2–13.3µm (8.7±2.2) NTM Z.5215 Darwin Harbour 178.5–891.1µm (427.1±199.8) x 4–12.3µm (7.7±2.3) NTM Z.5207 Bynoe Harbour 163.5–833.1µm (432.4±214.6) x 3.7–13.7µm (7.3±2.3) QM G314247 Gulf of Carpentaria 218.8–599.3µm (412.2±120.3) x 4.3–11.1µm (7.2±2.2) (e.g. pointy, hollow, pyramidal or volcano-shaped), some with a system of exhalant channels. They project from a semi-buried massive base and protrude through the substrate/sand. The skeletal architecture in these istulose projections, although similar, is considered here not to be homologous to the one observed in Ciocalypta species, which is characterised by a central spicular axis and extraaxial secondary tracts supporting the ectosomal skeleton (see below under Ciocalypta). Instead, the skeleton of the istulose projections of this species is formed by multiple axes, oriented longitudinally and radiating towards the surface. Based on these arguments, we suggest that the species is better allocated to the genus Amorphinopsis. It needs to be noted however, that the skeleton of the present species is composed totally by styles with a great size range and lacks of ‘true’ oxeas, a diagnostic characteristic of this genus. In the majority of specimens examined, including the holotype, a great proportion of the styles are ‘subacerate’ (styloids) similar to those observed in Aaptos (Hadromerida: Suberitidae) where the thickest part of the spicule occurs in the apical third and the basal end is substantially narrow. The relative proportion of styloids varies among the populations examined. Subtylostyle modiications, especially in the smaller styles are also common. We are unable to conirm whether this type of styles might be considered a derived form of oxea (as stated in the current deinition of the genus) and therefore we propose to expand slightly the deinition of Amorphinopsis to accept species with spicules differentiated into oxeas and/or styles in a large size range, with smaller ones concentrated at the surface. The examination of additional specimens allowed us to understand the concept of this species better and to conclude that the material from Beagle Gulf described by Hooper et al. (1997) under Ciocalypta oscitans, C. confossa and Amorphinopsis foetida, and the Indonesian species Suberites oculatus Kieschnick and Ciocalypta oculata maxima Hentschel are all conspeciic with A. fenestrata. Amorphinopsis subacerata (Ridley & Dendy, 1886) from the Philippine Islands is very similar to A. fenestrata and as stated by those authors they share many characteristics including the type of styles but differ in external form and in having larger and thicker styles. Study of local populations and examination of the type material is required to establish whether the two species are conspeciic. Distribution. Amorphinopsis fenestrata is common throughout the Northern Territory coast (Sahul Province), from Parry Shoals (NW of Darwin Harbour) to the Gulf of Carpentaria. The records of Lindgren (1897, 1898) from Vietnam and China, of Dendy (1922) from the Indian Ocean and of Burton (1928) from the Malay Archipelago need to be veriied. Amorphinopsis maculosa (Pulitzer-Finali, 1996) comb. nov. (Figs 1C, 4) Topsentia maculosa Pulitzer-Finali, 1996: 114. Axinyssa aplysinoides. – Hooper et al. 1997: 4. Not Axinyssa aplysinoides Dendy, 1922: 39. Material examined. HoloType – MSNG 48701, Laing Is., 4º 09’ S, 144º 52’ E, Papua New Guinea, 6 m depth, 23 August 1986. addiTional specimens – Gulf of Carpentaria: QM G300854, G301034, G313577, G314255, G314267, G315207, G320819, G320904. Shoal Bay: QM G303561. Description Shape. Thickly encrusting, following substrate with convoluted ridges and short projections. Colour. Light grey or yellow alive; yellow inside beige in ethanol. Oscula. Inconspicuous, of different diameter, lushed and irregularly distributed or aggregated in top of the short projections (Fig. 1C). Surface. Smooth, lumpy, with dermal skin of reticulated appearance. Consistency and texture. Firm but crumbly. 60 Halichondriidae from northern Australia A C B Fig. 4. Amorphinopsis maculosa (QM G313577): A, light microphotograph of tangential section of ectosomal skeleton showing brushes of small styles with ends projecting through surface; B, light microphotograph of section perpendicular to surface showing choanosomal skeleton; C, diagram of spicules. Scale bars: A, 100 µm; B, 500 µm; C, 100 µm. Skeleton. Ectosomal skeleton (Fig. 4A) consisting of tangential to paratangential crust, approx. 200–300 µm thick, supported by choanosomal tracts of spicules and with disorganised brushes of small styles with ends projecting through surface, forming a discontinuous palisade spaced roughly at regular distances. Choanosomal skeleton (Fig. 4B) halichondroid, forming oval to round lacunae, 300–1000 µm in diameter, becoming compact towards surface, with very little collagen and abundant spicule content. Spicule tracts long and multispicular, running either towards surface or parallel to surface at the subectosomal area. Spicules (Fig. 4C, Table 2). Oxeas and much less frequent styles in a large size variation, 207–994 x 5–38 µm; small ectosomal styles (and transitional to oxeas), 139–274 x 3–8 µm. Remarks. The holotype of Topsentia maculosa Pulitzer-Finali was examined and is comparable in external morphology and skeletal characteristics to material collected in the Gulf of Carpentaria and Shoal Bay. The species is redescribed using this material and assigned to the genus Amorphinopsis. Table 2. Comparison of spicule dimensions among specimens of Amorphinopsis maculosa. Specimen MSNG 48701 (Holotype of Topsentia maculosa) QM G313577 Locality Papua New Guinea Gulf of Carpentaria QM G314255 Gulf of Carpentaria Oxeas 240.2–994.6µm (633.7±255.1) x 7.4–37.6µm (21.7±9.9) 369.1–863.5µm (675.8±124.6) x 7.9–20.3µm (15.2±3.2) 207.9–819.5µm (560.1±166.3) x 4.5–22.1µm (12.6±4) 61 Styles 138.5–259.2µm (211±35.5) [16] x 3–9.7µm (6.6±1.8) [16] 143.7–273.8µm (187.7±35.2) x 3.2–7µm (4.8±1) 147.5–240.4µm (174.2±20.9) x 3–7.8µm (5.6±1) B. Alvarez and J. N. A. Hooper Amorphinopsis maculosa is very similar in habit and skeletal organisation to A. fenestrata (Ridley, 1884) but that species has only styles as megascleres and lacks the oxeas of variable sizes present in this species (see above). Fistulous projections, as observed in A. fenestrata, are not present in the examined material of A. maculosa. The specimen assigned to Axinyssa aplysinoides (Dendy) by Hooper et. al. (1997) was re-examined and has characteristics (i.e. habit, ectosomal skeleton, oxeas and smaller styloids) that agree with Amorphinopsis maculosa. Amorphinopsis maculosa was also compared to other species of Amorphinopsis recorded for northern Australia and adjacent biogeographical regions – Amorphinopsis excavans (Dendy, 1889) from the Indian Ocean, A. foetida (Dendy, 1889) from the Gulf of Manaar, A. maza (De Laubenfels, 1954), A. oculata (Kieschnick,1896), and A. sacciformis from Indonesia. Amorphinopsis excavans has similar habit, skeletal structure and spicule composition, but differs in the organisation of the ectosomal skeleton; in A. excavans it consists of a tangential layer of thick intercrossing tracts of large oxeas, and loose oxeas of all sizes with small styles illing up the spaces (Erpenbeck & Van Soest 2002). The material of A. excavans from northern Australia described above also has a clearly tangential and detachable ectosomal skeleton of large oxeas grouped in bundles and small styles tangentially to paratangentially oriented, sometimes in disorganised tufts. Other characteristics that differentiate A. maculosa from A. excavans are the predominance of small styles in the ectosome (instead oxeas and styles as reported in most populations of A. excavans) and the lacunar appearance of the choanosomal skeleton. Amorphinopsis foetida (type specimen BMNH 1889.1.21.55, examined) is also a massive and slightly lobose species but with a skeletal architecture and spicule geometry different to A. maculosa. The ectosomal skeleton of A. foetida is formed by a halichondroid dermal crust, tangential-paratangetially oriented, 500–900 µm thick, with a mixture of oxeas and small styles projecting through the surface, and with vague tracts and small rounded meshes approx. 100–300 µm. The choanosomal skeleton is halichondroid, cavernous, with large rounded lacunae, 500–700 µm approx., and ill-formed multispicular tracts and ibres irregularly oriented through the skeleton; less compact than the ectosome and bounded by little collagen. The spicules are mixture of oxeas of different thickness and sizes, characteristically curved or sinuous (165.4–720.6 µm x 3.3–16.6 µm). The smaller styles are curved, and some are transitional to oxeas (154–281.7µm x 2.7–7.9µm). Amorphinopsis maza was re-examined by Erpenbeck & Van Soest (2002) and differs from A. maculosa in habit, skeletal organisation and spicule composition. Suberites oculatus Kieschnick, 1896, accepted as Amorphinopsis oculata (Van Soest et al. 2008) is a synonym of Amorphinopsis fenestrata (see above). Ciocalypta sacciformis Thiele, 1900 (syntypes, SMF 685, 1818, Ternate Maluku Sea, 1894, coll. Kükenthal W., examined) was transferred from its original genus to Ciocalapata by De Laubenfels (1936) and related it to species-like Ciocalypta including oxeas and styles. The species was interpreted as Amorphinopsis by Hooper et al. (1997). The revision of the type material indicates that the specimens described by Thiele agree better with the current concept of Halichondria but not with Ciocalypta, Ciocalapata or Amorphinopsis as suggested by previous authors. The syntypes examined are small fragments with a pouch-like shape as described by Thiele. The ectosomal skeleton is a thick tangential to paratangential layer up to 500–800 µm, with single spicules and short tracts crisscrossing in a disorganised manner. The choanosomal skeleton is disorganised with bundles of spicules irregularly spaced and running towards the surface where they merge with the ectosomal skeleton. The skeleton is formed entirely by oxeas in a great size range 195–706 µm. The skeleton of SMF 1818 includes styles in low frequency 463–525 µm (491.2±17.4) x 6–16 µm. As admitted by Thiele, the styles might be modiications of oxeas. Distribution. Papua New Guinea (Eastern Coral Triangle Province), Gulf of Carpentaria and outer region of Shoal Bay (Sahul Shelf province) between 6–28 m depth. It is also found in Torres Strait (Northeastern Australian Province) (Alvarez & Hooper unpublished data). Remarks on Amorphinopsis. Amorphinopsis includes approx. 13 valid species (Van Soest et al. 2008) distributed mainly throughout the Indian Ocean but with species also recorded from the Atlantic and Mediterranean oceans. The genus is represented in northern Australia (Sahul Shelf Province) by three species – Amorphinopsis excavans, A. fenestrata comb. nov. and A. maculosa comb. nov. Amorphinopsis foetida was recorded by Hooper et al. (1997) from the Beagle Gulf but it is concluded here that the specimen is better allocated to A. fenestrata. The species originally described as Leucophloeus fenestratus Ridley, 1884 is common in the study area. Its habitat and skeletal characteristics have been interpreted as typical from the genus Ciocalypta but as discussed above, these similarities are considered not homologous and we propose to include the species in Amorphinopsis. The choanosomal skeleton of A. fenestrata is formed entirely by styloids and lacks ‘true’ oxeas as seen in other species of Amorphinopsis and other genera of Halichondriidae. We propose to expand the deinition of Amorphinopsis slightly to include species with spicules differentiated into oxeas and/or styles. Genus Axinyssa Lendenfeld, 1897 Gender feminine. Type species, by original designation, Axinyssa topsenti Lendenfeld, 1897: 116. Recent, western Indian Ocean. 62 Axinyssa bergquistae (Hooper et al., 1997) com. nov. (Figs 1D, 5) Halichondria bergquistae Hooper et al., 1997: 45. Halichondriidae from northern Australia membrane; becoming distinctively conulose at erect columns or digits. Conules up to 2 mm long organised in longitudinal rows along erect columns and digits, with brushes of larger choanosomal spicules protruding through surface. Texture and consistency. Hispid due to projection of spicules through surface, irm and incompressible. Skeleton (Fig. 5A,B). Ectosome without specialised skeleton, with lightly coloured collagenous skin. Choanosomal skeleton halichondroid, with high spicule density in deeper regions; becoming more organised at subectosomal region, with multispicular spicule tracts, 50– 200 µm running longitudinally and ascending towards surface, becoming more radial and plumose near periphery; ending in disorganised brushes that project through ectosome. Spicules (Table 3, Fig. 5C). Mixture of oxeas of variable thickness and length (354–948 x 4–27.8 µm), slightly bent Material examined. HoloType – QM G303351, East Point Bommies, Darwin Harbour, Northern Territory, Australia, 12°24.083’ S, 130°48.14’ E, 10 m depth, 23 September 1993, coll. Hooper, JNA & Hobbs, LJ. addiTional specimens – Cartier Island: QM G301059. Bynoe Harbour: Z.5224 (0M9H2375-X); Z. 5901. Darwin Harbour: Z.5976. Gulf of Carpentaria: QM 313572. Description Shape (Fig 1D). Massive-lobate, bulbous-digitate; with erect columns or irregular coalescent plates. Individuals approx. 70–100 mm high, 45–200 mm thick. Colour. Purple-mauve, grey-brown; some individuals with lighter tinges. Oscula. Variable in size (3–7mm diameter), conspicuous, discrete, with raised white and opaque membranous lips (Fig. 1D), irregularly distributed. Surface. Smooth to microconulose at base with shallow and meandering channels covered by a translucent A C B Fig. 5. Axinyssa bergquistae: A, light microphotograph of perpendicular section (NTM Z. 5976) showing choanosomal spicule tracts projecting through surface; B, light microphotograph of perpendicular section through surface (NTM Z.5224), showing choanosomal skeleton with longitudinal plumose tracts; C, diagram of spicules. Scale bars: A-B, 500 µm; C, 100 µm. 63 B. Alvarez and J. N. A. Hooper Axinyssa mertoni (Hentschel, 1912) com. nov. (Figs 1E, 6) Ciocalypta mertoni Hentschel, 1912: 424; Burton 1934: 564. Halichondria tyleri. – Hooper & Wiedenmayer 1994: 209. Halichondria mertoni. – Hooper et al. 1997: 52. Pseudaxinyssa pitys De Laubenfels, 1954:178; Bergquist 1965 : 175 [?]. Axinyssa pitys. – Hooper & Bergquist 1992: 102. Material examined. Type maTerial – Ciocalypta mertoni, holotype, SMF 1608, Aru Is, North of Penambulai; station 10, 8 m depth, 2 April 1908, coll. Merton exp. 1908. Pseudaxinyssa pitys, holotype, USNM 23103, Caroline Islands, Palau Is, Koror I., Iwayama Bay, 2 m depth, 1 September 1949, coll. De Laubenfels, M.W. addiTional specimens –Darwin Harbour: NTM Z.5221 (0M9H2189-V). Description Shape (Fig. 1E, 6A). Massive with conspicuous istules up to 50 mm long, 1–2 mm thick, projecting from semiburied basal portion up to 100 mm diameter. Colour. Grey alive. Oscula. On top of istules, lushed, less than 5 mm diameter. Surface. Regularly conulose with transluscent membrane stretching over conules; marked with choanosomal axes. Skeleton (Fig. 6B). Ectosomal skeleton absent. Choanosomal skeleton plumose to halichondroid with multispicular axes of spicules running longitudinally, nearly parallel and close to each other, anastomosing and diverging towards surface and becoming dendritic, bounded with collagen and ending in disorganised plumose brushes that project through surface. Spongin ibres ill-deined and short, direction-less, embedding spicule tracts. Spicules (Fig. 6C, Table 4). Oxeas, hastate, slightly bent, straight or crooked, 476–1470.2 x 11.2–29.7 µm. Style and strongylote modiications also present. Remarks. Ciocalypta mertoni is conspecific with Pseudaxinyssa pitys and is better allocated to the genus and sometimes slightly sinuous. Styloid modiications common. Remarks. The species was initially described under Halichondria. Examination of additional material considered conspeciic with the specimen described by Hooper et al. (1997) allowed us to conclude that the species is better allocated in Axinyssa. The plumose conules observed in some areas of the surface of A. bergquistae resembles Axinyssa mertoni (described below), but that species has a more lax skeleton with less spicular density and more collagen. It differs also in general shape, colour and the oscula morphology. Axinyssa bergquistae is comparable to A. valida (Thiele, 1899: 12) [holotype NHMB 13, examined] in external morphology, skeletal organisation and size of spicules [(290.3–859.3 µm (599±201.8) x 9.7–34.8 µm (20.6±8.3)] and could possibly be conspeciic. But both the skeleton and the size of spicules of most Axinyssa species are very similar and the separation of species is dificult and subjective (see below under remarks on the genus). The examination of more material from Indonesia (Alvarez & De Voogd in progress) will help to determine whether the northern Australian populations belong to the same species. Distribution. Darwin Harbour and Bynoe Harbour (Sahul Shelf Province). Probably present also in the Northeast Australian Shelf Province (Alvarez & Hooper unpublished data). Table 3. Comparison of spicule dimensions among specimens of Axinyssa bergquistae. Specimen QM G303351 (Holotype of Halichondria bergquistae) NTM Z.5224 Locality Oxeas Darwin Harbour 407.3–793.8µm (619.7±82.8) x 10.3–27.8µm (15.5±4.3) QM G313572 Gulf of Carpentaria A Bynoe Harbour 366.3–948.1µm (749.2±135.7) x 4.5–33.2µm (21.3±7) 353.6–747.3µm (618.3±77.5) x 4–20.9µm (14.7±4.2) C B Fig. 6. Axinyssa mertoni: A, preserved specimen (NTM Z.5221); B, light microphotograph of perpendicular section through surface showing choanosomal skeleton with longitudinal tracts projecting through surface (Holotype, SMF 1608); C, diagram of spicules. Scale bars: A,B, 500 µm; C, 100 µm. 64 Halichondriidae from northern Australia Axinyssa. The type material of both species were examined here, and they are identical in all their characteristics. Ciocalypta mertoni was considered a junior synonym of Halichondria tyleri by Hooper & Wiedenmayer (1994) (following Burton 1959), but this synonym was later rejected by Hooper et al. (1997) who considered it a valid species of Halichondria, but admitted that the lack of an ectosomal skeleton was atypical of that genus. The species agrees well with the concept of Axinyssa, as it lacks an ectosomal skeleton and has a halichondroid to vaguely plumose choanosomal skeleton bounded by relatively high amounts of collagen and formed by oxeas of variable size and common styles and strongyles. The growth form of this species however, is not reported for other species of Axinyssa, thus a slight expansion in the diagnosis of the genus is necessary to accommodate species with istulose projections. A single specimen found in Darwin Harbour is assigned to this species. It differs from the type material examined only on spicule dimensions. The oxeas of the Darwin specimen are in average longer and thicker, but this might correspond to intraspeciic variation within the species. Distribution. Axinyssa mertoni is rare within the study area with only one specimen recorded here. The distribution of the species is extended to the Sahul Shelf and the Tropical Northwestern Paciic provinces. The record of Bergquist (1965) from Palau needs to be veriied as there are several sympatric species of Axinyssa occurring in the area (Lori Bell Colin pers. comm.). Remarks on Axynissa. Axinyssa is represented in the area of the Indo-Paciic by several species: A. aculeata Wilson, 1925 (Philippines); A. aplysinoides (Dendy, 1922) (Seychelles); A. oinops (De Laubenfels, 1954) (central West Paciic); A. topsenti Lendenfeld, 1897 (Tanzania); A. variabilis Lindgren, 1897 (Malaysia); and A. valida (Thiele, 1899) (Indonesia). The new combinations established in this revision extend the list by two additional species – A. mertoni (Sahul Shelf and Tropical Northwest Paciic provinces) and A. bergquistae (Sahul Shelf and probably Northeast Australian Shelf). Two additional species from the central Indo-Pacific region are also referred to Axinyssa after examination of type material – Ciocalypta rutila gracilis Hentschel, 1912 (SMF 1566, Aru Is, examined; see under remarks of Ciocalypta), and Phycopsis terpnis De Laubenfels, 1954 (Caroline Is, Central Paciic, USNM 23061, examined). We were not able to revise all the type material of the Indo-Paciic species of Axinyssa thoroughly, therefore it remains inconclusive whether A. mertoni and A. bergquistae might be conspeciic with other species recorded from the region. As mentioned above, the skeletal organisation and the size of spicules among Axinyssa species is very similar and separation of species is subjective. The external morphological characteristics seem to be more discrete, but study of individual populations is necessary to determine the actual range of variability present within these species. Table 4. Comparison of spicule dimensions among specimens of Axinyssa mertoni. Specimen SMF 1608 Locality Aru Island, Indonesia Caroline Is, Central West Paciic USNM 23103 (holotype of A. pitys) NTM Z.5221 Darwin Harbour, NT Oxeas 515.6–779.6µm (691.4±61) x 11.2–21.7µm (18±2.5) 514.3–871µm (788.2±74.9) x 8.6–18.5µm (15±2.5) 476–1470.2µm (979.5±285.9) x 14.7–29.7µm (21.6±4.8) Study of different populations of Axinyssa through Indonesia (Alvarez & De Voogd unpublished data) is currently in progress and will help to re-deine the limits of Axinyssa species. The diagnosis of Axinyssa is here expanded to include species like Axinyssa mertoni with istulose projections. We note however, that the skeletal organisation of the istules observed in A. mertoni is considered not to be homologous with the organisation observed in species of Ciocalypta. The istulose projections of Ciocalypta spp. are transparent with the skeleton formed by a central axis of spicules and extraaxial tracts diverging towards the surface. The istulose projections of A. mertoni are opaque, tough-cartilagineous, arising from a massive base, and without a central axis of spicules which is diagnostic for Ciocalypta. Axinyssa mertoni shares with the Indian Ocean Ciocalypta digitata (Dendy, 1905) the lack of an ectosomal skeleton, and the presence of istulose projections; however, the istules of C. digitata are transparent with a spicular axis from which thick bundles of spicules diverge towards the surface ending in conules (Erpenbeck & Van Soest, 2002). In our view the expansion of the deinition of Axinyssa to include species with istular projections such as A. mertoni does not affect the current position of C. digitata or the deinition of the Ciocalypta. The current position of the genus Axinyssa within the family Halichondriidae is debatable as molecular data (Erpenbeck et al. 2005) indicate that some species currently allocated to this genus are related to other dictyonellid genera such as Acanthella and Dictyonella. From a morphological point of view, the lack of an ectosomal skeleton, the presence of abundant collagen in the skeleton and the common occurrence of styles, strongyles and transitional forms, also points to possible relationships with members of Dictyonellidae. These relationships should be further explored to conirm the placement of this genus within the family Halichondriidae. However, taxonomic veriication of the species of Axinyssa used in the molecular analyses should also be taken in consideration, particularly given the paucity of morphometric characters in this group and our still rudimentary understanding of character homology. 65 Genus Ciocalypta Bowerbank, 1862. Gender feminine. Type species, by monotypy, Ciocalypta penicillus Bowerbank, 1862. Recent, eastern Atlantic Ocean. B. Alvarez and J. N. A. Hooper Ciocalypta heterostyla Hentschel, 1912 (Figs 1F, 7) Ciocalypta heterostyla Hentschel, 1912: 424; Hooper et al. 1997: 36. Material examined. HoloType – SMF 1569, Aru Is, N Penanbuli, 8 m depth, 2 April 1908, coll. H. Merton. addiTional specimens – Darwin Harbour, NTM Z.5902. Description Shape (Fig. 1F). Fistulose, with semi-buried and massive base. Fistules pointed, projecting perpendicularly from base, 20–30 mm long, less than 10 mm diameter wide, slightly translucent especially at tips. Colour. Light yellow. Oscula. Apical on istules. Surface. Smooth, microconulose. Ectosomal skeleton (Fig. 7A). Thin layer formed by tangentially oriented pauci- to multispicular tracts of spicules, crossing over and forming a reticulation of triangular meshes; supported by choanosomal tracts. Choanosomal skeleton. Differentiated at fistules, with central column formed by thick multispicular tracts oriented longitudinally and expanding into thick brush at tip of istule (Fig. 7B). Secondary tracts, 20–100 µm thick, slightly plumose, diverging nearly perpendicularly from central column toward surface, regularly spaced and connected irregularly by unispicular-paucispicular tracts of spicules; becoming thicker and ending in fan-like brushes at the subectosomal area, generally with smaller spicules oriented with their ends towards surface. Skeleton at base C A B Fig. 7. Ciocalypta heterostyla (Holotype SMF 1569): A, light microphotograph of tangential section of ectosomal skeleton, showing tracts of spicules, forming a reticulation of triangular meshes; B, light microphotograph showing choanosomal skeleton at a istule. C, diagram of spicules. Scale bars: A-B, 500 µm; C, 100 µm. 66 Halichondriidae from northern Australia Ciocalypta stalagmites Hentschel, 1912 (Figs 1G–H, 8) Ciocalypta stalagmites Hentschel, 1912: 426. Halichondria tyleri.– Hooper et al. 1997. Material examined. lecToType – SMF 1567 (here designated, a representative specimen of the species, chosen from six examined specimens deposited at SMF and registered as syntypes with a single accession number), Aru Is, Mimien I., Indonesia, 15 m depth, 9 April 1908, coll. Meron, H. paralecToTypes – SMF 1537, Aru Is, Sungi Manumbai (Kapala Sungi), Indonesia, station 17, Merton Expedition Aru and Kei Is 1907-1908, 20 m depth, 5 May 1908, coll. Merton, H. SMF1574, Aru I., SW Lola I, Indonesia, station 9, Merton Expedition Aru and Kei Is 1907-1908, 8–10 m depth, 1 April 1908, coll. Merton, H. SMF 1595, Aru Is, Bambu I., Indonesia, station 11, Merton Expedition Aru and Kei Is 1907-1908, 10 m depth, 3 April 1908, coll. Merton, H. SMF 1623, Aru Is, SW Mariri and Leer Is, Indonesia, Merton Expedition Aru and Kei Is 1907-1908, 6–10 m depth, 31 March 1908, coll. Merton, H. SMF 1627, Aru Is, N Penambulai I., Indonesia, Merton Expedition Aru and Kei Is 1907-1908 , 8 m depth, 2 April 1908, coll. Merton, H. addiTional specimens – Parry Shoals, NT, NTM Z.3133. Charles Point, NT, NTM Z.5230 (0M9H2574-P). Bynoe Harbour: NTM Z.5226 (0M9H2511-V). Darwin Harbour: NTM Z.941, Z.5210 (0M9H2571-M), Z.5229 (0M9H2299-Q), Z.5977. Cobourg Peninsula: NTM Z.592, Z.1358, Z.1395, Z.3286. Wessel Is: Z.3920, Z.5218 (0M9H2666-P). Description Shape (Figs 1G–H, 8A). Flat cushion-shaped or massive base, buried or semi-buried, strongly attached to substrate, up to 35 mm thick, 4–100 mm diameter, with istules projecting perpendicularly above surface. Fistules halichondroid, with directionless multispicular tracts and single spicules in confused reticulation. Spicules (Fig. 7C, Table 5). Mixture of styles in a large range of sizes, straight to slightly curved, some sinuous, some with rounded ends, 184–809 x 4–19 µm. Remarks. Ciocalypta heterostyla was originally described from the Aru Is (Indonesia) and it has not been redescribed or recorded until this present study. The material from Darwin Harbour agrees closely with the type. Only the spicule dimensions are on average slightly larger in the specimen from Darwin. The species is not very conspicuous due to its cryptic habit. It is apparently rare in the study area with only one specimen collected so far. Hooper &Wiedenmayer (1994) followed Burton (1959) and considered this species a synonym of Ciocalypta tyleri Bowerbank. The two species are certainly similar in external morphology and skeletal organisation, however the skeleton of C. heterostyla is formed exclusively by styles instead of oxeas, and the skeleton within the istules is much more organised with a regular reticulation. Because of these characters we considered that C. heterostyla is not only a valid species but also it can be easily differentiated from its South African relative. Distribution. Known only from the type locality (Aru Is, Indonesia) and from Darwin Harbour (Sahul Shelf province). It is found between 8–12 m depth. Table 5. Comparison of spicule dimensions between specimens of Ciocalypta heterostyla. Specimen Locality SMF Aru Is, Indonesia 1569 Darwin Harbour NTM Z.5902 Styles 199.9–584.1µm (387.3±137) x 3.5–14.8µm (8.3±3.2) 184–809.2µm (466.5±229.1) x 3.9–18.6µm (9.4±5.5) A B D C Fig. 8. Ciocalypta stalagmites (Lectotype, SMF 1567): A, Preserved lectotype; B, light microphotograph showing choanosomal skeleton at istule; C, light microphotograph showing choanosomal skeleton at base; D, diagram of spicules. Scale bars: A, 2 cm; B-C, 500 µm; D, 100 µm. 67 B. Alvarez and J. N. A. Hooper diagnostic to separate C. stalagmites from other Ciocalypta species. Ciocalypta stalagmites is very similar to C. tyleri Bowerbank, 1873 from South Africa, and it was interpreted as such by Hooper et al. (1997) but allocated to the genus Halichondria (following Van Soest et al. 1990; Van Soest 1991). Both species are now referred to Ciocalypta under the revised concept of Halichondriidae (Erpenbeck & Van Soest 2002). The two species are similar in external morphology and skeletal organisation, however, the skeleton of C. stalagmites seems to be less organised. Despite similarities between these two species, it is unlikely that the Indonesian and northern Australian populations are conspeciic with their South African relatives, thus we propose to reserve C. stalagmites for those populations inhabiting the Sahul Province and adjacent areas. Future independent evidence might demonstrate whether C. tyleri and C. stalagmites belong to a complex of cryptic species that cannot be easily separated using traditional morphological characters, or are instead conspeciic and genuinely a widely distributed species. Ciocalypta stalagmites is also very similar to C. vansoesti (Hooper et al., 1997). comb. nov. The two species have similar growth form and skeletal characteristics, but they can be differentiated from each other by some distinctive characteristics (see below). Distribution. Ciocalypta stalagmites is very common throughout the northern Australian localities of the Sahul Shelf province and its distribution extends to adjacent provinces including the Northeast Australian Shelf, Papua New Guinea (Alvarez & Hooper unpublished), and Indonesia (Alvarez & De Voogd unpublished). It is found from the intertidal zone to 40 m. sharply pointed, globular or lattened, mammiform, tubular, rounded, 5–125 mm long, laterally fused sometimes or branching at tips. Colour. Base generally pale-beige. Fistules, transparent, mauve-brown-purple, greenish-yellow. Internally, beige. Oscula. Generally at apex of fistules or tubes but also observed irregularly distributed along istules, with membranous rims. Consistency and Texture. Compressible-spongy, easily torn. Surface. At istules, smooth to slightly conulose, marked with longitudinal rows of minute conules. Choanosomal tracts of spicules, visible through the ectosome of translucent specimens. At base smooth, opaque, rough, spiculous. Skeleton. Ectosomal skeleton, tangential layer of variable thickness (5–300 µm) easily peeled in some specimens, formed by a dense mass of smaller oxeas and supported by choanosomal skeleton. Choanosomal skeleton (Fig. 8C) at base, halichondroid, densely spiculous, with mixture of large and smaller oxeas and no distinct tracts of spicules. In istules (Fig. 8B), becoming distinctive and organised, with central column of large spicules oriented longitudinally and smaller spicules criss-crossing. Multispicular tracts generally formed by medium-size spicules radiating from central column towards surface, connected by shorter tracts and single spicules, forming a regular reticulation. Large subectosomal spaces (up to 5 mm diameter) in some areas. Spicules (Fig. 8D, Table 6). Two size classes of oxeas: I, smaller, thinner, fusiform, straight or slightly lexuous (147–321 x 4–10 µm); II, large, thick and slightly curved, fusiform, occasionally crooked-sinuous (378–886 x11–40 µm), and styles in equivalent size categories are common. Remarks. The growth form of Ciocalypta stalagmites is remarkably variable. Fortunately this variation is also well represented in the type material. The external colouration is also variable and is possibly related to the presence of different cyanobacterial associations. The skeletal organisation, composition and size of spicules, however, are consistent and very similar among specimens with different habits and colouration. A very consistent characteristic through all the populations we examined is that the oxeas are differentiated in size categories not only by their length but also by their thickness. Proper statistic and morphometric analyses could be employed to demonstrate whether this is Ciocalypta vansoesti (Hooper et al., 1997) comb. nov. (Figs 1I, 9) Halichondria vansoesti Hooper et al., 1997: 37. Material examined. As listed by Hooper et al. (1997). addiTional specimens – Darwin Harbour: Z.5904, Z.5217 (0M9H2568-J), Z.5903, Z.5978. Bynoe Harbour: Z.5212 (0M9H2498-I).Gulf of Carpentaria: QM G303524. Remarks. This species was well described by Hooper et al. (1997), but allocated to the genus Halichondria (following Van Soest et al. 1990; Van Soest 1991). It is Table 6. Comparison of spicule dimensions between specimens of Ciocalypta stalagmites. Specimen SMF 1567 (Lectotype) Locality Aru Is, Indonesia NTM Z.5212 Bynoe Harbour NTM Z.5210 Darwin Harbour NTM Z.5218 Wessel Is Oxea type I 167.8–240.2µm (200.8±17.4) x 4.4–10µm (7.9±1.3) 165.3–320.6µm (238.2±34) x 5.5–10.3µm (7.6±1.2) 188.3–258.7µm (214±19.1) x 5.6–9.2µm (7.4±0.9) 147.2–202.7µm (169.1±12.8) x 5.3–9.4µm (6.7±0.9) 68 Oxea type II 377.7–837µm (626.2±102.8) x 12.3–37.1µm (18.7±4.7) 411.4–659.5µm (524.9±56.8) x 11.1–26.3µm (18.9±4.1) 440.1–662.8µm (601.8±53.9) x 11.3–38.5µm (21.9±7.7) 437.7–886.5µm (588.1±135.1) x 8.7–39.5µm (21.7±7.9) Halichondriidae from northern Australia A B Fig. 9. Ciocalypta vansoesti: A, preserved paratype specimen (QM G303450); B, light microphotograph showing choanosomal skeleton at istule. Scale bars: A, 20 mm; B, 500 µm. now referred to Ciocalypta under the revised concept of Halichondriidae (Erpenbeck & Van Soest 2002). As mentioned by Hooper et al. (1997) and above, Ciocalypta vansoesti is closely related to C. stalagmites, both with similar growth form, skeletal organisation and type of spicules. Both species have istules projecting from a buried-semiburied mass, but in the case of C. vansoesti the istules are transluscent-white with the surface regularly conulose and subectosomal tracts and central column visible beneath (Figs 1I, 9A). However, the skeleton in the istules, when compared to C. stalagmites, is denser and slightly disorganised; the central column is not as condensed and the extra axial reticulation is vague (Fig. 9B, Hooper et al. 1997, igs. 22a, 23a). The dimensions of the oxeas are similar and overlap with those of C. stalagmites, but in C. vansoesti there is not a clear difference in size categories, with the two classes of spicules overlapping both in length and in width (Table 7). Ciocalypta vansoesti is also similar in some of the ield characteristics to Axinyssa mertoni, indicating once again that istule-like growth forms are common among halichondrids and so are not useful to differentiate species unless they are used in combination with other skeletal characters. Distribution. Ciocalypta vansoesti is common in Darwin Harbour and Bynoe Harbour. It is also recorded from Cobourg Peninsula and the Gulf of Carpentaria. It is found intertidally and subtidally to 40 m depth. Remarks on Ciocalypta. Burton (1959) considered many species of halichondriid genera with a fistulose habit conspeciic with Ciocalypta penicillus (type species of Ciocalypta). These synonyms were not properly substantiated and some of them have been rejected by Hooper et al. (1997) and the present revision, i.e. Ciocalypta heterostyla, C. stalagmites, C. tyleri, C. oculata maxima (referred here to Amorphinopsis fenestrata), and C. mertoni (referred here to Axinyssa mertoni). Additional species of Ciocalypta recorded from the Sahul Shelf Province and adjacent areas besides the three revised here (i.e. C. heterostyla, C. stalagmites and C. vansoesti) are C. rutila gracilis Hentschel, 1912 (see below), C. digitata (Dendy, 1905, as Collocalypta), C. melichlora Sollas, 1902, C. rutila Sollas, 1902, and C. simplex Thiele, 1900: 76. One of the syntypes of Ciocalypta rutila gracilis Hentschel, 1912 (SMF 1566, examined) belongs in Axinyssa. Both the external morphology (based on Hentschel’s description) and the arrangement of skeleton agree with the concept of that genus. The skeleton of the material examined is formed by two classes of oxeas: straight 480.8–706.4 µm (601.6±76.7)x 9–24 µm (17±3.1) and vermicular, crooked sinuous, relatively thinner, and often bent up to a 90 degree angle, 193–600.7 µm (406.7±110.6) x 5.3–16 µm (10.6±3) [18]; less often styles are also present. Specimens recorded for Northeast Australian Shelf and Indonesia correspond with the type examined (Alvarez & Hooper unpublished data; Alvarez & De Voogd unpublished data) and will be redescribed under the name of Axinyssa gracilis in forthcoming publications. Ciocalypta digitata (Dendy, 1905) resembles Ciocalypta stalagmites and C. vansoesti in habit but differs in skeletal Table 7. Comparison of spicule dimensions between specimens of Ciocalypta vansoesti. Specimen NTM Z.2648 QM G303450 Locality Darwin Harbour, East Point Bynoe Harbour QM G303524 Gulf of Carpentaria Oxea type I 185.3–459.7µm (320±60.9) x 6.4–9.5µm (8±0.9) 194.1–361.7µm (275.3±45.6) x 2.1–9.8µm (6.8±1.8) 237.5–429.2µm (314.9±56.5) x 6.4–11.9µm (8.4±1.3) 69 Oxea type II 475.2–616.4µm (546.2±38.3) x 14–21.6µm (17.9±2.3) 382–662.1µm (536.1±78.2) x 6.8–30.8µm (16.8±6) 432.8–677.3µm (541.2±58.7) x 8.2–31.9µm (17.5±4.8) B. Alvarez and J. N. A. Hooper subectosomal region, with a vague reticulation of ascending tracts connected by short bundles and single spicules. Near surface, spicule tracts become more deined and even slightly plumose, supporting ectosomal skeleton and occasionally protruding through surface (Fig. 11B). Skeleton of some specimens is obscured by granular pigments and cyanobacteria when examined under light microscope. Spicules (Fig. 11C, Table 8). Mixture of oxeas not separable into size categories 112–389 x 2–12 µm, hastate, pointed, straight or occasionally slightly bent in middle. Remarks. Halichondria carotenoidea is diagnosed by a unique combination of features (i.e. growth form, organisation of the skeleton, and the size and composition of spicules) not found in other species of Halichondria revised here or reported for adjacent areas (see Table 9). The skeleton and spicule composition of all the material examined is almost identical to Halichondria (H.) microbiana sp. nov. (see below). Both species are sympatric and include a high density of cyanobacteria in the choanosome. The two species clearly differ in shape, which is branching to arborescent in H.(H.) carotenoidea and massive to cushion shape in H. (H.) microbiana. Both species can be further distinguished by the size of the oxeas, which are separated into class categories in H.(H.) microbiana but not in H.(H.) carotenoidea. The material described by Hooper et al. (1997) as Halichondria stalagmites (Hentschel) is a composite of H. (H.) carotenoidea (specimens NTM Z.2651 and Z.3147) and H (H.) microbiana (specimens nTm Z. 131, Z.1097, Z. 1991), and it does not agree with the concept of the species described by Hentschel (1912) as Ciocalypta stalagmites (see above). Distribution. Halichondria carotenoidea is common in Darwin and Bynoe Harbours, and it was also observed at Wessel Is (Raragala I.). Etymology. Named after carotenoid, a pigment which might give the red-orange colour to the species. It is intended as a noun in apposition. organisation and composition and size of spicules. This species lacks an ectosomal skeleton, but the fistular processes have spicular axis and extra-axial tracts as in other species of Ciocalypta. Genus Halichondria Fleming, 1828 Gender feminine. Type species, by subsequent designation (of Bowerbank 1862), Spongia panicea Pallas, 1766. Recent, East England. Subgenus Halichondria (Halichondria) Fleming, 1828 Halichondria (Halichondria) carotenoidea sp. nov. (Figs 10A,B, 11) Halichondria stalagmites. – Hooper et al. 1997: 43 [in part, not Z.131, Z.1097, Z.1991 = Halichondria (Halichondria) microbiana sp. nov.] Phakellia sp. 614. – Hooper et al. 1992. Axinella sp. 244. – Hooper et al. 1992. Material examined. HoloType – nTm Z.5909 , off Dudley Point, Fannie Bay, Darwin Harbour, N.T., 12°24.96’ S, 130°48.83’ E, 4–7 m depth, 4 June 2002, coll. Alvarez, B and party. paraTypes.– NTM Z.4451 (0M9H2036-G), Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, N.T., 12°29.103’ S, 130°47.111’ E, 8–14 m depth, 7 May 2002, coll. Alvarez, B and party. NTM Z.5225 (0M9H2462-S), Spencer Point, Indian I., Bynoe Harbour, N.T., 12°35.351’ S, 130°31.454’ E, 6–8 m depth, 11 June 2003, coll. Alvarez, B and party. addiTional specimens – N Adele I., Collier Bay, NW Shelf: Z.712. Parry Shoals, NT: NTM Z.3147. Darwin Harbour NT: NTM Z.194, Z.919, Z.934, Z.986, Z.1979, Z.2086, Z.2245, Z.2651, Z.2697. Description Shape (Figs 10 A,B). Fan shaped, digito-palmate, multiplanar, arborescent, erect, stalked, bifurcated at base, with complex branching; branches or digits flattened generally with pointed tips but round tips also common; specimens reaching up to 350 mm high, 300 mm wide. Colour. Orange. Blue-grey, or beige in alcohol Consistency and texture. Soft, easily torn, rubbery. Oscula. Regularly distributed along lateral side of branches (Fig 10B) less than 5 mm in diameter, generally in rows, with membranous rims slightly elevated. Some specimens with subectosomal thin canals ending in oscula. Surface. Smooth, wrinkled, evenly covered with microconules when exposed to air. Marked with subectosomal drainage canals in some specimens. Skeleton (Fig 11 A, B). Ectosomal skeleton formed by 100–200 µm thick continuous layer, supported by choanosomal skeleton and packed mainly with smaller category of oxeas, in short, ill-deined and criss-crossing bundles, oriented tangentially- paratangentially, protruding occasionally through surface in disorganised manner (Fig. 11A). Choanosomal skeleton, halichondroid to plumose, formed by ill-deined bundles of larger spicules, criss-crossing in all directions in the inner region of choanosome. Skeleton becomes more organised at Table 8. Comparison of spicule dimensions between specimens of Halichondria (Halichondria) carotenoidea. Specimen NTM Z.5909 NTM Z.5225 70 Locality Darwin Harbour, Dudley Point Bynoe Harbour Oxeas 113.2–388.9µm (263.2±87.1) x 2.3–12.3µm (7.3±3) 111.7–379.5µm (245.3±80.2) x 2.3–10.7µm (6.2±2.4) Halichondria (Halichondria) darwinensis Hooper et al., 1997 (Fig. 10C) Halichondria darwinensis Hooper et al., 1997: 49. Material examined. Specimens as listed in Hooper et al. (1997). addiTional specimens – Darwin Harbour, NTM Z.5211. Remarks. Halichondria darwinensis was described by Hooper et al. (1997). Additional material, an in situ Halichondriidae from northern Australia A B C D E F G Fig. 10. Halichondria (Halichondria) carotenenoidea sp. nov.: A, specimen in situ at Dudley Point, Darwin Harbour, N.T.; B¸ specimen in situ at Spencer Point, Bynoe Harbour, N.T., showing oscula distributed along lateral side of branches. C, Halichondria (H.) darwinensis, specimen in situ at Weed Reef, Darwin Harbour, N.T. Halichondria (H.) phakellioides: D, specimen in situ exposed at the reef lat of East Arm, Darwin Harbour during the low tide of 18 October 2001; E, specimen in situ at Raragala I. Wessel Is, 20 m depth. F, Halichondria (H.) microbiana sp. nov, specimen in situ at Lee Point, Darwin Harbour, N.T. G, Topsentia dura, specimen in situ at Nightcliff bommies, Darwin Harbour, N.T. Photographs: A,D,G – B. Alvarez; B – M. Browne; C,F – H. Nguyen; E – P. Colin. 71 B. Alvarez and J. N. A. Hooper Table 9. Halichondria (Halichondria) species from the Central Indo-Paciic (after Van Soest et al. 2008 ), with remarks on current taxonomic allocation, type material and description. Species Remarks Halichondria (Halichondria) bergquistae Hooper, Cook, Hobbs & Kennedy, 1997 Referred here to Axinyssa (see Remarks of Axinyssa this revision). Halichondria (Halichondria) darwinensis Hooper, Cook, Hobbs & Kennedy, 1997 Valid (see under genus Halichondria, this revision). Unrecognisable from the description; no type material available. Halichondria (Halichondria) armata Lindgren, 1897 Halichondria (Halichondria) cartilaginea (Esper, 1794) Valid, common species through Indo-Paciic. Always associated to symbiotic green algae (Lim et al. 2008). Halichondria (Halichondria) fragilis Kieschnick, 1896 Unrecognisable from the description; no type material available. Halichondria (Halichondria) pelliculata Ridley & Dendy, 1886 Valid, recognisable from the description. Type material not examined. Distinctive shape and surface characteristics, not comparable to the new species recorded here. Halichondria (Halichondria) stalagmites (Hentschel, 1912) Referred here to Ciocalypta (see remarks of Cioclaypta this revision) Halichondria (Halichondria) incrustans Kieschnick, 1896 Unrecognisable from the description; no type material available. Halichondria (Halichondria) ridleyi Hooper, Cook, Hobbs & Kennedy, 1997 Referred in this revision to Topsentia (see Remarks of Topsentia this revision) Halichondria (Halichondria) syringea Pulitzer-Finali, 1996 Halichondria (Halichondria) vansoesti Hooper, Cook, Hobbs & Kennedy, 1997 Valid. Type material examined (MSNG 48700). Coalescent istules on a buried base [?]. Detachable ectosome differentiated into a thick layer, with spicules oriented paratangentially, disorganised. Choanosome halichondroid with some ill deined, multispicular, ascending to surface and supporting ectosome. Oxeas several sizes, 256.4-706.7µm (539.3±137) x 5.2-19µm (13±3.7); few styles. Refered here to Ciocalypta (see under genus Ciocalypta, this revision) photograph (Fig 10C), and spicule measurements (Table 10) are provided here to complement the original description. Halichondria darwinensis is very inconspicuous and represented so far by only three individuals (including the holotype and the paratype). Its encrusting habit with small and insubstantial digits makes it inconspicuous and hard to ind. The re-examination of the available material and additional measurements of spicules, indicate that the differences in the thickness of the spicules reported by Hooper et al. (1997) as a distinctive character for the species, no longer appear to be signiicant (see Table 10). Both the length and the thickness of the oxeas are variable, but not divisible into size classes. The species seems to have afinities with the genus Axinyssa. The skeleton is poorly developed, with very little spongin and relatively less spicule density when compared to other Halichondria species; the ectosomal skeleton is quite undifferentiated and the spicules are similar in shape and dimensions to other Axinyssa species. Therefore, the assignment of this species to the genus Halichondria is inconclusive. Distribution. Halichondria darwinensis is presently known only from Darwin Harbour. It occurs intertidally and subtidally to 10 m. Table 10. Comparison of spicule dimensions among specimens of Halichondria (Halichondria) darwinensis. Specimen NTM Z.3205 (Holotype) Locality Darwin Harbour, East Point QM G303252 Darwin Harbour, East Arm (Paratype) NTM Z.5211 Darwin Harbour, Weed Reef Oxeas 272.3–659.6µm (467.2±112) x 3.7–13.2µm (8.5±2.5) 307–627.5µm (538.8±90.6) x 4.3–16µm (10.2±3.2) 386.2–663.7µm (524±68) x 7–14.9µm (11.8±1.9) 72 Halichondria (Halichondria) phakellioides Dendy & Frederick, 1924 (Figs 10 D, E, 12) Halichondria phakellioides Dendy & Frederick, 1924: 498; Burton 1934: 600; Hooper et al. 1997. Material examined. Specimens as listed in Hooper et al. (1997). addiTional specimens – Bynoe Harbour, NT: Z.241, Z.5223 (0M9H2321-P). Darwin Harbour, NT: Z.2026, Z.4100, (0CDN-8022-F), Z.5219 (0M9H2057-C), Z.5905, Z.5906, Z.5915, Z.5923, Z.5925, Z.5928, Z.5948, Z.5949, Z.5950, Z.5965, Z.5970. Parry Shoals, N.T., QM G310137. Boucat Bay, E of Maningrida, Arnhem Land, N.T.: Z.5623. Wessel Is: Z.5228 (0M9H2655-C). Remarks. Halichondria phakellioides was described by Hooper et al. (1997). Additional material, illustrations (Figs 10D, E, 12), and spicule measurements (Table 11) are provided here to complement the original description. Halichondriidae from northern Australia A C B Fig. 11. Halichondria (Halichondria) carotenenoidea sp. nov. (NTM Z.5909): A, light microphotograph of tangential section of ectosomal skeleton showing ill-deined and criss-crossing bundles of spicules; B, light microphotograph of perpendicular section through surface showing organisation of choanosomal skeleton and part of ectosomal skeleton layer (left upper corner of section); C, diagram of spicules Scale bars: A, 100 µm; B, 500 µm; C, 50 µm. depth, 7 August 2003, coll. Nguyen, H. paraTypes.– nTm Z.1097, Dudley Point Reef, East Point, Darwin, N.T., 12°25.0001’ S, 130°48.01’ E, 6–7 m depth, 22 December 1982, coll. Hooper, JNA. NTM Z.5908, Moira Reef, Bynoe Harbour, N.T., 12°30.799’ S, 130°30.527’ E, 5–8 m depth, 25 June 2003, coll. Nguyen, H. addiTional specimens.– Darwin Harbour NT: NTM, Z.1991. Cobourg Peninsula, N.T.: NTM Z.131. Description Shape (Fig.10F). Cushion-shaped with globular, subspherical or massive base, sometimes semi-buried Distribution. Halichondria phakellioides is widely distributed through the Northwest Australian Shelf and Sahul Shelf provinces. It is found from the intertidal to 20 m depth. Halichondria (Halichondria) microbiana sp. nov. (Figs 10F, 13) Halichondria stalagmites.–Hooper et al. 1997: 43 [in part; not Z.2651 and Z.3147 = H. (H.) carotenoidea]. Material examined. HoloType. –NTM Z.5907, Lee Point, Darwin, N.T., 12°20.538’ S, 130°52.184’ E, 9–12 m Table 11. Comparison of spicule dimensions among specimens of Halichondria (Halichondria) phakellioides. Specimen NTM Z.5223 Locality Bynoe Harbour NTM Z.5219 Darwin Harbour NTM Z.5228 Wessel Is Oxea type I 170.1–308.4µm (232±30.7) x 4.1–10.3µm (8±1.5) 154.2–293.2µm (228.6±34.5) x 5–10.3µm (7.3±1.4) 162.1–313.9µm (242.4±48.6) x 3.7–7.5µm (5.6±0.9) 73 Oxea type II 363.2–524.6µm (440.6±40.8) x 11.5–24.4µm (17.9±3.7) 356.7–499.6µm (425.5±38.7) x 10.5–20.2µm (16.3±3) 326.5–519.3µm (440.5±45.5) x 10.6–21.1µm (15.6±3) B. Alvarez and J. N. A. Hooper A B Fig. 12. Halichondria (Halichondria) phakellioides (NTM Z.5228): A, light microphotograph of tangential section showing organisation of ectosomal skeleton; B, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton. Scale bars: A-B, 500 µm. Paciic (Table 7) and those reported in this revision. It is diagnosed by a unique combination of features (i.e growth form, organisation of the skeleton, size and composition of spicules and presence of cyanobacteria in the ectosome and the choanosome) not found in those species. All the specimens examined here have a high density of ilamentous cyanobacteria in the choanosome, generally concentrated at the subectosomal region, a character shared with H. (H.) carotenoidea. As mentioned above, this new species is also very similar in skeletal characteristics to H.(H.) carotenoidea sp. nov., but it differs in habit and size of spicules, which are larger and separated in size categories in H. (H.) microbiana. Hooper et al. (1997) interpreted this species as Halichondria stalagmites. However, a thorough re-examination of the type material of that species indicates that it is not conspeciic with H. (H.) microbiana (see description of C. stalagmites above). Distribution. Halichondria microbiana is relatively common at Darwin and Bynoe Harbours, and it was also observed at Wessel Is (Raragala I.) and recorded from the Northeastern Australian Shelf (Alvarez and Hooper, unpublished data). It occurs between 6–12 m depth. Etymology. Named after the symbiotic microorganisms hosted by the species. The speciic name is intended as a noun in apposition. Remarks on Halichondria (Halichondria). The subgenus Halichondria is large with approx. 95 valid species (Van Soest et al. 2008). Table 7 lists species distributed within the Central Indo-Paciic Realm (following Spalding et al. 2007) which includes the study area and adjacent areas. Some of those species, in particular the in substrate, and small tapering erect projections (from 10 mm long and less than 5 mm diameter) at apex. Erect projections vary in shape, i.e. istulose-like, convoluted, bifurcating and lobate; generally small 30–70 mm diameter, 10–20 mm thick. Colour. Orange. Blue-grey, in alcohol. Consistency and texture. Soft, easily torn, rubbery. Oscula. Apical and conspicuous on small digits, with membranous rims. Surface. Semitransparent at apex of digits in some specimens. Skeleton (Fig. 13A,B). Ectosomal skeleton, halichondroid, formed by a thin (less than 50 µm), wavy, loose membranous layer, with oxeas tangentially oriented and supported by choanosomal tracts (Fig. 13A). Choanosomal skeleton, halichondroid, formed by ill-deined bundles of larger spicules, criss-crossing and without any particular orientation connected by short bundles and single spicules. In subectosomal region, skeleton becoming cavernous with large lacunae, 100–600 µm diameter (Fig. 13B) and vague reticulation of pauci-multispicular, illdeined tracts, diverging towards surface or condensed and running nearly parallel to it, supporting ectosomal skeleton. High densities of ilamentous cyanobacteria present in both ectosome and choanosome, especially near surface. Spongin or collagen scarce. Spicules (Fig. 13C; Table 12). Oxeas, hastate, straight, possibly in two size categories, larger and thicker (249–587 x 5–14 µm) and smaller and thinner (88–217 x 3–7 µm). Style modiications slightly common within larger category. Remarks. We compared Halichondria microbiana with the valid species recorded for the Central West Table 12. Comparison of spicule dimensions between specimens of Halichondria (Halichondria) microbiana. Specimen NTM Z.5907 Locality Darwin Harbour, NT NTM Z.5908 Bynoe Harbour, NT Oxea type I 87.9–203µm (126.5±24.8) x 3.3–7.1µm (5±0.9) 91.3–217µm (137.8±32) x 2.9–7.2µm (5.1±1) 74 Oxea type II 249.2–587µm (482.9±75.1) x 6.3–14.2µm (8.9±1.8) 282.5–572.1µm (446.6±93.5) x 5.2–13.2µm (8.8±2) Halichondriidae from northern Australia A C B Fig. 13. Halichondria (Halichondria) microbiana sp. nov. (NTM Z.5908): A, light microphotograph of tangential section of ectosomal skeleton, showing oxeas on membranous thin layer; B, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and large lacuna at subectosomal level (left upper corner of section); C, diagram of spicules. Scale bars: A, 200 µm; B, 500 µm; C, 50 µm. ones described by Kieschnick (1896) and Lindgren (1897), are unrecognisable and their type material has never been relocated. Other species reported for the area have been allocated to other genera as result of this revision, thus Halichondria (Halichondria) is represented within this realm by nine species including the two new species described above. The subgenus Eumastia Schmidt, 1870 is not represented in the study area at all and it is reserved for Halichondrialike species from high latitudes (Erpenbeck & Van Soest 2002). Genus Hymeniacidon Bowerbank, 1858 Gender: feminine. Type species, by subsequent designation of Bowerbank (1864), Hymeniacidon caruncula Bowerbank, 1958. Recent, Tenby, Wales. 75 Hymeniacidon gracilis (Henstschel, 1912) (Fig. 14) Stylotella digitata gracilis Henschel, 1912: 356. Hymeniacidon gracilis. – Hooper et al. 1997. Material examined. As listed by Hooper et al. (1997). Remarks. The species was well described by Hooper et al. (1997) and elevated to full species rank in the genus Hymeniacidon. We provide a new illustration of the type material (Fig. 14) and additional spicule measurements (Table 13) to complement the description given by Hooper et al. (1997). Only three specimens are recorded for the study area; no new material was located in recent collections. Distribution. As recorded by Hooper et al. (1997). Remarks on the genus Hymeniacidon. Hymeniacidon gracilis is the only valid species of the genus from the study area. Other species of Hymeniacidon recorded for the B. Alvarez and J. N. A. Hooper A B Fig. 14. Hymeniacidon gracilis (Synype SMF 970). A, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton; B, diagram of spicules. Scale bars: A, 500 µm; B, 50 µm. these style modiications are common among the genus. Population genetics and additional morphometric analyses might reveal whether these style modiications of Stylissa are reliable characters for the separation of species. Sahul Shelf Province and adjacent areas are H. vernonensis Hooper et al., 1997 and H. laccida Pulitzer-Finali, 1996. The material described under H. vernonensis by Hooper et al.(1997) was revised and it does not agree with the current concept of the genus. The species is formally transferred here to the dictyonellid genus Stylissa. As admitted by Hooper et al. (1997), is very similar to Stylissa labelliformis but S. vernonensis includes distinctive styles, curved at the centre, sinuous or rhabdose and frequent anisoxeas with one telescoped point. It should be noted however, that these spicule modiications are common among species of Stylissa and therefore they are not reliable for the delimitation of species. Both species are similar in growth form, surface characteristics and skeletal organisation. The choanosomal skeleton of S. vernonensis however, is nearly halichondroid with only vague tracts of spicules and with a much higher spicule density and no spongin ibres (whereas in S. labelliformis is vaguely plumo-reticulated and with well developed spongin ibres). The type specimen of Hymeniacidon laccida (MSNG 48703) from Laing Is., Papua New Guinea, was re-examined and it does not correspond to the genus Hymeniacidon. The species belongs also to the dictyonellid genus Stylissa and is likely to be conspeciic with Stylissa massa (Carter, 1887). The specimen examined however, includes distinctive subtylostyles transitional to strongyles with tylote modiications. As is the case with S. vernonensis, Genus Topsentia Gender feminine. Type species, by original designation, Anisoxya glabra Topsent, 1898. Recent, Azores Is. Topsentia dura (Lindgren, 1897) (Figs 10G, 15) Halichondria dura Lindgren, 1897 : 480. Topsentia dura.– Hooper et al. 1997: 14. Material examined. As listed by Hooper et al. (1997). addiTional maTerial.– Darwin Harbour, N.T.: Z.5209, Z.5233. Wessel Is: Z.5234. Remarks. Hooper et al. (1997) assigned material from the Beagle Gulf to this species under the genus Topsentia. Additional specimens from recent collections agree also with this material and are assigned to this species. Further illustrations (Figs 10G, 15) and spicule measurements (Table 14) are provided here to complement that description. The description agrees with the current concept of Topsentia, however it remains inconclusive whether the material from the Beagle Gulf is conspeciic with Lindgren’s species from Indonesia as the type was not examined. Examination of additional specimens from Indonesia (Alvarez & De Voogd unpublished data) and a re-description of the type might provide additional evidence to conirm if these populations belong in the same species. Distribution. Indonesia [?], Darwin Harbour and Wessel Is. Topsentia dura is occurs the intertidal zone to 25 m depth. Remarks on Topsentia. Hooper et al. (1997) described material under the Red Sea species Topsentia halichondrioides (Dendy, 1905) that seems very similar Table 13. Comparison of spicule dimensions between specimens of Hymeniacidon gracilis. Specimen SMF 970 Locality Indonesia NTM Z.883 Darwin Harbour Styles 220.5–261.1µm (238.4±9.6) x 3.3–8.9µm (6.1±1.5) [25] 243.1–279.1µm (265.6±11.1) x 3–7.4µm (5.1±1.1) 76 Halichondriidae from northern Australia A B C D E F Fig. 15. Topsentia dura (NTM Z.5209): A, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and palisade of erect oxeas at surface; B, diagram of spicules. Topsentia halichondrioides (QM G303442); C, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton and oxeas oriented perpendicularly at surface level; D, diagram of spicules. Topsentia ridleyi (QM G303309, Holotype); E, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton; F, diagram of spicules. Scale bars: A,C,E, 500 µm; B,D,F, 100 µm. to T. dura (Fig. 15B,C). These species are massive, of hard consistency with skeletons made of a confused mass of oxeas of similar dimensions, not clearly differentiated into size classes (see Table 14) and differing only in colouration and shape of oscules (i.e. volcano-shaped in 77 T. halichondrioides, and sunken and small in T. dura). Halichondria ridleyi Hooper et al., 1997 is referred here to the genus Topsentia (comb. nov.) and it is also very similar to T. dura (Fig. 15E,F). It differs from T. dura in having some surface istules and processes. B. Alvarez and J. N. A. Hooper Table 14. Comparison of spicule dimensions among specimens assigned to Central Paciic species of Topsentia. Species Topsentia ridleyi Specimen Locality QM G303309 Darwin Harbour Topsentia ridleyi NTM Z.3262 Cobourg Peninsula Topsentia halichondrioides G303442 Bynoe Harbour Topsentia halichondrioides NTM Z.5233 East Point Topsentia dura NTM Z.5234 Wessel Is Topsentia dura NTM Z.5209 Darwin Harbour Topsentia dura NTM Z.3178 Darwin Harbour Topsentia dura NTM Z.1442 Gunn Point Topsentia indica SMF 997 Aru Is, Indonesia Oxea type I 142.3–273.5µm (207.7±39.4) x 3.3–7.6µm (5.4±1.2) 160.6–331.2µm (257.4±50.3) x 4.2–12.3µm (8.2±2.3) 111.4–165.9µm (142.3±13.8) x 5.9–8.9µm (7.2±0.9) 142.4–333.5µm (220.1±47.2) x 3.7–10.3µm (7.3±1.7) 151.2–399.7µm (248.3±61.9) x 3–11.7µm (7.6±2.3) 189.4–357.5µm (269.9±41.9) x 4.1–9.7µm (7.1±1.4) 189.6–387.1µm (254.5±52.5) x 5.5–12.5µm (7.8±2) An additional species of Topsentia recorded from Aru Is, Indonesia, is T. indica Hentschel, 1912 (syntype SMF 995 and 997, examined). The differentiation of these species using traditional morphological characters is extremely subjective. Molecular and morphometric studies of local populations might contribute to a better understanding of the concept of this species. Other species of Topsentia recorded for the Sahul Province and adjacent areas that are better placed elsewhere include Topsentia maculosa Pulitzer-Finali, 1996 from Papua New Guinea (it belongs in Amorphinopsis, see above) and Topsentia plurisclera Pulitzer-Finali, 1996 (holotype, MSNG 48702, examined) is a species of Petrosia. DISCUSSION This revision of species of Halichondriidae from northern Australia recognises a total of 15 species belonging to the genera Amorphinopsis, Axinyssa, Ciocalypta, Halichondria (Halichondria), Hymeniacidon and Topsentia. Other genera of the family (i.e. Epipolasis, Laminospongia, Vosmaeria, Ciocalapata and Spongosorites are not represented in the Sahul Shelf Province. Epipolasis and Spongosorites are however represented in the Northeast Australian Shelf (Alvarez and Hooper, unpublished data). Of the species reported in this revision, Axinyssa bergquistae, Ciocalypta vansoesti and the two new species Halichondria (Halichondria) carotenoidea and H (H.) microbiana are so far known only from northern Australian waters. The rest of the species have extralimital distributions through the Central Indo-Paciic realm. Axinyssa mertoni (Hentschel, 1912) in its new generic combination is recorded for northern Australia and it represents a new record for the study area. 78 Oxea type II 335.6–576.7µm (443.9±52.7) x 6.7–16.6µm (12±2.6) 503–848.1µm (653.8±83.5) x 13.4–31.5µm (20.9±4.6) 421.6–716.5µm (555.3±62.7) x 14–24.4µm (19.3±2.6) 487.6–914.9µm (711.2±99.1) x 13–30µm (18.9±4.4) 348.6–568.1µm (448.9±57.8) x 11–23.7µm (16.8±3.2) 373.9–656.5µm (477.9±79.2) x 10–23.9µm (15±3.2) 352.5–525.1µm (441.4±46.2) x 9.9–18.5µm (13.7±2.4) 341.2–611.2µm (482.6±69.2) x 8.3–22.7µm (15.6±3.4) 493.7–1078.1µm (734.4±125) x 11.1–39.4µm (21.7±5.5) As in other members of the order Halichondrida, particularly in Axinellidae and Dictyonellidae, species within and across all genera of Halichondriidae are extremely dificult to delimit. This problem is demonstrated by the large number of misidentiications in previous studies due to the lack of adequate generic deinitions and also to the poor understanding of the importance, or indeed relevance, of some of the alleged pivotal characters that currently differentiate both species-groups and genera within the Halichondriidae. The additional information obtained from the collection of new material plus the revised generic deinitions of the family (Erpenbeck & Van Soest 2002) allowed us to clarify the concept of halichondriid species of northern Australia and to allocate them to more appropriate genera. Nevertheless, differentiating species within Halichondriidae continues to be ambiguous based solely on the present limited suite of accepted morphological characters, with a number of them shared among species and even genera. For example, Ciocalypta heterostyla, C.vansoesti, and Axinyssa mertoni are species with istulelike growth form, nearly indistinguishable in the ield (Fig 1). However, they all are easily diagnosed based on skeletal characteristics. This suggests that the growth form of these species in particular, might be an adaptation to the habit where they occur (i.e. soft and muddy sediments). Amorphinopsis foetida and A. maculosa are also very similar in habit and in their skeletal characteristics but they can be distinguished by the predominance of oxeas and styles. Separation of species based on the dominance of styles or oxeas however might be debatable, as this could possibly be related to intraspeciic variation as seen in some species of Axinellidae (Alvarez et al. 1998; Alvarez & Hooper 2009). Spicule morphometrics (i.e. size variation of spicules) within Halichondriidae seems to be a useful tool for the Halichondriidae from northern Australia ACKNOWLEDGEMENTS differentiation of species. Ciocalypta stalagmites, for example, is distinguished from the other species by the distinctive and nearly constant size of two categories of oxeas. Similarly, Halichondria (H.) microbiana can be distinguished from H. (H.) carotenoidea by the size of the larger category of oxeas. On the other hand, all the Axinyssa species we have studied have always a mixture of oxeas in a large size range, thus spicule sizes is not a useful character for the distinction of species within this genus. These examples indicate that the characters currently used to separate genera and species within the family are extremely homoplastic and suggest that genera and species within Halichondriidae might be non-monophyletic. The study of local populations using both morphological and genetic methods will help to clarify whether these taxa are monophyletic. This study represents the inal contribution to the present taxonomic revision of the order Halichondrida of northern Australia, restricted to the marine province identiied as the Sahul Shelf in the classiication of Spalding et al. (2007). The result of this and previous studies (Alvarez & Hooper 2009, 2010) indicates that the group is represented in the study area mainly by the families Axinellidae (Alvarez & Hooper 2009), Dictyonellidae (Alvarez & Hooper 2010) and Halichondriidae (this revision). One additional family, the Heteroxyidae (formerly Desmoxyidae see Van Soest & Hooper 2005), is represented in the area by two very common species: Myrmekioderma granulatum (Esper, 1830), which is documented and illustrated by Hooper et al. (1997); and Higginsia mixta (Hentschel, 1912). The type of H. mixta (SMF 968) was examined and it agrees with material deposited at the collections of QM and NTM and recorded for the area of study. The ectosomal skeleton of the specimens studied have a relatively thick tangential crust formed by a dense mass of spined microxeas and interrupted by disorganised brushes of thin raphidiform oxeas and extra long stylesstrongyles (mostly broken in the preparations) projecting through surface. The similarities of this type of skeletal organisation with raspailiid genera such as Ceratopsion are remarkable and worth further investigation. Phylogenetic afinities based on molecular data (Erpenbeck et al. 2005) indicated that members of Heteroxyidae (i.e. Didiscus and Myrmekioderma) are closely related to the axinellid genera Reniochalina and Ptilocaulis. The position of these genera within Axinellidae (Halichondrida) and its relationships based on molecular data with other raspailiid genera such as Axechina has already been discussed (Alvarez 2009 and references within). Sequencing data of additional genera currently allocated to Heteroxyidae, and in particular of the species represented in the Sahul Shelf province will help to clarify these relationships and classiication. The family Bubaridae is the only family of Halichondrida not represented in the area of study. This work was funded by an Australian Biological Research Studies Research grant (No 205-10) and by the ‘Collection and Taxonomy of Shallow Water Marine Organisms’ program for the U.S. National Cancer Institute (Contract N02-CM-27003 and Contract N02CO-2009-00012) subcontracted to the MAGNT through CRRF. We thank the following people: Michael Browne and Huy Nguyen, for their invaluable assistance during 2002–2004 MAGNT ield collections; Dr Pat Colin, CRRF, and Don DeMaria, for their assistance and photographic work during the MAGNT ield collections in the 2004 Wessel Is; Terry Yumbuluy, Wessel Is, for allowing collections of sponges in his sea area; Merrick Ekins, QM, for his assistance interrogating QM database and making specimens available for study and Dr Kathryn Hall for substantial re-checking voucher specimens housed in the collections of the QM to conirm or refute previous species diagnoses. Drs Rob W.M. Van Soest (ZMA) and Nicole De Voogd are thanked for their input and valuable discussions on Indo-Paciic halichondrid sponges. Mr Swee-Cheng Lim is thanked for his assistance locating type material at MSNG. Drs Richard Willan and Chris Glasby (MAGNT) are thanked for their continuous advice and suggestions during the preparation of this manuscript. And lastly, we thank the two referees for their valuable suggestions. REFERENCES Alvarez, B. & Hooper, J.N.A. 2009. Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) from northern Australia. Family Axinellidae. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 25: 17–42. Alvarez, B. & Hooper, J.N.A. 2010. Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) from northern Australia. Family Dictyonellidae. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 26: 13–36. Alvarez, B., Van Soest, R.W.M. & Rützler, K. 1998. A revision of the species of Axinellidae (Porifera: Demospongiae) in the Central-West Atlantic region. Smithsonian Contributions to Zoology 598: 1–47. Bergquist, P.R. 1965. The sponges of Micronesia, Part 1. The Palau Archipelago. Paciic Science 19: 123–204. Bowerbank, J.S. 1862. On the anatomy and physiology of the Spongiadae. Part III: On the generic characters, the speciic characters and on the method of examination. Philosophical Transactions of the Royal Society of London 152: 1087–1135, pls. 72–74. Bowerbank, J.S. 1864. A monograph of the British Spongiadae. Volume 1. The Ray Society, London. Bowerbank, J.S. 1873. Contributions to a general history of the Spongiadae. Part 4. Proceedings of the Zoological Society of London for the year 1873: 3–25 pls. 1–4. 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: 109–138, pls.1–2. 79 B. Alvarez and J. N. A. Hooper Kieschnick, O. 1896. Silicispongiae von Ternate nach den sammlungen von Herrn Prof. Dr. W. Kükenthal. Zoologischer Anzeiger 19: 526–534. Lendenfeld, R.V. 1897. Spongien von Sansibar. Abhandlungen Herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 21: 93–133, pls 9–10. Lim, S-C., De Voogd, N. & Tan, K-S. 2008. A guide to sponges of Singapore. Science Centre Singapore: Singapore. Lindgren, N.G. 1897. Beitrag zur kenntniss der spongienfauna des Malaiischen Archipels und der Chinesischen Meere. Zoologischer Anzeiger 20: 480–487. Lindgren, N.G. 1898. Beitrag zur kenntniss der spongienfauna des Malayischen Archipels und der Chinesischen Meere. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Thiere 11: 283–378, pls 17–20. Pulitzer-Finali, G. 1996. Sponges from the Bismarck Sea. Bolletino dei Musei e degli Istituti Biologici dell’Università di Genova 60–61: 101–138. Ridley, S.O. 1884. Spongiida. Pp. 366–684. Report on the Zoological Collections made in the Indo-Paciic Ocean during the Voyage of H.M.S. ‘Alert’ 1881-2. British Museum, Natural History: London. Ridley, S.O. & Dendy, A. 1886. Preliminary report on the Monaxonida collected by H.M.S ‘Challenger’. Part I. The Annals and Magazine of Natural History (Series 5) 18: 325–351. Sollas, I.B.J. 1902. On the sponges collected during the Skeat expedition in the Malay Peninsula (1899-1900). Proceedings of the Zoological Society of London 2: 210–221. Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S.A., Martin, K.D., McManus, E., Molnar, J., Recchia, C.A. & Robertson, J. 2007. Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. Bioscience 57: 573–583. Thiele, J. 1899. Studien über Paziische Spongien. II. Zoologica 24: 1–33, pls. 1–5. Thiele, J. 1900. Kieselschwämme von Ternate. I. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 25: 19–80. Van Soest, R.W.M. 1991. Demospongiae higher taxa classiication re-examined. Pp. 54–71. In: Reitner, J. & Keupp, H. (eds) Fossil and Recent Sponges. Springer-Verlag: Berlin. Van Soest, R.W.M, Boury-Esnault, N., Hooper, J.N.A., Rützler, K, de Voogd, N.J., Alvarez de Glasby, B., Hajdu, E., Pisera, A.B., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R., Klautau, M., Picton, B., Kelly, M., Vacelet, J. 2008. World Porifera database. Available online at http://www. marinespecies.org/porifera. Last consulted 27 July 2011. Van Soest, R.W.M., Díaz, M.C. & Pomponi, S.A. 1990. Phylogenetic classiication of the halichondrids (Porifera, Demospongiae). Beaufortia 40: 15–62. Van Soest, R.W.M. & Hooper, J.N.A. 2005. Resurrection of Desmoxya (Porifera: Halichondrida), with the description of a new species from Rockall Bank bathyal coral reefs, North Atlantic. Journal of the Marine Biological Association of the United Kingdom 85: 1367–1371. Wilson, H.V. 1925. Silicious and horny sponges collected by the U.S. Fisheries Steamer ‘Albatross’ during the Philippine expedition, 1907-10. Bulletin of the United States National Museum 2: 273–532. Burton, M. 1934. Sponges. Scientiic Reports of the Great Barrier Reef Expedition 1928-29 4: 513–621, pls.1–2. Burton, M. 1959. Sponges. Scientiic Reports of the John Murray Expedition 1933-34 10: 151–281. Carter, H.J. 1887. Report on the marine sponges, chiely 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, pls 5–7. De Laubenfels, M.W. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular, and the West Indies in general, with material for a revision of the families and orders of the Porifera. Papers from the Tortugas Laboratory 30: 1–225, pls. 1–22. De Laubenfels, M.W. 1954. The Sponges of the West-Central Paciic. Oregon State Monographs, Studies Zoology: 1–306, pls. 1–12. Dendy, A. 1889. Report on a second collection of sponges from the Gulf of Manaar. Annals and Magazine of Natural History 3: 73–99, pls. 3–5. Dendy, A. 1905. Report on the sponges collected by Professor Herdman, at Ceylon, in 1902. Pp. 57–246, pls 1–16. In: Herdman, W.A. (ed.) Report to the Government of Ceylon on the pearl oyster Fisheries of the Gulf of Manaar. Royal Society: London. Dendy, A. 1922. Report on the Sigmatotetraxonida collected by H.M.S. ‘Sealark’ in the Indian Ocean. Transactions of the Linnean Society of London 18: 1–164, pls. 1–18. 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 pls. 25–26. Erpenbeck, D., Breeuwer, J. & Van Soest, R.W.M. 2005. Implications from a 28S rRNA gene fragment for the phylogenetic relationships of halichondrid sponges (Porifera: Demospongiae). Journal of Zoological Systematics and Evolutionary Research 43: 93–99. Erpenbeck, D. & Van Soest, R.W.M. 2002. Family Halichondriidae. Pp. 773–786. In: Hooper, J.N.A. & Van Soest, R.R.M. (eds) Systema Porifera. A guide to the supraspeciic classiication of the phylum Porifera. Plenum Press: New York. Hentschel, E. 1912. Kiesel-und hornschwämme der Aru und KeiInseln. Abhandlungen Senckenbergiana Naturforschende Gessellschaft: Hamburg. Hooper, J.N.A. 2005. Porifera. Australian Faunal Directory. Australian Biological Resources Study, Canberra. Last consulted 18 May 2011. http://www.environment.gov.au/ biodiversity/abrs/online-resources/fauna/afd/taxa/. Hooper, J.N.A. & Bergquist, P.R. 1992. Cymbastela, a new genus of lamellate coral reef sponges. Memoirs of the Queensland Museum 32: 99–137. 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 order Poecilosclerida and Axinellida (Porifera: Demospongiae). Invertebrate Taxonomy 6: 261–301. Hooper, J.N.A., Cook, S.D., Hobbs, L.J., Hooper, L.G. & Kennedy, J.A. 1997. Australian Halichondriidae (Porifera: Demospongiae): I. Species from the Beagle Gulf Marine Park. Pp. 1–65. In: Hanley, J.R., Caswell, G., Megirian, D. & Larson, H.K. (eds) The marine lora and fauna of Darwin Harbour, Northern Territory, Australia. Museums and Art Galleries of the Northern Territory: Darwin. Hooper, J.N.A. & Wiedenmayer, F. 1994. Porifera. Pp. 1–624. In: Wells, A. (ed.) Zoological Catalogue of Australia. Volume 12. Pp 1–624. CSIRO: Melbourne. Accepted 4 November 2011 80 Halichondriidae from northern Australia APPENDIX I G300854 G301034 G303252 G303287 G303309 G303351 G303442 G303450 G303524 G303541 G303558 G303560 G303561 G303595 G303658 G303677 G310137 G310170 G313543 G313572 G313577 G314246 G314247 G314255 G314267 G315205 G315207 G320819 G320904 Collection and locality data of material examined in the collections of QM and NTM QM material Gulf of Carpentaria, northern central region, QLD, 9°36.0001’ S, 136°6.01’ E, 52 m , 23 Nov 1991, coll. Cook, SD. on CSIRO RV Southern Surveyor SW Vrilya Point, SW, Gulf of Carpentaria, QLD, 11°29.0167’ S, 142°55.09’ E, 18 m , 1 Dec 1991, coll. Cook, SD. on CSIRO RV Southern Surveyor South Shell I., reef N of boatramp, East Arm, Darwin Harbour, NT, 12°29.1334’ S, 130°53.09’ E, 0 m , 19 Sep 1993, coll. Hooper, JNA & Hobbs, L.J. South Shell I., reef N of boatramp, East Arm, Darwin Harbour, NT, 12°29.1334’ S, 130°53.09’ E, 0 m, 19 Sep 1993, coll. Hooper, JNA & Hobbs, LJ Dudley Point Reef, East Point, Darwin, NT, 12°25.05’ S, 130°49.01’ E, 0 m , 20 Sep 1993, coll. Hooper, JNA & Hobbs, L.J. East Point Bommies, Darwin Harbour, NT, 12°24.0834’ S, 130°48.14’ E, 10 m , 23 Sep 1993, coll. Hooper, JNA & Hobbs, LJ Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 m , 26 Sep 1993, coll. Hooper, JNA & Hobbs, LJ Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 m , 26 Sep 1993, coll. Hooper, JNA & Hobbs, LJ Duyfken Point, W Gulf of Carpentaria, QLD, 12°41.0501’ S, 141°3.01’ E, 42 m, 11 Nov 1993, coll. Cook, SD. & Kennedy,J. on CSIRO RV. Southern Surveyor Vernon Is, W of South West Vernon I., NT, 12°6.15’ S, 131°4.14’ E, 13 m, 10 Oct 1993. Cape Hotham, NW of cape, NT, 12°1.05’ S, 131°13.16’ E, 34 m, 9 Oct 1993. Bynoe Harbour, 2 nmls E Fish Reef, NT, 12°24.1334’ S, 130°28.16’ E, 17 m, 6 Oct 1993, coll. CCNT Ocean Rescue 2000 Program Shoal Bay, outer region of bay, NT, 12°6.15’ S, 130°49.16’ E, 18 m, 12 Oct 1993, coll. CCNT Ocean Rescue 2000 Program Vernon Is, W of Knight Reef, NT, 12°1.0334’ S, 131°3.16’ E, 22 m, 11 Oct 1993. Vernon Is, N marsh Shoal, NT, 12°07.0001’ S, 130°56.1’ E, 16 m, 11 Oct 1993, coll. CCNT stn. 138. Dredge Shoal Bay, middle of bay, NT, 12°13.0167’ S, 130°56’ E, 17 m, 12 Oct 1993, coll. CCNT Ocean Rescue 2000 Program Parry Shoals 35nm W Bathurst I., NT, 11°7.0321’ S, 129°25.9’ E, 16 m, 12 Aug 1987, coll. mussig, AM and NCI team Darwin Harbour, NT, 12°15.1834’ S., 130°29.11’ E., 9 m depth, 17 August 1987, coll. mussig, AM and NCI team N Bathurst I., Timor Sea, NT, 11°13.98’ S, 130°34.21’ E, 41.2 m, 5 Oct 1997, coll. Cook, SD. on RV Southern Surveyor SW Groote Eylandt, NT, 14°25.0801’ S, 135°58.51’ E, 20.3 m, 13 Oct 1997, coll. Cook, SD. on RV Southern Surveyor SW Groote Eylandt, NT, 14°20.22’ S, 136°34.98’ E, 19.6 m, 14 Oct 1997, coll. Cook, SD. on RV Southern Surveyor N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136°18.13’ E, 20 m, 27 Sep 1998, coll. Leys, SP. on RV Southern Surveyor N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136°18.13’ E, 21.7 m, 27 Sep 1998, coll. Leys, SP. on RV Southern Surveyor W of Groote Eylandt, Gulf of Carpentaria, NT, 14°8’ S, 136°8’ E, 13 m, 6 Oct 1998, coll. Leys, SP. on RV Southern Surveyor SW of Groote Eylandt, Gulf of Carpentaria, QLD, 14°20’ S, 136°2’ E, 22.1 m, 6 Oct 1998, coll. Leys, SP. on RV Southern Surveyor W Groote Eylandt, Gulf of Carpentaria, NT, 14°8.5667’ S, 136°16.51’ E, 21.4 m, 13 Oct 1998, coll. Wassenberg T SW of Groote Eylandt, Gulf of Carpentaria, NT, 14°22.5’ S, 136°9.12’ E, 22.2 m, 13 Oct 1998, coll. Wassenberg T Gulf of Carpentaria, QLD, 15°20.037’ S, 140°19.84’ E, 28 m, 24 may 2003, coll. Bartlett C, Cook S on RV Southern Surveyor 2380403 CSIRO “Effects of Trawling” Gulf of Carpentaria, QLD, 15°20.037’ S, 140°19.84’ E, 28 m, 11 mar 2003, coll. Bartlett C, Cook S on RV Southern Surveyor 2380403 CSIRO “Effects of Trawling” 81 B. Alvarez and J. N. A. Hooper APPENDIX I (continued) Z.84 Z.131 Z.194 Z.241 Z. 592 Z 712 Z.919 Z.934 Z.941 Z.945 Z.986 Z.987 Z.1097 Z.1358 Z.1391 Z.1395 Z.1442 Z.1979 Z.1991 Z.2018 Z.2026 Z.2086 Z.2215 Z.2245 Z.2648 Z.2651 Z.2697 Z.3106 Z.3133 Z.3147 Z.3178 Z.3195 Z.3205 Z.3262 Z.3920 Z.4085 Z.4093 Z.4100 Z.4122 Z.4123 Z.4125 Z.4451 Collection and locality data of material examined in the collections of QM and NTM NTM material Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.5001’S, 132°2’E, 18 Oct 1981, coll. Hooper, JNA & Alderslade, PN Sandy I. No.2, Cobourg Peninsula, NT, 11°5.5001’S, 132°17’E, 10 m, 21 Oct 1981, coll. Hooper, JNA & Alderslade, PN Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, 0–0.5 m, 13 Sep 1981, coll. Hooper, JNA and party Indian I., Bynoe Harbour, NT, 12°35’S, 130°33.01’E, 3 m, 18 Nov 1981, coll. Byers,P.,F.V. Skeleton Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5’S., 132°10.51’E., 3 m depth, 4 May 1982, coll. Hooper, JNA N Adele I.,Collier Bay, NW Shelf, WA 15°58.0167’S., 122°39.07’E., 59 m depth, 21 April 1982, coll. R.V.SPRIGHTLY, dredge. Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, 10 m, 31 Aug 1982, coll. Hooper, JNA East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 26 Oct 1982, coll. Hooper, JNA Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 26 Oct 1982, coll. Hooper, JNA Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, m, 22 Dec 1982, coll. Hooper, JNA Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3’S, 132°3.71’E, .5–6 m, 16 May 1983, coll. Hooper, JNA Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3’S, 132°3.71’E, 6 m, 17 May 1983, coll. Hooper, JNA Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°10.4’S, 132°2.8’E, 2 m, 19 May 1983, coll. Hooper, JNA Blue Hole, Gunn Point, NT, 12°9.0001’S, 131°0’E, 25 m, 19 Aug 1983, coll. Alderslade, PN West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E , 11 May 1984, coll. Hooper, JNA and party Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 20 Jul 1984, coll. Hooper, JNA Vestey’s Beach, Bullocky Point, Darwin, NT, 12°26.2’S, 130°49.89’E 21 Jan 1985, coll. Hooper, JNA Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 10 m, 12 Apr 1985, coll. Hood, C and party Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 3 Apr 1986, coll. Hooper, JNA and party Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party Parry Shoals, Arafura Sea, NT, 11°12.5167’S, 129°42.07’E, 20 m, 15 Aug 1987, coll. Mussig, AM and NCI team Parry Shoals, Arafura Sea, NT, 11°11.4’S., 129°43.01’E., 18 m depth, 13 August 1987, coll. Mussig, AM and NCI team Parry Shoals, Arafura Sea, NT, 11°12.5167’S, 129°42.07’E, 16 m, 15 Aug 1987, coll. Mussig, AM and NCI team East Point Reef, East Point, Darwin, NT, 12°29.5’S, 130°48.01’E, 0.5 m, 10 Sep 1987, coll. Smit, N Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 9 m, 16 Sep 1987, coll. Smit, N Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 25 Sep 1987, coll. Smit, N Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5’S, 132°10.51’E, 11 Sep 1986, coll. Hooper, JNA & Johnson,C Cumberland Strait, NE bay, Wessel Is, Gove Peninsula, NT, 11°26.8’S, 136°30.2’E, 13 m, 14 Nov 1990, coll. Hooper, JNA Near Boat Ramp, East Arm Port, Darwin, NT, Australia, 12º29.8’S, 130º53.5’E, intertidal, 20 September 2001, coll. B. Glasby & party, by hand Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 20 September 2001, coll. B. Glasby & party, by hand Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 18 October 2001, coll. B. Glasby & party, by hand Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 18 October 2001, coll. B. Glasby & party, by hand Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 19 October 2001, coll. B. Glasby & party, by hand Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 19 October 2001, coll. B. Glasby & party, by hand Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.103’S, 130°47.111’E, 8–14 m, 7 May 2002, coll. Alvarez, B and party 82 Halichondriidae from northern Australia Z.5206 Z.5207 Z.5208 Z.5209 Z.5210 Z.5211 Z.5212 Z.5213 Z.5215 Z.5216 Z.5217 Z.5218 Z.5219 Z.5221 Z.5222 Z.5223 Z.5224 Z.5225 Z.5226 Z.5228 Z.5229 Z.5230 Z.5233 Z.5234 Z.5736 Z.5901 Z.5902 Z.5903 Z.5904 Z.5905 Z.5906 Z.5907 Z.5908 South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 10–12 m, 21 Aug 2002, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5–10 m, 23 May 2003, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 7–12 m, 24 Jul 2003, coll. Alvarez, B and party Nightcliff bommies, off Nightcliff jetty, Darwin Harbour, NT, 12°22.751’S, 130°50.116’E, 5–8 m, 8 Aug 2003, coll. Alvarez, B and party Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22 Aug 2003, coll. Alvarez, B and party Weed Reef, entrance to West Arm, Darwin Harbour, NT, 12°29.25’S, 130°47.54’E, 9–12 m, 6 Sep 2003, coll. Nguyen, H off Herbert Point, Indian I., Bynoe Harbour, NT, 12°34.586’S, 130°31.419’E, 0–5 m, 24 Jun 2003, coll. Alvarez, B and party South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7–11. m, 19 Aug 2002, coll. Alvarez, B and party West Arm, 2.5 km N of Stokes Point, Darwin Harbour, NT, 12°31.300’S, 130°48.500’E, 4–5 m, 3 Aug 2002, coll. Alvarez, B and party Wickham Point, 2.5 km SW of East Arm Wharf, East Arm, Darwin Harbour, NT, 12°30.12’S, 130°52.39’E, 4–7 m, 15 Sep 2002, coll. Alvarez, B and party Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22 Aug 2003, coll. Alvarez, B Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2001’S, 130°47.1’E, 5–19 m, 8 May 2002, coll. Alvarez, B and party South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7–14 m, 20 Aug 2002, coll. Alvarez, B and party East Arm Wharf, East Arm, Darwin Harbour, NT, 12°29.19’S, 130°53.35’E, 0.6 m, 1 Mar 2002, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5–10 m, 23 May 2003, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 3–7 m, 25 May 2003, coll. Alvarez, B and party Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.351’S, 130°31.454’E, 6–8 m, 11 Jun 2003, coll. Alvarez, B and party Moira Reef, Bynoe Harbour, NT, 12°30.799’ S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Browne, M Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 12–18 m, 16 Sep 2002, coll. Alvarez, B and party Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, 130°38.371’E, 9–12 m, 23 Aug 2003, coll. Browne, M off Dudley Point, Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4–7 m, 4 Jun 2002, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Mandorah jetty, NW Cox Peninsula, Darwin Harbour, NT, 12°26.55’S, 130°46.05’E, 9–12 m, 5 Sep 2003, coll. Alvarez, B and party Moira Reef, Bynoe Harbour, NT, 12°30.799’S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Alvarez, B and party Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 12–24 m, 3 Sep 2002, coll. Alvarez, B and party Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, 130°38.371’E, 9–12 m, 23 Aug 2003, coll. Nguyen, H West Arm, 2.5 km N of Stokes Point, Darwin Harbour, NT, 12°31.300’S, 130°48.500’E, 4–5 m, 3 Aug 2002, coll. Alvarez, B and party Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m, 7 Nov 2006, coll. Alvarez, B Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m, 7 Nov 2006, coll. Alvarez, B Lee Point, Darwin, NT, 12°20.538’S, 130°52.184’E, 9–12 m, 7 Aug 2003, coll. Nguyen, H Moira Reef, Bynoe Harbour, NT, 12°30.799’S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Nguyen, H 83 B. Alvarez and J. N. A. Hooper Z.5909 Z.5915 Z.5923 Z. 5925 Z.5928 Z.5948 Z.5949 Z.5950 Z.5965 Z.5970 Z.5976 Z.5977 Z.5978 off Dudley Point, Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4–7 m, 4 Jun 2002, coll. Alvarez, B and party Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 20 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009, coll. Sultana, S, SCUBA. Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA. East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12º25.01’S., 130º48.88’E., 6 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12º25.01’S., 130º48.88’E., 6 m depth, 21 May 2009, coll. Alvarez, B and Sultana, S, SCUBA Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22 Aug 2003, coll. Alvarez, B and party Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 13–16 m, 5 Sep 2003, coll. Alvarez, B and party Larrakeyah sewerage outfall, Darwin Harbour, NT, 12º28.04’S, 130º49.77’E, 5.8 m depth, 22 July 2010, coll. Sultana, S, SCUBA 84

RELATED PAPERS

RELATED TOPICS

Log In

Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae

Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of northern Australia. Family Halichondriidae

Related Papers

RELATED PAPERS

RELATED TOPICS