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

John N . A . Hooper; Belinda Glasby

Nine species in five genera of the family Axinellidae, including three new species, Axinella loribellae sp. nov. A. sinoxea sp. nov. and Phakellia tropicalis sp. nov., are recorded for the tropical northern Australian waters of Western Australia, the Northern Territory and the Queensland coast as part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. One species, Dragmacidon durissimum (Dendy, 1905), generally found in the Indian Ocean, represents a new record for Australia. Taxonomic descriptions and discussion of those species are presented here. The position of Reniochalina within the Axinellidae is also discussed based on new evidence found in this and other studies.

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2009 25: 17–42 Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) from northern Australia. Family Axinellidae BELINDA ALVAREZ1 and JOHN N.A. HOOPER2 1 Museum and Art Gallery 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 johnh@qm.qld.gov.au ABSTRACT Nine species in ive genera of the family Axinellidae, including three new species, Axinella loribellae sp. nov. A. sinoxea sp. nov. and Phakellia tropicalis sp. nov., are recorded for the tropical northern Australian waters of Western Australia, the Northern Territory and the Queensland coast as part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. One species, Dragmacidon durissimum (Dendy, 1905), generally found in the Indian Ocean, represents a new record for Australia. Taxonomic descriptions and discussion of those species are presented here. The position of Reniochalina within the Axinellidae is also discussed based on new evidence found in this and other studies. Keywords: Sponge, Porifera, Halichondrida, Axinellidae, northern Australia, new species, taxonomy. Taxonomic knowledge of northern Australian sponges is limited to a few studies. The irst sponges collected from this area were described by Ridley (1884) and included 24 species, of which only 17 are currently recognised as valid species. Bergquist and Tizard (1967) later described 19 species from the rich intertidal area of Darwin Harbour. Since 1967, there have been 50 additional records to the fauna of northern Australia and only one revision of a particular group, i.e. the family Halichondriidae (Hooper et al. 1997 and references within ). Recent descriptions of some species have also been included in major taxonomic revisions of the demosponge families Raspailiidae (Hooper 1991) and Microcionidae (Hooper 1996). The order Halichondrida is presently represented in northern Australia by 41 nominal species (Hooper and Wiedenmayer 1994; Hooper et al. 1997), and a large number of specimens recently collected and deposited in the Museum and Art Gallery Northern Territory and the Queensland Museum (see abbreviations below). Data gathered from these collections clearly indicate that new species and records are represented in the area, and that species previously recorded also need to be revised using more sophisticated taxonomical tools. The Halichondrida is a group with an uncertain classiication and deinition. As with many other sponge groups, it is deined by traditional morphological characters, such as growth form, surface characteristics and skeletal features. But in the Halichondrida, however, these characters are extremely simple, polymorphic and few, and as a consequence the discrimination of taxa within this group is ambiguous. Halichondrid sponges have diverse growth forms (e.g. encrusting, massive, ramose, tubular, labellate). INTRODUCTION The northern marine region of Australia, or the Northern Province as deined by the Interim Marine and Coastal Regionalisation of Australia (IMCRA, version 3.3, www.environment.gov.au/coasts/mpa/imcra/index. html) includes tropical waters off the Northern Territory (from the Admiralty Gulf in the west) and the Queensland coasts (western coast of Cape York to Torres Strait in the east). The continental shelf of this area is generally shallow (less than 70 m) and extensive, reaching approximately 400 km in width in the Timor Sea and adjoining the coast of New Guinea in the Arafura Sea and Torres Strait (Bunt 1987; Ferns 1999). The area is part of the central Indo-West Paciic, which is well known for its high species-richness, high levels of endemism and is considered to be centre of origin of many tropical marine species (Veron 1995). Sponges are one of the most diverse and prevalent groups of marine invertebrates of northern Australia, but also one of the most poorly known in terms of proportions of known and new species, and levels of endemism. According to Hooper et al. (1997) the northern sponge fauna includes approximately 800 species, 60% of which remain undescribed. Further studies based on ‘presence-absence’ analyses of the diversity of tropical Australian sponges (Hooper et al. 2002) identiied at least two ‘hot spots’ of biodiversity for the northern area, one in the region of Darwin and Cobourg Peninsula and the other in the Wessel Islands region. Only 30% of the sponge species included in that biodiversity study could be assigned to a known taxon indicating that a great percentage of the fauna of that region is not well known. 17 B. Alvarez and J. N. A. Hooper The skeletons are plumoreticulate, dendritic or confused, constructed with three types of spicules (strongyles, styles and oxeas), or transitional forms, in any combination and not functionally localised. The order includes ive families (Axinellidae, Dictyonellidae, Heteroxyidae, Halichondriidae and Bubaridae), and 45 genera, most of which remain poorly deined despite recent efforts to clarify and redeined the taxonomy of these families (Alvarez and Hooper 2002; Alvarez and Van Soest 2002; Hooper 2002a; Van Soest et al. 2002; Van Soest and Hooper 2002). Evidence from molecular studies (Alvarez et al. 2000) indicates also that some of the genera are not monophyletic. Moreover, species allocated to some genera (e.g. Axinella, Acanthella, Phakellia) have fuzzy boundaries and overlapping characters and include numerous forms (or varieties/morphs). Similarly, some allegedly widely distributed species may represent complexes of cryptic species hiding under morphotypes that span a continuum, and which cannot be resolved easily using morphometric data alone. The taxonomic confusion around the Halichondrida, has generated long-lasting debates at higher levels of sponge classiication. Further studies using larger groups of species, revisions at the regional level and different kinds of genetic and chemical approaches have been recommended to reine the current concept of this taxon (Van Soest and Hooper 2002). The aim of this study is to revise the fauna of the Halichondrida from northern Australia and the status of all the nominal halichondrid species in this region. The present paper represents the irst part of this revision and includes the family Axinellidae. Revision of the remaining families represented in the area (i.e. Dictyonellidae, Halichondriidae and Heteroxyidae) will follow in separate papers. 0˚ 10˚ 20˚ S 120˚ 140˚ E Fig. 1. Study area included in this taxonomic revision. Barrier Reef were also examined if the species distribution was included in the studied area. Complete locality and collection data of material included in this revision is indicated under the species description as usual, whereas non type voucher material deposited at the Queensland Museum and the Museum and Art Gallery Northern Territory, is listed in Appendix 1. Specimens were prepared for light microscopy using the usual methods (e.g. Hooper 1996; Van Soest and Hooper 2005). Spicule measurements are in micrometres, based on 25 spicules (otherwise indicated in brackets), of each category and denoted as range (and mean ± 1 S.E.) of spicule length x spicule width. Measurements were made using a digital video camera attached to a light microscope in combination with the software V++ Precision Digital Imaging System v 4.0 (© Digital Optics Ltd). Scanning Electron Microscope photographs were taken in a JEOL JSM 5610LV. The higher systematic arrangement follows classification in the current version of World Porifera Database (Van Soest et al. 2008). Terminology used here follows Boury-Esnault and Rützler (1997) and Alvarez and Hooper (2002). MATERIALS AND METHODS This revision includes material of the family Axinellidae recorded for the tropical northern Australian waters of the Western Australia, 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 meridians (Fig. 1). The area does not represent a true biogeographical area and it was delimited based on the marine bioregions deined by IMCRA. This area also corresponds with two of the ecoregions (i.e. Arafura Sea, Arnhem Coast to Gulf of Carpenteria) of the Sahul Shelf marine province as deined by Spalding et al. (2007). All specimens recorded for the selected area and registered under Axinellidae at the Queensland Museum and the Museum and Art Gallery Northern Territory sponge collections were examined and identiications were veriied. Specimens and relevant type material from adjacent areas including western Australia, the Ashmore, Cartier and Hibernia reefs on the Sahul Shelf, Lesser Sunda Islands, Aru Islands, the south coast of Papua New Guinea and Great 130˚ ABBREVIATIONS 18 Abbreviations used in the paper are: AIMS, Australian Institute of Marine Sciences; BMNH, Natural History Museum, London (formerly British Museum Natural History); CRRF, Coral Reef Research Foundation, Palau; GBR, Great Barrier Reef, NTM, Museum and Art Gallery Northern Territory, Darwin, Australia (formerly Northern Territory Museum); MONZ, Museum of New Zealand; NTM, Museum and Art Gallery Northern Territory (formerly Northern Territory Museum), Darwin; SMF, Senckenberg Research Institute and Natural History Museum, Frankfurt; QLD, Queensland, Australia; QM, Queensland Museum, Brisbane; WA, Western Australia, Axinellidae from northern Australia Australia; ZMA, Zoologisch Museum, University of Amsterdam, Amsterdam. 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’ program, by the Australian Institute of Marine Sciences, CRRF and NTM (subcontracted through CRRF), respectively. Dragmacidon durissimum (Dendy, 1905) Phakellia tropicalis sp. nov. Reniochalina stalagmitis Lendenfeld, 1888 Genus Axinella Schmidt, 1862 Gender feminine. Type species, by subsequent designation of De Laubenfels (1936), Axinella polypoides Schmidt, 1862. Recent, Adriatic Sea. Axinella aruensis (Hentschel, 1912) (Figs 2 A–F, 3, 4, Table 1) Phakellia aruensis Hentschel, 1912: 420; Hooper et al. 1992 [in part]; Pulitzer-Finali 1993: 283. Axinella aruensis. – Hooper and Wiedenmayer 1994:72; Alvarez et al. 2000 [form II, see below]; Alvarez, Krishan and Gibb 2007[form II]; Holmes and Blanch 2007. Material examined. The material examined for this species is separated according to the morphotypes described below. Holotype – SMF 953, E side, Aru I., Indonesia, 31 August 1908, coll. Merton, H.. AdditionAl specimens – Lacepede Is, NW Shelf, WA: NTM Z.2284, Z.2304, Z.2331, Z.2345. Joseph Bonaparte Gulf: QM G301197, Cartier I.: QM G301092. Melville I. NT: NTM Z.615, TAXONOMY Order Halichondrida Gray, 1867 Family Axinellidae Carter, 1875 Nine species of Axinellidae, listed below, were recorded within the studied area; three of these being new species. Axinella aruensis (Hentschel, 1912) Axinella loribellae sp. nov Axinella sinoxea sp. nov Cymbastela stipitata (Bergquist and Tizard, 1967) Cymbastela vespertina Hooper and Bergquist, 1992 Dragmacidon australe (Bergquist, 1970) Table 1. Comparison of spicule dimensions among specimens and varieties of Axinella aruensis. Measurements in micrometres. Specimen Locality Axinella aruensis SMF 953 Aru Is, Indonesia Z.2304 Lacepede Is, WA G301092 Cartier Is, WA Z.619 Melville I., NT Z.3141 Parry Shoals, NT Z.5053 Darwin Harbour, NT Z.4465 Wessel Is, NT Oxeas Styles 257.1–423.9 (360.6±38.1) x 13.8–21.4 (16.5±1.9) 245.5–337.6 (285.4±21) x 8.6–17.2 (13±2.6) 281.2–450.2 (360.7±39.9) x 10.3–19.4 (15.6±2.2) 236.1–406 (302.4±39.4) x 9.3–17.5 (13.9±2.3) 267.1–372.9 (307.1±23.3) x 9.4–17.2 (13.6±2.3) 297.6–498.6 (392.1±47) x 7.3–22.5 (16.5±3.7) 194.4–396.4 (299.5±48.6) [24] x 5.6–17.4 (12±3.3) 249.1–382.2 (313.6±40.7) [12] x 14.1–21.7 (17.4±2.2) [12] 213.2–271.4 (244.8±20.9) [9] x 11.9–15.7 (13.5±1.6) [9] 242.8–419 (301.1±36) x 12.6–20.3 (16.8±1.9) 186–362.8 (267.2±43.3) [22] x 9.8–17.1 (14.1±1.9) [22] 248.6–294.6 (270.4±18.8) [4] x 11.1–16.7 (13.9±2.3) [4] 263.6–417.2 (342.7±37.6) x 12–23.5 (17.1±2.9) 204.6–331.7 (269.7±44.1) [15] x 8.6–18 (13.8±2.8) [15] 305–451.7 (376.7±36.3) x 13.1–25.9 (18.3±3.9) 187.1–318.2 (237.2±26.6) x 8.5–17.2 (13.2±2.4) 266.7–354.5 (312.1±23.3) x 13.1–18.8 (16.3±1.4) 274.8–392.8 (333±32.8) x 8.3–16.4 (13.3±2.3) 283.1–406.6 (334.7±40.7) [6] x 14.7–19.6 (16.5±1.8) [6] 167.2–222.5 (200±15.3) x 9.3–15.5 (13.2±1.7) 200.1–353.2 (260.7±34.1) x 10.3–22.8 (15.4±2.9) 248.6–363.3 (297±31) [10] x 11.5–18 (15.1±2.3) [10] 209.1–278.6 (246.3±15.8) x 12.7–19.9 (15.1±1.8) 173.8–247.4 (214.7±18.9) x 7.5–16.7 (10.3±1.9) 163.6–231.4 (191.1±17.4) x 6.5–12.5 (9.8±1.6) 166.4–262.3 (218.2±22.4) x 7.8–17.9 (13.9±2.4) Axinella aruensis form I Z.5816 Bynoe Harbour, NT Z.3068 Parry Shoals, NT Z.5819 East Point, Darwin, NT Z.3946 Wessel Is, NT Axinella aruensis form II Z.4490 Stevens Rock, Darwin Z.5054 Wessel Is 19 B. Alvarez and J. N. A. Hooper A C B D F E G H Fig. 2. Axinella aruensis: A, B, specimens at Raragala I., Wessel Is; C, specimen at South Shell I., Darwin Harbour; D, form I, Z.5816, Dawson Rock, Bynoe Harbour; E, form II, specimen at Raragala I, Wessel Is; F, Axinella sinoxea sp.nov., NTM Z.2719. Axinella loribellae sp. nov.; G, Holotype, NTM Z.4427; H, NTM Z.5662. Photos: A–B, P. Colin; C–D, G, B. Alvarez; E, D. DeMaria; F, J. Hooper; H, A. Ayling. 20 Axinellidae from northern Australia Z.619, Z.630, Z.632. Parry Shoals, Arafura sea, NT: QM G310136 (Q66C0514-X), NTM Z.3062 (Q66C87-0514-X), Z.3141.Bynoe Harbour, NT: NTM Z.5071 (0M9H2464-U). Darwin Harbour, NT: NTM Z.5053 (0M9H2168-X), Z.5057 (0M9H2665-O), Z.5058 (0M9H2675-Y), Z.5072 (0M9H2579-U), Z.5830. Cobourg Peninsula, NT: NTM Z.1363, Z.1388, Z.2511, Z.2526, Z.2529. English Company Is., NT: NTM Z.3956. Wessel Is, Gove Peninsula, NT: QM G3.609 (Q66C4762-R), G300768 (=Q66C4737P, QM G311873 and NTM Z.3945), Z.3922 (Q66C4687-L), Z.3935 (=Q66C4785-R, QM G300752), Z.3936 (Q66C4831-R), Z.4465 (0M9H2770-C), Z.5055 (0M9H2650-W). Papua New Guinea: QM G312913, G312935. Axinella aruensis, form I, Bynoe Harbour, NT: NTM Z.5816, Z.5817, Z.5818. Darwin Harbour, NT: NTM Z.2156, Z.5819-Z.5823. Wessel Is, Gove Peninsula, NT: NTM Z.3925, Z.3946. Axinella aruensis, form II. Darwin Harbour, NT: QM G303332, Z.1961, Z.2249, Z.2402, Z.2632, Z.4425 (0M9H2044-O), Z.4490, Z.4491, Z.5824-Z.5829, Z.5831, Z.5232. Parry Shoals, Arafura sea, NT: NTM Z.3137, Z.3068. Wessel Is, Gove Peninsula, NT: QM G300759 (Q66C-4831-R), NTM Z.5054 (0M9H2648-U). Yampy sound, WA, NTM Z.665. Description. Three different morphotypes of this species with one corresponding to the holotype are recognisable among the material examined and they will be described below separately under the heading of ‘forms’. Axinella aruensis, typical form. Shape (Fig. 2A–C). Thickly labellated, on broad and short, or long and narrow, peduncle, uni or bi-planar, sometimes folded, with round margins projecting in most cases into short and broad extensions with square, round or pointed tips, or in long rounded to flat branches which tend to fuse laterally. Specimens up to 400 mm high. Colour. Orange, pale yellow or yellowish brown alive. Dark brown in alcohol. Oscula. Regularly distributed in one or both sides of fan, stellate, lush or with elevated rims, less than 5 mm diameter. Surface. Evenly microconulose-conulose, nodulose, rough, marked with primary longitudinal choanosomal ibres. Skeleton (Fig. 3A). Plumose, vaguely reticulated to halichondroid, very compact, with plumose columns up to 600 µm thick, diverging toward surface, ending in fan-shapped spicule brushes and projecting through ectosome. Axial skeleton differentiated only towards base of attachment or peduncle, halichondroid. Spicules (Fig. 3B). Oxeas with blunt, pointed or telescoped tips; slightly bent and sometimes slightly sinuous, 195–498 x 5–22 µm. Styles, less frequent or rare, similar in size to oxeas (see Table 1) with blunt ends, enlarged or slightly narrow bases, straight or slightly bent. Transitional forms (e.g. styloids, strongyles) are common. A B C D E F Fig. 3. Axinella aruensis: light microphotograph of skeleton and diagram of spicules: A, B, SMF 953, holotype; C, D, Z.5819 (form I); E, F, NTM Z.5054 (form II). Scale bars: A, 200 µm; B, D, F, 50 µm; C, E, 500 µm. Axinella aruensis, form I. Shape (Fig. 2D). Erect, fan-shaped or narrow long and lat digits with few simple ramiications, generally with square margins. Specimens up to 20 cm high and 10 cm wide. Colour. Light orange, brown, beige or yellow. Same colour in alcohol. Oscula. Regularly distributed, less than 5 mm diameter, with distinctive raised margins. Surface. Minutely hispid, marked with choanosomal skeletal tracts in a regular reticulation or with radial grooves. Skeleton (Fig. 3C). Plumose and slightly compressed at axial region, with thick plumo-echinated multispicular columns, up to 1 mm thick, and radiating outwards towards surface, anastomosing or connected irregularly by short and thick paucispicular or multispicular tracts, or by single spicules oriented in any direction. Main tracts end at surface in fan-shaped brushes with spicules projecting shortly through ectosome; light spongin embedding tracts. Spicules (Fig. 3D, Table 1). Oxeas, 187–451 x 8–25 µm, with pointed or blunt ends, straight or bent; thinner forms are common. Styles less frequent or rare, slightly smaller, 21 B. Alvarez and J. N. A. Hooper including intermediate forms similar to styloids, anisoxeas or strongyles. Fused spicules are characteristically common. Axinella aruensis, form II. Shape (Fig. 2E). Thick fans or lamellae with round margins, folding in more than one perpendicular plane, or joining at angles from 45-90 degrees; or short single or digitate projections, stipitate, on short narrow peduncles or on broad base. Generally small with individuals reaching up to 13 cm high. Colour. Bright or light orange alive. Light beige in alcohol. Oscula. Stellate with minute drainage canals, sometimes located at margin of fans or evenly distributed in both sides of fan, 3-5 mm in diameter. Surface. Pierced uniformly with minute ostia. Microconulose; minutely hispid, firm but some has mucous consistency after collection. Marked by regular choanosomal reticulation. Skeleton (Fig. 3E). Thick and dense plumo-echinated multispicular tracts up to 600 µm wide, forming regular, nearly radial reticulation which is marked on surface. Main columns end at surface in fan-shaped brushes, with spicules projecting shortly through ectosome. Axial skeleton not differentiated. Spicules (Fig. 3F, Table 1). Oxeas and styles in nearly equal proportions, 174-279 x 7-20µm. Styles are dominant and slightly smaller than oxeas. Remarks. Although some features allowed distinction of two additional morphotypes within this species, the limits among them are not clear and some individuals could be considered intermediate forms. Axinella aruensis sensu stricto is distinguished from its two other forms by shape and colour, generally observed to be thickly labellate and orange when alive, by the change of colour in alcohol (it turns brown) and by the dominance of oxeas in relation to styles. Form I differs slightly in shape from A. aruensis; the colour in life is always beige or pale yellow and does not change in alcohol; oxeas are also dominant and styles are relatively more common. Form II is always beige in alcohol and styles are dominant relative to oxeas. Some other features of shape and surface consistency and texture are also distinctive within this form. Some data included in Hooper et al. (1992) indicate there are some differences in the biochemistry between populations (i.e. North West Shelf versus Darwin Harbour) of this species, however the published results do not seem to be related to the forms distinguished here. Alvarez et al. (2007) detected up to 29% of intra-genomic polymorphism within the Internal Transcribe Spacer (ITS) of the rDNA in individuals of Axinella aruensis from Darwin Harbour. These levels of intra-genomic variation are so far the highest reported for Porifera and correspond in most cases to hybrid species reported for other groups, including corals of the genus Acropora. Thus, it is possible that the forms here distinguished to document the variability present within species are the result of a hybridisation processes with sympatric species or populations. Future population genetic studies will help to determine whether the variability observed across these forms are signiicant to justify their recognition as different species or as hybrids. Axinella aruensis is very similar in shape, skeletal architecture and spicule composition and dimensions to some Axinella species recorded from the Indian Ocean (e.g. A. donnani (Bowerbank, 1873); A. manus Dendy, 1905 and A. symmetrica (Dendy, 1905, as Phakellia)). Skeletal reticulation of the Indian Ocean species, in particular A. donnani, is much more regular, with thicker primary lines. Interesting also is the change in colouration, from orange to brown after few hours of collection, reported for A. donnani (Bowerbank, 1873), a characteristic also seen in A. aruensis. A detailed revision of the Indian Ocean species complemented with population genetic studies is essential to deine their limits and phylogenetic relationships with the northern Australian populations of A. aruensis and its forms. Distribution. Axinella aruensis and its forms appear to occur sympatrically and are common throughout northern Australia (Fig. 4). The type locality is the Aru Is, Indonesia, but the species is also known from other Indonesian localities (Alvarez and de Voogd, unpublished data) and from Papua New Guinea. It is found in subtidal areas from 5 to 76 m. Pulitzer-Finali’s (1993) record for East Africa, is dubious and requires conirmation. Axinella loribellae sp. nov (Figs 2G–H; 5) Material examined. H olotype – NTM Z.4427 (0M9H2041-L), Stevens Rock, Weed Reef, Darwin Harbour, 12°29.2001´S, 130°47.1´E, NT, 5–19 m depth, 8 May 2002, coll. B. Alvarez and party. pArAtypes – ntm Z.5834 Stevens Rock, Weed Reef, Darwin Harbour, Northern Territory, Australia, 12°29.1667´S, 130°47.19´E, 17 m depth, 8 May 2006, coll. B. Alvarez. Additional specimens. Melville I., NT, NTM Z.631. Darwin Harbour, NT: QM G303388, NTM Z.822, Z.868, Z.5662. Wessel Is, NT: NTM Z.3938, Z.5059 (0M9H2771-F). 120˚ E 128˚ 136˚ E 8˚S 16˚ S Fig. 4. Distribution of Axinella aruensis (open circles) and its forms (I, black circles; II, grey circles) based on conirmed records from QM and NTM. 22 Axinellidae from northern Australia A tracts oriented perpendicular to surface and laterally close, connected by single spicules or uni-paucispicular tracts up to 2 spicules long, cemented with thin and clear spongin, to each other and ending in brushes of spicules that protrude shortly through ectosome. Spicules (Fig. 5C; Table 2). Styles slightly bent; strongyles straight, slightly sinuous, bent in middle; oxeas fusiform, straight. Transitional shapes between monoactins and diactins are common. All types in a wide range of sizes and thickness. Dominant types vary among specimens (see below). Remarks. A great variability in the shape and size of spicules was observed among specimens of this species. Styles and strongyles of 190–270 by 7–13 µm, are the dominant types, however they were absent in some of the examined specimens (i.e. NTM Z.631 and NTM Z.3938). Apart from these differences in spicule composition and dimensions, no other characters seem to vary among specimens examined here, and consequently they are considered at this stage to belong to a single species. Further genetic studies may help to conirm whether or not variability in spicule dimensions and composition is indicative of sibling species differentiation. The new species differs from Axinella aruensis and its related species from the Indian Ocean (see above) mainly in shape (thinly instead of thickly labellate); in skeletal architecture (clearly differentiated into axial and extraaxial skeleton with thinner extra-axial spicular tracts and more regular reticulation when compared to the thicker multispicular columns present in A. aruensis) and in spicule composition (with common transitional shapes between oxeas and styles not observed in A. aruensis). Axinella ceylonensis (Dendy, 1905, as Phakellia) from the Gulf of Manaar is similar to A. loribellae in shape (although shortly stipitate) and in thickness of the lamellae. Both species share the variability observed in the size and shape of oxeas and styles. The two species differ in skeletal and surface characteristics. The skeleton of A. ceylonensis is not differentiated in axial and extra-axial region but described as plumose, with columns radiating outwards into small surface conules. No other species in the study area or in the Indian Ocean is similar to this new species. Indeed, most nominal species of Axinella reported for the Indian Ocean, do not agree with the current diagnosis of the genus and need to be re-examined. Some of these are massive forms with styles and trichodragmata in the skeleton and might belong in the axinellid genus Dragmacidon (e.g. Axinella C B Fig. 5. Axinella loribellae sp. nov.: A, Paratype, NTM Z.5834; B, light microphotograph of skeleton; C, diagram of spicules. Scale bars: A, 2 cm; B, 100 µm; C, 50 µm. Description. Shape (Figs 2G–H, 5A). Fan-shaped; thin lamellae, 1–5 mm thick, single or bifurcated, sometimes convoluted with rounded margins and indentations on short stalks or broad base. Specimens are 300 mm high and up to 400 mm wide. Colour. Burnt orange alive, brown in alcohol. Oscula. Small, 2–5 mm diameter, with stellate drainage canals, evenly distributed. Consistency. Flexible, easy to tear, rubbery. Surface. Smooth, velvety, marked irregularly with ribs. Skeleton (Fig. 5B). Plumose, differentiated in extraaxial and axial region. Axial skeleton compressed with wavy longitudinal paucispicular-multispicular tracts, lightly embedded in collagenous spongin, interwoven, and radiating towards the extra-axial region. Extra-axial region a close-set reticulation of plumose paucispicular Table 2. Comparison of spicule dimensions among specimens of Axinella loribellae sp. nov. Measurements in micrometres. Specimen Z.4427 (Holotype) Locality Darwin Harbour Z.5059 Wessel Is Z.631 Melville Is Styles 196.3–352.9 (274.6±47.7) x 8.33–18.3 (13.5±2.7) 159.7–365.4 (250.6±50.4) x 5.8–14.43 (10.0±2.1) 187.6–307.9 (228.9±29.0) x 4.3–11.3 (7.3±1.8) 23 Strongyles 103.6–396.3 (190.1±74.0) x 6.01–13.09 (10.3±1.9) 93.7–531.8 (243.1±128.6) x 5.8–13.3 (8.8±2.3) – – Oxeas 148.5–440.2 (226.8±63.6) x 4.8–12.9 (8.7±2.1) 100.8–302.2 (208.5±50.1) x 3.7–11.4 (6.5±2.1) 142.8–351.6 (217.8±44.1) x 3.5–13.2 (7.1±2.7) B. Alvarez and J. N. A. Hooper surface; slightly compressed in axial region. Spicule tracts bound only slightly with clear collagenous spongin. Spicules (Fig. 6D; Table 3). Styles robust, bent, or less often, straight, enlarged in the middle section, 159–245 x 7–17 µm; thinner category, 97–201 x 2–6 µm also present. Long thin raphids abundant. Smaller oxeas and thick and short strongyles, very rare. Remarks. This species conforms in most of its characteristics with the current concept of Axinella. The absence or low frequencies of oxeas observed in the examined specimens are also seen in other Caribbean species of the genus (e.g. Axinella waltonsmithi (de Laubenfels, 1953) and A. pomponiae Alvarez, Van Soest and Rützler, 1998), which might be considered a common feature among Axinella species. The new species resembles Axinella aruensis in gross morphology and as such can be easily mistaken for it in the ield; both are fan-shaped on a common stalk and both have a similar surface pierced with minute ostia and microconulose. But A. sinoxea is clearly different from A. aruensis in skeletal architecture and spicule composition, having a regular plumoreticulated skeleton of ascending tracts and long thin raphids in the skeleton. As is the case with A. loribellae, no other species recorded in the study area or in the Indian Ocean was found to be related to A. sinoxea. Distribution. Common in the vicinity of East Point Sponge Gardens, Darwin Harbour, but also found in deeper waters (down to 40 m) of Western Australia. Etymology. Latin, sine- without; sinoxea referring to the lack of proper oxeas characteristic of the species. It is intended as a noun in apposition. Remarks on Axinella. Axinella is a widespread genus of sponges with approximately 100 accepted species (Van Soest et al. 2008), many of which, however, need to be veriied against the current deinition of the genus (Alvarez and Hooper 2002). Ongoing revisions of species of Axinella and related genera by one of the authors (BA) are undertaken on a regional basis with the purpose to verify the identity of the reported species and the monophyly of the genus, which is currently proven as polyphyletic based on molecular studies (Alvarez et al. 2000; Erpenbeck et al. 2005). Three species of Axinella (A. aruensis, A. loribellae, A. sinoxea) are reported in this work. No other species of the genus, as far as we know, have been reported within the area of northern Australia that is the subject of this present study. Axinella echidnaea reported by Ridley 1884 is accepted as Reniochalina stalagmitis (see below). bidderi Burton, 1959 and A. massalis Burton, 1959). Some others are Stylissa-like, or other dictyonellid genera, with the surface marked by ridges or conules and with a dense and irregular skeleton of multispicular tracts of styles (e.g. Axinella bubarinoides Dendy, 1922; A. dragmaxioides Burton, 1959 [?]; A. labelloreticulata (Burton, 1959); A. labyrinthica Dendy, 1889; A. minor Thomas, 1981; A. proliferans Ridley, 1884; A. tenuidigitata Dendy, 1905; A. ventilabrum Burton, 1959), or closer to halichondrid genera (e.g. Axinella halichondrioides Dendy, 1905, which is encrusting and has only oxeas) or to the raspailiid genus Ceratopsion (i.e. Axinella lamellata Dendy, 1905, with a dermal and tangential layer of small oxeas). Distribution. Axinella loribellae seems to be restricted to northern Australia between Darwin Harbour and the Wessel Is. It is found between 11–32 m depth. Etymology. Named after Lori Bell, Coral Reef Research Foundation, Palau, for her considerable contribution to the knowledge of Indo-Paciic sponge diversity and distribution. It is intended as a noun in apposition. Axinella sinoxea sp. nov. (Figs 2F, 6A–D) Material examined. Holotype – NTM Z.940, East Point, Darwin Harbour, NT, 12°24.05´S,130°48.01´E, 12 m depth, 13 September 1982, coll. Hooper, J.N.A. pArAtypes – Z.5833, East Point, Darwin Harbour, NT, 12º 24.484´S, 130º 48.471´E, 11 m depth, 7 June 2007, coll. B. Alvarez. Additional specimens. NW Shelf, WA: NTM Z.2310, Z.2322. Darwin Harbour, NT: NTM Z.2246, Z.2719. Description. Shape (Figs 2F, 6A). Single or multiple fans, 4–6 mm thick, 8–14 cm long and up to 30 cm wide, on common stalk, 3–5 cm long and 7–10 mm in diameter; erect, uniplanar with digitate to irregular margins or bifurcate tips. Colour. Orange, pale yellow or beige with light pink tinge alive; brown-grey in alcohol. Oscula. Regularly distributed in one or both sides of fan, round to elongated or irregularly shaped, some stellate, with slightly elevated rims, less than 1 mm diameter. Consistency. Soft, floppy, flexible, slightly compressible. Surface. Smooth but slightly rough to touch; pierced regularly by minute pores, microhispid due to projections of brushes of choanosomal spicules. Encrusted irregularly with detritus in some specimens. Skeleton (Figs 6B-C). Plumoreticulated, with ascending multispicular tracts connected regularly by single spicules or unispicular tracts, 1 or 2 spicules long, ending in brushes at Table 3. Comparison of spicule dimensions among specimens of Axinella sinoxea sp. nov. Measurements in micrometres. Specimen Z.940 (Holotype) Locality East Point, NT Z.2310 NW Lacepede Is WA Thick styles Thin styles Raphids 184.5–245.1 (223.7±12.92) 97.77–201.3 (179.64±22.84) [23] 192.9–249.6 (227.2±14.9) 7.35–17.4 (13.11±2.49) x 2.48–5.6 (3.8±1) [23] x 0.8–3.0 (2.0±0.6) 153.4–197.6 (183.4±10.2) 131.3–191.2 (152.1±14.8) 131.9–258.6 (201.5±35.1) x 7.8–12.2 (10.2±1) x 2.2–6.4 (4.4±1.) x 0.37–2.69 (1.6±0.5) 24 Axinellidae from northern Australia A B C D Fig. 6. Axinella sinoxea sp. nov. NTM Z.940: A, photograph of the holotype; B, light microphotograph of the skeleton; C, SEM, raphids in choanosomal skeleton; D, scale bars: A, 5 cm; B, 500 µm; C, 100 µm; D, 20 µm. In the present study area, the genus Axinella seems to be less speciose than in other taxonomically revised regions: seven species in the Western Central Atlantic (Alvarez et al. 1998); six (recorded) species and nine possible new species (Kelly et al. 2009); at least ive species in Indonesia (Alvarez and De Voogd, unpublished data). Unfortunately little is known about the biology of these species to explain why the genus might be more diverse in some areas than others. Many more putative Axinella species are known for the GBR (pers. obs.), but these remain unresolved pending future studies. Distinction of Axinella species continues to be subjective and is based on a combination of characters as discussed by Alvarez et al (1998). Variability and plasticity of all the morphological characters that characterise the species are seen in all the species described above. The morphological variability of A. aruensis, for example, is remarkable and suggestions from molecular data (Alvarez et al. 2007) that it may be due to hybridisation should be further explored. Genus Cymbastela Hooper & Bergquist, 1992 Gender feminine. Type species, by original designation, Pseudaxinyssa stipitata Bergquist and Tizard, 1967. Recent, Darwin Harbour, Arafura Sea. 25 Cymbastela stipitata (Bergquist and Tizard, 1967) (Figs 7A–B) Pseudaxinyssa stipitata Bergquist and Tizard, 1967:189; Hooper et al. 1992: 265. Cymbastela stipitata. – Hooper and Bergquist 1992: 106; Hooper and Wiedenmayer 1994 : 75; Alvarez et al. 2000: 195; Alvarez and Hooper 2002: 733. Material examined. Specimens as listed in Hooper and Bergquist (1992). AdditionAl specimens – Bynoe Harbour, NT: Z.5065 (0M9H2333-C). Darwin Harbour, NT: QM G303262, NTM Z.4078 (0CDN8001-H, Fig. 7A), Z.4104 (0CDN8026-J), Z.4131, Z.4435 (0M9H2008-Y), Z.5064 (0M9H2134-M), Z.5835, Z.5836, Wessel Is, NT: Z.5066 (0M9H2658-H), Z.5067 (0M9H2785-T). Remarks. Cymbastella stipitata, was re-described extensively by Hooper and Bergquist (1992) and the type material re-examined by Alvarez and Hooper (2002). This is one of the most common sponges in the studied area. It is particularly abundant in the intertidal zone of Darwin Harbour, which becomes greatly exposed during the nocturnal king tides of the dry season (May–June) and the diurnal king tides during the wet season (SeptemberNovember, Fig. 7B). It is found, but less commonly, in subtidal areas down to 19 m depth. New records indicated that its distribution within northern Australia extends from B. Alvarez and J. N. A. Hooper A B C D E F H G Fig. 7. Cymbastela stipitata: A, NTM Z.4078 (0CDN-8001-H); B, specimens exposed at the reef lat of East Arm, Darwin Harbour during the low tide of 20 September 2001. Dragmacidon australe: C, specimen at Channel I. Darwin Harbour; D, QM G304246, Lizard I, GBR, QLD. Phakellia tropicalis sp. nov.; E, Holotype (NTM Z.5847); F, Paratype (NTM Z.5845). Reniochalina stalagmites: G, specimen at East Point, Darwin; H, specimen at Cotton I., Wessel Is. Photos: A, B, E, F, B. Alvarez; C, H. Nguyen; D, J. Hooper; G, A. Ayling; H, P. Colin. 26 Axinellidae from northern Australia 125˚E 135˚ Genus Dragmacidon Hallmann, 1917 Gender neuter. Type species, by original designation, Thrinacophora agariciformis Dendy, 1905. Recent, Gulf of Manaar, Indian Ocean. 145˚ 5˚ Dragmacidon australe (Bergquist, 1970) (Figs 7 C–D, 9 A–B) Pseudaxinella australis Bergquist, 1970: 20; Hooper and Lévi 1993: 1441; Hooper and Wiedenmayer 1994: 80; Alvarez et al. 2000: 196. Dragmacidon australe. – Alvarez and Hooper 2002: 735; Kelly et al. 2009 (In press). Material examined. Holotype – NMNZ Por. 26, Takatu Channel, Northland, New Zealand, 11 m. AdditionAl specimens – Cartier I, WA: QM G301089. Bynoe Harbour, NT: G303444. Darwin Harbour, NT: NTM Z.5068. Coral Sea, GBR, QLD: QM G300295, G304182, G304246, G304253, G320664, NTM Z.2727. Description. Shape (Fig. 7 C–D). Thickly encrusting, following substrate, globular, bulbous or semispherical, approx. 100 mm in diameter by 20 mm thick. Colour. Bright red, orange alive. Consistency. Slightly compressible or stiff. Mucous surface. Oscula. Irregularly distributed, less than 1 mm diameter, with slightly elevated rims surrounded by thin drainage channels in stellate arrangement. Surface. Highly conulose; evenly pierced by pores 120–400 µm in diameter. Conules, approx. 1–3 mm long, single or grouped in reticulated pattern, unevenly echinated by spicules, 2–3 mm apart. Skeleton (Fig. 9A). Plumoreticulate to halichondroid; formed by thick plumose or plumo-echinated multispicular tracts, up to 500 µm thick, ascending nearly perpendicularly from base and becoming thicker and bushy near surface; projecting through ectosome into surface conules. Main tracts connected by shorter and thinner plumose tracts, sometimes ill-deined, forming irregular reticulation of large round meshes. Spicules (Fig. 9B; Table 4). Oxeas and styles in equal proportions, 176–510 x 7–21 µm (Table 4). Remarks. The species was originally assigned to Pseudaxinella and transferred to Dragmacidon by Alvarez 15˚ S Fig. 8. Distribution of Cymbastela stipitata, based on conirmed records from QM and NTM. Bynoe Harbour to the Wessel Is (Fig. 8), inferring it is a narrow range endemic within northern Australia. Cymbastela vespertina Hooper and Bergquist, 1992 Cymbastela vespertina Hooper and Bergquist, 1992: 110; Hooper and Wiedenmayer 1994: 75; Alvarez et al. 2000: 195; Alvarez and Hooper 2002: 733. Pseudaxiyssa sp. nov. Hooper et al. 1992: 265. Material examined. Specimens as listed in Hooper and Bergquist (1992). Remarks. Cymbastela vespertina is a sibling species of C. stipitata. Separation of the two species based in morphology is dificult and very subjective. Biochemical and molecular evidence (Hooper et al. 1992; Alvarez et al. 2000) indicates the sympatric populations are heterogeneous. Future genetic population studies might reveal whether or not these populations can be reliably separated into different species, but based on current external gross morphological differences and skeletal characters the two taxa are maintained as distinct. Remarks on Cymbastela. Two species of Cymbastela are present within the area of the present study (i.e. C. stipitata (Bergquist and Tizard, 1967) and C. vespertina Hooper and Bergquist, 1992). Other species of Cymbastela represented in other regions including Australia are: C. cantharella (Lévi, 1983), New Caledonia; C. concentrica (Lendenfeld, 1887), Queensland coast; C. coralliophila Hooper and Bergquist, 1992, GBR; C. marshae Hooper & Bergquist, 1992, Houman-Abrolhos, WA; C. notiaina Hooper and Bergquist, 1992, South Australia; and C. tricalyciformis (Bergquist, 1970) from New Zealand. Comprehensive descriptions of these species are given by the respective authors of the species. Phylogenetic relationships of the genus with other axinellid species based on molecular characters indicate that the northern Australian species of Cymbastela are closely related to other members of Dictyonellidae, such as Acanthella (Alvarez et al. 2000; Erpenbeck et al. 2005). There is no doubt, however, that species of Cymbastela are related to the Axinellidae based on their morphology. Thus the phylogenetic relationships derived from these molecular analyses remain enigmatic at this stage. A B Fig. 9. Dragmacidon australe: A, light microphotograph of skeleton; B, diagram of spicules. Scale bars: A, 500 µm; B, 50 µm. 27 B. Alvarez and J. N. A. Hooper Table 4. Comparison of spicule dimensions among specimens of Dragmacidon australe. Measurements in micrometres. Specimen Locality Styles Oxeas G303444 Bynoe Harbour 254.1–510.4 (349.1±78.8) x 7.3–20.9 (13.6±3) 286.2–434.5 (361.5±39.8) x 7.3–17.9 (13.4±2.4) Z.5068 Darwin Harbour 176.4–397.3 (290.1±61.4) x 7–17.2 (13±2.8) 271.9–412.5 (340.7±31.9) x 9.5–18.9 (14.3±2.4) and Hooper (2002) because it conformed more closely with the type species of that genus. Hooper and Lévi (1993) compared specimens from the GBR with the holotype of Dragmacidon australe from New Zealand and with material from New Caledonia described as D. debitusae (Hooper and Lévi, 1993). Very subtle differences were found between the two species. The material from northern Australia reported here agrees with D. australe in the majority of its features and is therefore assigned to this species. This species is also very similar to D. reticulatum (Ridley and Dendy, 1886) from the central West Atlantic both in external morphology and spicule composition. Distribution. Dragmacidon australe was irst recorded for New Zealand and additional records from the GBR were reported in Hooper and Lévi (1993). The present revision extends the distribution range of this species into northern Australia. The species is not very common in this region with only isolated records registered through the extension of the studied area (Fig. 10) and thus is probably at the edge of its range. It is also found along more temperate areas of the Queensland coast (Hooper pers. obs.). 125˚E 135˚ 145˚ 5˚ 15˚ S Fig. 10. Distribution of Dragmacidon australe in northern Australia, based on conirmed records from QM and NTM. Species distribution extends along the Queensland coast (Hooper pers. obs.) and New Zealand (type locality). A C Dragmacidon durissimum (Dendy, 1905) (Figs 11 A–C) Thrinacophora durissima Dendy, 1905:187. Sigmaxinella durissima. – Dendy 1922: 113. Axinella durissima. – Burton 1959: 259. Pseudaxinella durissima. – Alvarez et al. 2000: 196. Dragmacidon durissima. – Hallmann 1917: 639; Alvarez and Hooper 2002: 735. Material examined. Ashmore Reef, WA: QM G300181. Description. Shape (Fig. 11A). Hemispherical, cushionshaped. Colour. Orange alive, red on deck, beige in ethanol. Oscula. Round, irregularly distributed at top, with slightly elevated rims. Surface. Very rugose, composed of minute projections or conules, compact and close-knit; membranous skin stretched over conules. Skeleton (Fig. 11B). Plumoreticulate. Multispicular, plumose or plumoechinated spicule tracts, ascending toward surface and connected by shorter and thinner ones, or loose spicules, forming irregular reticulation of oval to square meshes; projecting through ectosome in surface conules or projections. B Fig. 11. Dragmacidon durissimum: A, QM G300181; B, light microphotograph of skeleton; C, diagram of spicules. Scale bars: A, 1 cm; B, 500 µm, C, 50 µm. 28 Axinellidae from northern Australia Spicules (Fig. 11C). Styles 203.1–312.5 µm (251.8±33.4) by 11.7–16.4 µm (13.7±1.2). Oxeas in equal proportions, 229.8–312.7 µm (283.2±18) by 7.4–19 µm (13.4±3.1). Trichodragmata short and thick, 15–20 by 5–10 µm. Distribution. Indian Ocean, including Seychelles Is, Amirante, Providence, Saya de Malha (Dendy 1905; Dendy 1922; Burton 1959), Maldive Is (Alvarez and de Voogd, unpublished data) and Ashmore Reef, Australia. Remarks. The material examined here agrees in all its characteristics with Dragmacidon durissimum, an Indian Ocean species never previously recorded in Australia. Only one specimen from WA was found among the collections examined in this revision. More isolated populations might be present along the WA coast, given that the species is widely distributed throughout the Indian Ocean. The species was included originally in Thrinacophora due the presence of trichodragmata, later transferred to Sigmaxinella by Dendy (1922), and subsequently to Axinella by Burton (1959) without suficient justiication. Hallman (1917) erected Dragmacidon for D. agariciformis (Dendy, 1905), D. durissimum, D. clathriformis (Lendenfeld, 1888) and D. incrustans (Whitelegge, 1897). All these species are very similar in habitat, spicule composition and all include trichodragmata. They closely resemble to the West African species D. lunaecharta (Ridley and Dendy, 1886). Also similar are the Western Atlantic species D. reticulatum (Ridley and Dendy, 1886) and D. australe (see above), but they lack trichodragmata. Remarks on Dragmacidon. An additional species of Dragmacidon (described as Pseudaxinella sp. in Alvarez et al. 2000: 196) remains undescribed as no additional material has yet been found to fully characterise the species. The existing material is a fragment of a thin asymmetric lamella, found detached from original substrate that does not agree with the characteristically thickly encrusting shape of Dragmacidon species. A C B Fig. 12. Phakellia tropicalis sp nov.: A, light microphotograph of skeleton in cross section; B, schematic drawing of skeleton; C, diagram of spicules. Scale bars: A, 500 µm; B, 100 µm. Genus Phakellia Bowerbank, 1862 Gender feminine. Type species, by original designation, Spongia ventilabrum Linnaeus, 1767. Recent, Lervig, Norway, North Sea. Phakellia tropicalis sp. nov (Figs 7 E–F, 12 A–B) Phakellia sp. Alvarez et al., 2000: 195; Holmes and Blanch 2007: 761; Alvarez et al. 2007: 1600. Material examined. H olotype – NTM. Z.5847 (Fig. 7e), Stevens Rock, West Arm, Darwin Harbour, 12°29.1667´S, 130°47.19´E, NT, 9 m depth, 8 May 2006, coll. Alvarez, B. pArAtypes – NTM Z.5845 (Fig. 7F), Stevens Rock, West Arm, Darwin Harbour, 12°29.1667´S, 130°47.19´E, NT, 9 m depth, 8 May 2006, coll. B. Alvarez. AdditionAl specimens – Bynoe Harbour, NT:, NTM Z.4198, Z.4486, Z.4488. Darwin Harbour, NT: NTM Z.866, Z.877, Z.1948, Z.4197, Z.4428, Z.5665, Z.5839–Z.5842, Z.5844, Z.5848, QM G303365, G303383. Wessel Is: NTM Z.4463. 29 Papua New Guinea: QM G312926, G312937. Malaysia: NTM Z.5843. Description. Shape (Fig. 7E–F). Small convoluted thin fans, up to 250 mm high and 300 mm wide, on short and thin peduncle, lexible, less than 5 mm thick, arranged in multiple planes, with luted or planar langes, ragged or crenulated margins. Colour. Bright orange, beige-orange, or yellow-brown (Darwin and Bynoe Harbour specimens) alive. Oscula. Star-shape oscula, minute. Surface. Velvety, felty, with choanosomal spicules projecting shortly; marked with ine network of excurrent channels ending in oscula and reinforced close to the peduncle by thick choanosomal axes or ‘veins’. Skeleton (Fig. 12A, B). A core of interwoven spicules, occupying most of specimen’s thickness, laterally compressed, echinated by a dense palisade of single spicules, sometimes aggregated in loose brushes, protruding through surface. Spicules (Fig. 12C). Strongyles wavy, 222–800 in length by 3–8 µm thick and styles 231–703 in length by 6–16 µm thick (Table 5). Remarks. (see Carvalho et al. 2007 for extensive review of Phakellia species.) B. Alvarez and J. N. A. Hooper The species is atypical of Phakellia, although it agrees with the diagnosis given by Alvarez and Hooper (2002) in most aspects. It is fan-shaped, with styles projecting through the ectosome, and includes the typical spicule composition of the genus. However the reticulation of thick ‘veins’ or thick axes observed in most species of Phakellia, seems to be either incomplete or obscured by the habit of the species here described. The skeleton of this species (especially when seen in cross section, as in Fig. 12A) resembles some species of Acanthella (Dictyonellidae), a genus often confused with Phakellia. Phakellia tropicalis however, lacks two of the main diagnostic features of Acanthella – the cartilaginous to membranous surface and the cavernous structure of the choanosomal skeleton, with sheets of aspiculous collagen joining the primary axes in the skeleton. Instead, the surface of P. tropicalis is velvety, felty, with choanosomal spicules projecting shortly through the surface, a character shared with other axinellid genera such as Axinella and Cymbastela, and the skeleton is formed by a core of interwoven strongyles with a regular and dense palisade of erect styles that occupies most of the thickness of the sponge. A phylogenetic analyses based on morphological characters by Alvarez et al. (2000) showed this species to be closely related to other species of Acanthella (i.e. A acuta, A. cavernosa and A. pulcherrima). In the same study, however, an analysis based on 28S rDNA sequences with the same set of species, indicated that P. tropicalis was closely related to typical axinellid genera such as Axinella and Dragmacidon supporting its allocation to Axinellidae instead of Dictyonellidae. Furthermore, the skeletal architecture of P. tropicalis corresponds with the basic plan observed in species of Bubaris Gray, 1867, a genus of the family Bubaridae and currently used in the strict sense to include encrusting forms (Alvarez and Van Soest 2002). Thus, one could interpret the new species as an ‘erect Bubaris’ and be tempted to allocate it to that genus. It is possible that species with erect forms originally described under Bubaris, but transferred to Acanthella and Phakellia in order to preserve the revised concept of Bubaridae by Alvarez and Van Soest (2002), might be related to the new species. If that is the case, those species could be grouped under a new genus following a revised version of Hentschel’s (1923) concept of Bubaridae, which accepted sponges of erect forms, where the core of interwoven strongyles echinated by styles is placed in the centre (or in the axis) instead of at the base, as in the encrusting forms. However, it would be premature to erect a new genus here based on the characteristic of one single species and without re-examining species currently hidden under Acanthella or Phakellia which might also share such characteristics and could justify the creation of the new genus. In the absence of such evidence, we assign the new species provisionally to Phakellia and propose to expand the deinition of Alvarez and Hooper (2002) (and modiied by Carvalho et al. 2007) as: Axinellidae of planar habit, with skeleton formed by multiple axes or a single core of sinuous megascleres (frequently strongyles), echinated either by single spicules or by secondary tracts of a second class of megascleres (frequently styles). Phakellia tropicalis is the only species of Phakellia recorded in this study and as far as we know the irst one recorded from warm waters and shallow depths. Note that some species currently accepted under Phakellia from similar habitats are likely to be misidentiications (see Carvalho et al. 2007 for an extensive review of Phakellia species). Other species of Phakellia recorded from northern Australia and adjacent areas are currently accepted under Acanthella (Van Soest et al. 2008) or Axinella (i.e. Phakellia aruensis Hentschel, 1912, see above). Yet other species described under Phakellia in the Indian Ocean are not typical of the genus: P. ceylonensis Dendy, 1905: 192 is an Axinella (see above); P. crassistylifera Dendy, 1905: 192 is likely to belong in Stylissa and Phakellia ridleyi Dendy, 1887: 159 is currently accepted as a species of Phakettia. Distribution. This species is found along the NT coast and is very common in both Darwin and Bynoe harbours. It is also recorded for Papua New Guinea and Malaysia. It is found between 5–20 m depth Etymology. Referring to its tropical habitat. It is intended as a noun in apposition. Genus Reniochalina Lendenfeld, 1888 Gender feminine. Type species, by subsequent designation of Hallmann (1914), Reniochalina stalagmitis Lendenfeld, 1888. Recent, Western Australia. 30 Reniochalina stalagmitis Lendenfeld, 1888 (Figs 7G–H, 13A–F, 14 A–E, 15) Reniochalina stalagmitis Lendenfeld, 1888: 82; Whitelegge 1902: 283; Hallmann 1914: 346; Hooper and Wiedenmayer 1994: 81; Hooper and Lévi 1993: 1404; Alvarez, et al. 2000: 197; Alvarez and Hooper 2002: 746; Holmes and Blanch 2007. Axinella echidnaea. – Ridley 1884: 462; Kieschnick 1896: 533; Hentschel 1912: 419 [misidentiication; not Ridley and Dendy 1887: 183] Reniochalina lamella Lendenfeld, 1888: 83; Whitelegge 1902: 283; Hallmann 1914: 346. Axiamon folium Hallmann, 1914: 441 [objective synonym, see Wiedenmayer (1989: 49) and Hooper and Lévi (1993: 1403)] Material examined. type mAteriAl – Reniochalina stalagmitis: Lectotype, BMNH 1887.4.27.122, Western Australia, Fig. 13A; paralectotype, AM G9004, wet, West Australia, Fig. 13B [also holotype of Axiamon folium]. Reniochalina lamella: AM B5478, syntype, wet, no locality data [also paratype of A. folium]. AdditionAl specimens – Ridley’s (1884) material: BMNH 1882.2.23.261, Prince of Wales Channel, Torres strait, North Queensland, coll. HMS Alert; BMNH 1881.10.21.259, Thursday I., Torres strait, North Queensland, coll. HMS Alert. Hentschel’s Axinellidae from northern Australia Table 5. Comparison of spicule dimensions among specimens of Phakellia tropicalis. Measurements in micrometres. Specimen Locality Z.5847 Holotype, Stevens Rock Z.4488 Bynoe Harbour Z.4463 Wessel Is G312926 Papua New Guinea Strongyles Styles 284.1–651.2 (480.4±105.3) x 3.8–8.4 (5.7±1.2) 222.8–670.7 (435.5±135) x 3.7–9.5 (7±1.6) 293.1–800 (553.4±134.6) x 4.4–8.4 (6.6±1.1) 277.8–696.3 (476.4±117.6) [24] x 4.2–8.4 (6.3±1.1) 231.9–549.2 (385.7±82.5) x 7.3–430.9 (29.8±83.6) 353.9–703.6 (484.6±102.2) x 9–16.4 (13.8±2) 273.6–658.2 (439.6±111.1) x 8.3–16 (11.2±2.2) 239.6–490.6 (343.5±69.5) x 5.9–11.5 (8.7±1.8) (1912) specimen, SMF 1687, Aru-Inseln, bei Pulu Bambu, Indonesia, 10 m depth, 3 April 1908, coll. H. Merton, dredge. NW Shelf, WA: NTM Z.2358, Z.2361, Z.2273, Z.724, Z.738. Ashmore reef, WA: QM G301093, G301112, G301139. NE Joseph Bonaparte Gulf, QM G301202. Fog Bay, NT, QM G303548. Bynoe Harbour, NT: NTM, Z.4462 (0M9H2388-N), Z.5074 (0M9H2451-H), Z.5853. Darwin Harbour, NT: QM G303329, G303362, G303374, G303579, NTM Z.227, Z.285, Z.474, Z.483, Z.815, Z.1107, Z.1989, Z.2686, Z.4448 (0M9H2005-V), Z.5854, Z.5855, Gunn Point, NT, QM G303535. Parry Shoals, NT, Z.525. Melville I., NT, NTM Z.608. Cobourg Peninsula, NT: NTM Z.67, Z.135, Z.537, Z.565, Z.1335, Z.2527. Groote Eylandt, NT, G313555. Wessel Is, Z.5075 (0M9H2647-T). Gulf of Carpenteria, NT, QM G300817. Torres Strait, QM G316882. Description. Shape (Figs 7G–H, 13 A–B). Arborescent, branching or fan-shaped, generally stalked, and erect with specimens up to 60 cm high. Branches, lat to cylindrical, dichotomous or fused, 1–2 cm diameter, generally with pointed tips, dividing and anastomosing irregularly in different planes. Colour. Two colour forms, orange-red and beige-yellow. Always brown in alcohol. Oscula. Small, less than 5 mm diameter, with elevated rims thin, membranous and transparent, irregularly distributed through the branches. Surface. Long surface processes or conules with projecting spicules, up to 5 mm long, evenly distributed and separated by more-or less longitudinal and parallel channels, 1 mm apart, 1 mm deep. Skeleton (Fig. 13C–F). Specialised ectosomal skeleton absent; choanosomal skeleton differentiated into axial and extra-axial regions. Extra-axial skeleton reticulated with ascending spongin ibres, 50–100 µm interconnected at all angles by single spicules or short ibres, or anastomosing and forming oval to round meshes up to 200 µm in diameter; spongin fibres slightly developed and cored with paucispicular tracts of spicules, sometimes plumose; projecting into surface processes and becoming dense and disorganised at tips. Axial skeleton slightly condensed and reticulated as in extra-axial region. Spicules (Fig. 14A–E; Table 6). Oxeas or anisoxeas, occasionally modiied to styles, straight, bent or slightly sinuous; with tips surmounted by microspines, which might be rudimentary or absent; microspines at one end might be half the compared size to those at other end (175–450 x 6–20 µm). Thin, sinuous styles or oxeas with smooth or slightly spined ends, 156–288 x 3–6 µm (Fig. 14A–C), scattered through the choanosomal skeleton, rare in most examined specimens. Few long styles, projecting through ectosome, present in some specimens but extremely rare. Remarks. The thin and sinuous styles (Fig. 14) were irst mentioned by Hallmann (1914) in his description of Axiamon folium. The presence of these spicules was conirmed in all the specimens examined here and point out relationships of Reniochalina with other members of the family Raspailiidae (order Poecilosclerida). Long and slightly sinuous styles, most of which were broken, were also observed in the type material of Axiamon folium (AM G9004) and in some of the specimens examined. These were extremely rare and scattered throughout the extraaxial skeleton and projecting through the surface, indicating further afinities with raspailid taxa and challenging the position of the genus within Axinellidae (see below). Distribution. Reniochalina stalagmitis is one of the most abundant sponge species of northern Australia. It is found from the intertidal to depths of 60 m. Its distribution extends well beyond the boundaries of the studied region (Fig. 15) with validated records from the northern region of WA (down to W Buccaneer Archipelago) and QLD (down to the Howick Is region). It also occurs in Indonesia. Notes on Reniochalina. Reniochalina was deined by Alvarez and Hooper (2002) as ‘Axinellidae with extra-axial spongin ibres projecting into surface processes and cored with paucispicular tracts of oxeas, anisoxeas and styles. Oxeas with tips surmounted by micro-spines’. The genus was considered to be closely related to other axinellid genera (i.e. Ptilocaulis and Phycopsis) by Alvarez and Hooper (2002), based on the shared presence of conspicuous, long, ilamentous surface processes and their skeletal features. The close afinities with Ptilocaulis were further conirmed by molecular phylogenetic analyses (Alvarez et al. 2000). Recent molecular studies (Erpenbeck et al. 2007; Holmes and Blanch 2007) also showed strong afinities of Reniochalina stalagmitis with the raspaillid species Axechina raspailoides. As previously indicated (Hooper 1991; Hooper 2002b), these species have similar growth forms and choanosomal skeletons, and it is now confirmed that they also share the presence of styles 31 B. Alvarez and J. N. A. Hooper Fig. 13. Reniochalina stalagmitis: A, lectotype, BMNH 1887.4.27.122; B, paralectotype, AM G9004; C, lectotype, BMNH1887.4.27.122, light microphotograph of skeleton; and D, SEM of skeleton; E, paralectotype, AM G9004, light microphotograph of skeleton and F, SEM of skeleton. Scale bars: A, B, 2 cm; C–F, 500 µm. 32 Axinellidae from northern Australia with spined tips and identical shape (Fig. 14B, C), a fact overlooked by previous authors with the exception of Hallmann (1914). Additional molecular analysis based on the CO1 fragment (Erpenbeck 2007) also indicated afinities of the Caribbean species Ptilocaulis marquezi (Duchassaing and Michelotti, 1864) with other raspailiid species (i.e. Pandaros acanthifolium (Duchassaing and Michelotti, 1864) and Ecyoplasia ferox (Duchassaing and Michelotti, 1864) suggesting strongly that both Ptilocaulis and Reniochalina are closely related to the Raspailiidae. These relationships however, remain unresolved as neither Reniochalina nor Ptilocaulis have the typical raspailiid ectosomal skeleton which is clearly present in Axechina and other raspailiid taxa. Unfortunately, the range of taxa sampled in the molecular analyses mentioned above was inadequate to conclude further on the afinities of the axinellid genera Reniochalina and Ptilocaulis (and likely Phycopsis too) with the family Raspailiidae. Therefore, if more evidence from molecular analyses becomes available to support the current results, these genera might be relocated to the Raspailiidae. Other species of Reniochalina reported in the literature do not agree with the current deinition of the genus and are referred here to more appropriate genera: Reniochalina condylia Hooper and Lévi, 1993, to Dragmacidon; R. plumosa Lévi and Lévi, 1983 to Axinella and, R. sectilis Wiedenmayer, 1989 to Rhaphoxya in Dictyonellidae. Additional species of Reniochalina including Reniochalina sp., reported in Alvarez et al. (2000), remain to be described from other areas of Australia (Hooper, pers. obs.). It is likely also that some species of Reniochalina are misidentiied as species of Ptilocaulis (e.g. P. rigidus Carter, 1883:322). initial stages of this revision indicated that the Axinellidae was represented in the area by a larger number of species. This observation turned out to be contrary to what was found after a thorough examination of recorded material. Many of the species thought to belong to the Axinellidae were in fact found to be members of other families and orders, particularly Raspailiidae (Poecilosclerida).This relects that separation of species within this group is extremely subjective and could be erroneous if is taken in an isolated context. Examination of the taxonomic characters across a large number of specimens is critical to detect the variability and plasticity of morphological characters present in this group and to avoid incorrect splitting of taxa. It is possible that cryptic species or hybrid forms are hidden within the continuum of variability commonly observed in species of Axinellidae; but only results of population genetic studies can reveal such cryptic species, and indeed verify the occurrence of hybridisation among the Porifera, as has been demonstrated for the Cnidaria (e.g. Veron 1995, Van Oppen 2000 and references within). The ive remaining genera of the Axinellidae (i.e. Auletta, Dragmaxia, Pipestela, Ptilocaulis and Phycopsis) are curiously not represented in the studied material despite the fact that they do include tropical species. Axinella loribellae, Cymbastela stipitata and C. vespertina are the only species of axinellids reported here with distributions restricted to northern Australia, reflecting some degree of endemism in the area. The latter sibling species pair represent western components of east-west coast species pairs of the genus Cymbastela, with C. coralliophila and C. concentrica of the east coast, presumed remnants of Pleistocene separation of northern Australian faunas during low strand sea levels (e.g. Hooper and Ekins 2004). Axinella sinoxea displays a similar distribution but can be found in deep waters of WA. The remaining species seem to have a more widespread distribution throughout Indonesia, Papua New Guinea and Malaysia. Axinella aruensis and Reniochalina stalagmitis are widely distributed in the northern region of Australia and Indonesia. Dragmacidon australe has a disjunct distribution with isolated records from New Zealand, GBR and northern DISCUSSION The results of this revision indicate that the Axinellidae is represented in the area covered by this study by only ive genera (i.e. Axinella, Cymbastela, Dragmacidon, Phakellia and Reniochalina) and nine species, three of which are new. Data gathered from NTM and QM collections during the Table 6. Comparison of spicule dimensions among specimens of Reniochalina stalagmitis. Measurements in micrometres. Specimen Locality Lectotype BMNH1887.4.27.122 WA AM G9004 WA AM B5478 WA G303362 Darwin Z.4462 Bynoe Harbour Wessel Is Z.5075 Oxeas Thin styles/oxeas 190–315 (248.6±27.9) x 8–15.2 (12.1±2.4) 175–450 (243.9±61.8) x 7.9–20 (10.7±3) 197–376.6 (248.3±49.5) x 10.1–19 (14.4±1.9) 209.1–356.6 (253.3±43.4) x 9.2–17 (13±2.2) 195.1–379.1 (293.5±39.1) x 7.1–18.2 (13.9±2.6) 199.6–380.2 (286.4±44.6) x 6.3–18 (12.4±2.8) 157.1–287.6 (240.2±35.3) [18] x 2.6–5.8 (4.5±0.9) [18] 211.7–268 (244±22.7) [9] x 2.6–5 (3.8±0.8) [9] 156.4–235 (203.7±25.3) [12] x 3.5–6.3 (4.9±0.8) [12] 206.5–232.6 (219.5±18.4) [2] x 3.9–4 (3.9±0) [2] 158.2–244.4 (213.2±40.2) [4] x 2.6–5.2 (4.2±1.1) [4] 205.8x3.7 [1] 33 B. Alvarez and J. N. A. Hooper A C B D E Fig. 14. Reniochalina stalagmitis: A, diagram of spicules; B, C, SEM of sinuous style with spined tip (AM G9004); D, E, tip of oxeas, (AM G9004). Scale bars: A, 50 µm; B, 5 µm; C, 20 µm; D, E, 2 µm. Australia. Dramacidon durissimum is a species common from Indian Ocean but its distribution is now extended to the Ashmore Reef, WA of northern Australia. The position of Reniochalina within the Axinellidae is now debatable. The presence of sinuous styles with spiny tips in R. stalagmitis and the recent evidence based on molecular data (see above) suggest strongly that the species is closely related to the raspailiid species Axechina raspailioides (Poecilosclerida) and challenges the position of Reniochalina stalagmitis within the family. The typical ectosomal skeleton present in raspailiid species and considered a synapomorphy for that family is absent in R. stalagmitis, thus no deinitive conclusions can be made at this stage. New studies as suggested above are needed 120˚ E 130˚ 140˚ 5˚ 15˚ S Fig. 15. Distribution of Reniochalina stalagmitis in northern Australia, based on conirmed records from QM and NTM. Species distribution extends along the Queensland coast (Hooper, unpublished data). 34 Axinellidae from northern Australia to decide whether Reniochalina should remain classiied under the Axinellidae. Phylogenetic relationships within the family Axinellidae and other related groups have been explored previously, using either morphological, molecular or chemical characters (Alvarez et al. 2000; Erpenbeck et al. 2002; Erpenbeck et al. 2005; Erpenbeck et al. 2006; Erpenbeck et al. 2007). The relationships shown in those studies remain inconclusive at this stage but suggest strongly that the Axinellidae is a polyphyletic taxon with uncertain afinities. The taxonomic revision of species of the Axinellidae and its sister groups is critical to support conclusions derived from those studies and is currently the focus of ongoing studies undertaken at regional basis (e.g. Indonesia, Eastern Australia, CentralWest Paciic, Southern Australia and New Zealand). 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Erpenbeck, D., Breeuwer, J., Parra-Velandia, F. and Van Soest, R.W.M. 2006. Speculation with spiculation? Three independent gene fragments and biochemical characters versus morphology in demosponge higher classiication. Molecular Phylogenetics and Evolution 38: 293–305. ACKNOWLEDGEMENTS This work was funded by an Australian Biological Research Studies (ABRS) research grant (Grant No 20510) and by the ‘Collection and Taxonomy of Shallow Water Marine Organisms’ program for the US National Cancer Institute (Contract N02-CM-27003) subcontracted to NTM through CRRF. 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Journal of the Marine Biological Association of the United Kingdom 85: 1367–1371. 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. and Robertson, J. 2007. Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas. Bioscience 57: 573–583. Veron, J.E.N. 1995. Corals in space and time. the biogeography and evolution of the Scleractinia. University of New South Wales Press: Australian Institute of Marine Science, Townsville. Whitelegge, T. 1897. The Sponges of Funafuti. Memoirs of the Australian Museum 3: 323–332, pl. 18. Whitelegge, T. 1902. Notes on Lendenfeld’s Types described in the Catalogue of Sponges in the Australian Museum. Records of the Australian Museum 4: 274–288. Wiedenmayer, F. 1989. Demospongiae (Porifera) from Northern Bass Strait, Southern Australia. Memoirs of the Museum of Victoria 50: 1–242. Ridley, S. O. and 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. Ridley, S.O. and Dendy, A. 1887. Report on the Monaxonida collected by H.M.S. “Challenger” during the Years 1873–76 Report on the Scientiic Results of the Voyage of H.M.S. ‘Challenger’ during the Years 1873–76. Pp. 1–275 pls 1–51. Her Majesty’s Stationery Ofice: Edinburgh, Dublin, London. Thomas, P. A. 1981. A second collection of marine Demospongiae from Mahe Island in the Seychelles Bank (Indian Ocean). Musée Royal de l’Afrique Centrale: Tervuren, Belgique. Soest, R.W.M. Van, Boury-Esnault, N., Hooper, J., Rützler, K., De Voogd, N.J., Alvarez, B., Hajdu, E., Pisera, A., Vacelet, J., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R. and Klautau, M., 2008. World Porifera database. Available online at http://www.marinespecies.org/porifera. Last consulted on 18 October 2009. Soest, R.W.M. Van, Erpenbeck, D. and Alvarez, B. 2002. Family Dictyonellidae. Pp. 773–786. In Hooper, J.N.A. and Soest, R.W.M. Van (eds) Systema Porifera. A guide to the supraspeciic classiication of the phylum Porifera. Plenum: New York. Soest, R.W.M. Van and Hooper, J.N.A. 2002. Order Halichondrida. Pp. 773–786. In: Hooper, J.N.A. and Soest, R.W.M. Van (eds) Systema Porifera. A guide to the supraspeciic classiication of the phylum Porifera. Plenum: New York. Accepted 21 October 2009 37 B. Alvarez and J. N. A. Hooper APPENDIX Collection and locality data of material examined in the collections of QM and NTM. QM material G300181 Passage West I., outer reef, Ashmore Reef, WA, 12°14´S, 122°56´E, 15.5 m, 27 Jul 1986, coll. Hooper, JNA G300295 Snake Reef, Howick Group, GBR, QLD, 14°27´S, 145°1´E, 12.5 m, 14 Dec 1990, coll. Hooper, JNA G300609 N side of Cumberland Strait, Wessel Is, Gove, NT, 11°28´S, 136°29´E, 13 m, 14 Nov 1990, coll. NCI, AIMS G300759 Marinbar I, SE Cape Wessel, Wessel Is, NT, 11°1.13´S, 136°46.04´E, 20 m, 17 Nov 1990, coll. NCI, AIMS G300768 Gugari Rip 100m NE, E Guluwuru I, Wessel Is, NT, 11°34´S, 136°22.12´E, 8 m, 13 Nov 1990, coll. NCI, AIMS G300817 Duyfken Point, W Gulf of Carpentaria, QLD, 12°34´S, 141°0´E, 58 m, 26 Nov 1991, coll. Cook, SD. on CSIRO RV Southern Surveyor G301089 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 14 m, 6 May 1992, coll. Hooper, JNA G301092 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 22 m, 7 May 1992, coll. Hooper, JNA G301093 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 22 m, 7 May 1992, coll. Hooper, JNA G301112 Cartier I, outer reef slope, S side of reef, WA, 12°32.15´S, 123°33.12´E, 23 m, 8 May 1992, coll. Hooper, JNA G301139 Hibernia Reef, entrance to lagoon, NE side reef, WA, 11°57.13´S, 123°22.06´E, 23 m, 10 May 1992, coll. Hooper, JNA G301197 Flattop Bank, NE Joseph Bonaparte Gulf, NT, 12°16´S, 129°15´E, 32 m, 17 May 1992, coll. Hooper, JNA G301202 Flattop Bank, NE Joseph Bonaparte Gulf, NT, 12°16´S, 129°15´E, 32 m, 17 May 1992, coll. Hooper, JNA G303262 South Shell I., reef N of boat ramp, East Arm, Darwin Harbour, NT, 12°29.1334´S, 130°53.09´E, 0 m, 19 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303322 East Point Bommies, Darwin Harbour, NT, 12°24.08´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303329 East Point Bommies, Darwin Harbour, NT, 12°24.08´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303332 East Point Bommies, Darwin Harbour, NT, 12°24.0834´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303362 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303365 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303374 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303383 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303388 Stevens Rock, West Arm, Darwin Harbour, 12°29.1667´S, 130°47.19´E, NT, 19 m depth, 24 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303444 Fish Reef, west side, Bynoe Harbour, NT, 12°26.01´S, 130°26.09´E, 11 m, 26 Sep 1993, coll. Hooper, JNA and Hobbs, LJ G303535 Shoal Bay, W Gunn Point, NT, 12°9.15´S, 130°56.02´E, 14 m, 11 Oct 1993 G303548 Fog Bay, 1 nml E Point Blaze, NT, 12°54.15´S, 130°7.16´E, 7 m, 4 Oct 1993 G303579 Lee Point near Anglers Reef, Darwin Harbour, NT, 12°18.13´S, 130°52.14´E, 10 m, 11 Oct 1993 G304182 Granite Bluff, Lizard I., S headland Mermaid Cove, QLD, 14°39´S, 145°27´E, 18 m, 4 Apr 1994, coll. Hooper, JNA and party G304246 Cobia Hole, Mrs Watson´s Bay, Lizard I., QLD, 14°39.03´S, 145°26.15´E, 18 m, 5 Apr 1994, coll. Hooper, JNA and party G304253 Palfrey I., W side, Lizard I., QLD, 14°42.03´S, 145°26.09´E, 16 m, 6 Apr 1994, coll. Hooper, JNA and party G310136 Parry Shoals 35nm W Bathurst I., NT, 11°7.03´S, 129°25.9´E, 16 m, 12 Aug 1987 G311873 100m NE Gugari Rip, East side Guluwuru IS, Wessel Is, NT, 11°20.4´S, 136°13.63´E, 8 m, 13 Nov 1990 G312926 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.05´S, 148°10.14´E, 20 m, 15 Dec 1996, coll. Hooper, JNA G312913 Coutance Islet, Kupiano, SE. Papuan Barrier Reef, Papua New Guinea, 10°14.0167´S, 148°6.14´E, 41 m, 14 Dec 1996, coll. Hooper, JNA G312935 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.0501´S, 148°10.14´E, 20 m, 15 Dec 1996, coll. Hooper, JNA G312937 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.05´S, 148°10.14´E, 20 m, 15 Dec 1996, coll. Hooper, JNA G313555 S Groote Eylandt, NT, 14°27.1801´S, 136°14.29´E, 22.5 m, 12 Oct 1997, coll. Cook, SD. on CSIRO RV Southern Surveyor G316882 Torres Strait, QLD, 10°46.8´S, 142°15´E, 16.4 m, 19 Jan 2004, coll. TSMap_GM_01_2004 Gwendoline May G320664 Munro Reef, Coral Sea, QLD, 14°18.15´S, 144°48.82´E, 23 m, 2 Jul 2003, coll. Hooper, JNA and party 38 Axinellidae from northern Australia APPENDIX (continued) Collection and locality data of material examined in the collections of QM and NTM. NTM material Z.67 Z.135 Z.227 Z.285 Z.474 Z.483 Z.525 Z.537 Z.565 Z.608 Z.615 Z.619 Z.630 Z.631 Z.632 Z.665 Z.724 Z.738 Z.815 Z.822 Z.866 Z.868 Z.877 Z.1107 Z.1335 Z.1363 Z.1388 Z.1948 Z.1961 Z.1989 Z.2156 Z.2246 Z.2249 Z.2273 Z.2284 Z.2304 Z.2310 Z.2322 Z.2331 Z.2345 Z.2358 Z.2361 Z.2402 Z.2511 Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.50´S, 132°3.01´E, 17 Oct 1981, coll. Hooper, JNA & Alderslade, PN Sandy I. No.2, Cobourg Peninsula, NT, 11°5.50´S, 132°17´E, 10 m, 21 Oct 1981, coll. Hooper, JNA & Alderslade, PN Lee Point, Darwin, NT, 12°19.0167´S, 130°53´E, 14 Nov 1981, coll. Hooper, JNA Dudley Point Reef, East Point, Darwin, NT, 12°25.00´S, 130°48.01´E, 1 m, 18 Sep 1981, coll. Hooper, JNA & Murray, P Fannie Bay, Darwin, NT, 12°25.00´S, 130°50´E, 9 Feb 1982, coll. Hooper, JNA Fannie Bay, Darwin, NT, 12°25.00´S, 130°50´E, 9 Feb 1982, coll. Hooper, JNA Parry Shoals 35nm W Bathurst I., NT, 11°7.03´S, 129°25.9´E, 1 m, 30 Apr 1982, coll. Hooper, JNA & Alderslade, PN Port Bremer, Cobourg Peninsula, NT, 11°8.5´S, 132°18.8´E, 1 May 1982, coll. Hooper, JNA & Alderslade, PN Sandy I. No.2, Cobourg Peninsula, NT, 11°5´S, 132°16.51´E, 14 m, 2 May 1982, coll. Hooper, JNA Cootamundra Shoals, North of Melville I., NT, 10°49.07´S, 129°12.09´E, 31 m, 6 May 1982, coll. Thom, B & Lockyer, R Cootamundra Shoals,North of Melville I., NT, 10°50.22´S, 129°13.17´E, 22 m, 10 May 1982, coll. Lockyer, R Unnamed shoal N Melville I, NT, 11°38.23´S, 129°51.00´E, 24 m, 17 May 1982, coll. Thom, B & Lockyer, R Unnamed shoal N Melville I, NT, 11°32.58´S, 130°02.50´E, 18 m, 25 May 1982, coll. Lockyer, R Unnamed shoal N Melville I, NT, 11°32.58´S, 130°02.50´E, 18 m, 25 May 1982, coll. Lockyer, R Unnamed shoal N Melville I, NT, 11°32.57´S, 130°2.51´E, 18 m, 25 May 1982, coll. Lockyer, R NW Yampi Sound, NW Shelf, WA, 15°27.0334´S, 121°49.01´E, 76 m, 29 Apr 1982, coll. CSIRO R.V. Sprightly N Adele I.,Collier Bay, NW Shelf, WA, 15°58.02´S, 122°39.07´E, 59 m, 21 Apr 1982, coll. CSIRO R.V. Sprightly N Adele I.,Collier Bay, NW Shelf, WA, 15°58.02´S, 122°39.07´E, 59 m, 21 Apr 1982, coll. CSIRO R.V. Sprightly Channel I., Middle Arm, Darwin, NT, 12°32.02´S, 130°51.02´E, 11 m, 16 Jul 1982, coll. Scott Chidgey (Caldwell Connell Ass Channel I., Middle Arm, Darwin, NT, 12°33.08´S, 130°51.04´E, 20 m, 18 Jul 1982, coll. Scott Chidgey (Caldwell Connell Ass) Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN. Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN. Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN. Dudley Point Reef, East Point, Darwin, NT, 12°25.00´S, 130°48.01´E, 22 Dec 1982, coll. Hooper, JNA Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5´S, 132°10.51´E, 11 May 1983, 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 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2´S, 130°47.1´E, 27 Apr 1984, coll. Hooper, JNA Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2´S, 130°47.1´E, 27 Apr 1984, coll. Hooper, JNA West side of Weed Reef, Darwin, NT, 12°29.2001´S, 130°47.1´E, m, 11 May 1984, coll. Hooper, JNA and party Northern tip of Weed Reef, outer reef slope, Darwin Harbour, NT, 12°29.2´S, 130°37.61´E, 5 Oct 1984, 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, 10 m, 12 Apr 1985, coll. Hood, C and party NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish project) NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC Dudley Point Reef, East Point, Darwin, NT, 12°24.5´S, 130°48.01´E, 8 m, 29 Jul 1985, coll. Hooper, JNA Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3´S, 132°3.71´E, 15 Sep 1985, coll. Hooper, JNA 39 B. Alvarez and J. N. A. Hooper APPENDIX (continued) Collection and locality data of material examined in the collections of QM and NTM. NTM material Z.2526 Z.2527 Z.2529 Z.2632 Z.2686 Z.2719 Z.2727 Z.3062 Z.3068 Z.3137 Z.3141 Z.3922 Z.3925 Z.3935 Z.3936 Z.3938 Z.3946 Z.3956 Z.4078 Z.4104 Z.4131 Z.4197 Z.4198 Z.4425 Z.4428 Z.4435 Z.4448 Z.4462 Z.4463 Z.4465 Z.4486 Z.4488 Z.4490 Z.4491 Z.5053 Z.5054 Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 18–20 m, 16 Sep 1985, coll. Hooper, JNA Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 18–20 m, 16 Sep 1985, coll. Hooper, JNA Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 17 Sep 1985, coll. Hooper, JNA 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 Dudley Point Reef, East Point, Darwin, NT, 12°24.5´S, 130°48.01´E, 3 Apr 1986, coll. Hooper, JNA and party Myrmidon Reef,GBR, QLD, 18°10.00´S, 147°23´E, 15 m, 1 Jan 1985, coll. Wilkinson,CR Parry Shoals, Arafura Sea, NT, 11°11.72´S, 129°43.26´E, 16 m, 12 Aug 1987, coll. Mussig, AM and NCI team Parry Shoals, Arafura Sea, NT, 11°11.72´S, 129°43.26´E, 16 m, 12 Aug 1987, coll. Mussig, AM and NCI team Parry Shoals, Arafura Sea, NT, 11°12.27´S, 129°42.71´E, 16 m, 14 Aug 1987, coll. Mussig, AM and NCI team Parry Shoals, Arafura Sea, NT, 11°12´S, 129°43.01´E, 16 m, 14 Aug 1987, coll. Mussig, A.M. and NCI (AIMS) Cumberland Strait, northern bay, Wessel Is, Gove Peninsula, NT, 11°27.5´S, 136°28.8´E, 20 m, 14 Nov 1990, coll. Hooper, JNA Cumberland Strait, northern bay, Wessel Is, Gove Peninsula, NT, 11°27.5´S, 136°28.8´E, 20 m, 14 Nov 1990, coll. Hooper, JNA N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper, JNA N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper, JNA N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper, JNA S W headland, Rimbija I., Cape Wessel, Wessel Is, Gove Peninsula, NT, 11°0.5´S, 136°43.79´E, 15 m, 16 Nov 1990, coll. Hooper, JNA N side Pugh Shoal, reef slope, NE of Truant I., English Company IS, Gove Peninsula, NT, 11°36.57´S, 136°53.39´E, 20 m, 18 Nov 1990, coll. Hooper, JNA Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand “Town Hall” hole, SW Channel I., Middle Arm of Darwin Harbour, NT, 12°33.74´S, 130°51.67´E, 19.5 m, 9 Sep 2004, coll. Alvarez, B Sand Island, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°35.291´S, 130°52.264´E, 7 m, 9 Sep 2004, coll. Alvarez, B Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.09´S, 130°47.1´E, 5–19 m, 8 May 2002, coll. Alvarez, B and party Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.09´S, 130°47.1´E, 5–19 m, 8 May 2002, coll. Alvarez, B and party Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°33.09´S, 130°52.43´E, 4 –8 m, 6 May 2002, coll. Alvarez, B and party Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°33.09´S, 130°52.43´E, 4 –8 m, 6 May 2002, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 5–10 m, 26 May 2003, coll. Alvarez, B and party Rimbija I., 2.8 km W of Cape Wessel, Wessel Is, eastern Arnhem Land, NT, 11°00.21´S, 136°43.84´E, 17–20 m, 1 Apr 2004, coll. Colin, P Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.57´S, 136°17.86´E, 11–20 m, 5 Apr 2004, coll. Alvarez B and party Raft Point, Bynoe Harbour, NT, 12°37.69´S, 130°32.16´E, 5–8 m, 26 Jun 2003, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 5 m, 1 Jun 2005, coll. Alvarez, B Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 5 m, 8 May 2006, coll. Alvarez, B Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B South Shell I., East Arm, Darwin Harbour, NT, 12°29.87´S, 130°53.12´E, 4–11 m, 18 Aug 2002, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party 40 Axinellidae from northern Australia APPENDIX (continued) Collection and locality data of material examined in the collections of QM and NTM. NTM material Z.5055 Z.5057 Z.5058 Z.5059 Z.5064 Z.5065 Z.5066 Z.5067 Z.5068 Z.5071 Z.5072 Z.5074 Z.5075 Z.5662 Z.5665 Z.5816 Z.5817 Z.5818 Z.5819 Z.5820 Z.5821 Z.5822 Z.5823 Z.5824 Z.5825 Z.5826 Z.5827 Z.5828 Z.5829 Z.5830 Z.5831 Z.5832 Z.5835 Z.5836 Z.5839 Z.5840 Z.5841 Z.5842 Z.5843 Z.5844 Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Raragala I., 700 m off NE tip Wessel Is, eastern Arnhem Land, NT, 11°32.85´S, 136°21.28´E, 13–16 m, 31 Mar 2004, coll. Alvarez, B Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.55´S, 136°17.96´E, 25–30 m, 5 Apr 2004, coll. Alvarez, B and party Weed Reef, entrance to West Arm, Darwin Harbour, NT, 12°29.25´S, 130°47.54´E, 9–15 m, 3 Aug 2002, coll. Alvarez, B and party Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.24´S, 130°35.56´E, 5–10 m, 23 May 2003, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.28´S, 136°17.52´E, 13–14 m, 5 Apr 2004, coll. Alvarez B. and party “Town Hall” hole, SW Channel I., Middle Arm of Darwin Harbour, NT, 12°33.74´S, 130°51.67´E, 10–18 m, 17 Sep 2002, coll. Alvarez, B and party Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.35´S, 130°31.45´E, 6–8 m, 11 Jun 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. Alvarez, B Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.49´S, 130°31.29´E, 9–10 m, 11 Jun 2003, coll. Alvarez, B and party Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll. Alvarez, B and party East Point, Darwin Harbour, NT, 12º24.16´S, 130º47.66´E, 11 m depth, 8 November 2008, coll. Ayling, A East Point, Darwin Harbour, NT, 12º24.16´S, 130º47.66´E, 11 m depth, 8 November 2008, coll. Ayling, A Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 3–12 m, 1 Jun 2005, coll. Alvarez, B Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.459´E, 6 m, 27 Apr 2007, coll. Alvarez, B Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.459´E, 6 m, 27 Apr 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43´E, 14 m, 25 May 2007, coll. Alvarez, B South Shell I., East Arm, Darwin Harbour, NT, 12°29.87´S, 130°53.14´E, 7–11. m, 19 Aug 2002, coll. Alvarez, B and party Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 17 m, 8 May 2006, coll. Alvarez, B Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B 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 Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.071´S, 130°47.103´E, 10–15 m, 9 May 2002, coll. Alvarez, B and party East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B Mengalum I., off Kota Kinabalu, Malaysia, 6 10.87´N, 115 35.97´E, 10–13 m, 24 Oct 2005, coll. Alvarez, B 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 41 B. Alvarez and J. N. A. Hooper APPENDIX (continued) Collection and locality data of material examined in the collections of QM and NTM. NTM material Z.5848 Z.5853 Z.5854 Z.5855 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12º42.2´S, 130º35.459´E, 3–12 m depth, 1 June 2005, coll. Alvarez, B Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12º42.2´S, 130º35.459´E, 3–12 m depth, 1 June 2005, coll. Alvarez, B 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 42

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