Axinellida (Porifera :Demospongiae) from the New Caledonia Lagoon John N. A. ~ o o p e r *and Claude ~ P v i ~ Queensland Museum, P.O. Box 3300, South Brisbane, Qld 4101, Australia. Laboratoire de Biologie des InvertCbrCs marins et Malacologie, Museum national d'Histoire naturelle, 57, rue Cuvier, 75005 Paris, France. A Abstract Sixteen species of axinellid demosponges, including seven new species and seven new locality records, are described from the shallow-water New Caledonian lagoon and reefs [families Axinellidae (12 species, Cyrnbastela, Reniochalina, Axinella, Phakellia, Stylissa, Ptilocaulis, Pseudaxinella, Rhaphoxya) and Desmoxyidae (4 species, Myrrnekioderrna, Higginsia)],bringing the total number of described axinellid species in this region to 25. Brief revisions are provided for several of these genera, based primarily on the Indo-west Pacific fauna, in order to place these New Caledonian species. Non-endemic New Caledonian axinellids belong predominantly to the north-eastern Australian (Solanderian province) and Indo-Malay fauna, usually representing the easternmost extent of these species' distributions in the Indo-west Pacific. Two species, Axinella carteri (Dendy) and Astrosclera willeyana Lister, were found to be truly widely distributed throughout the Indo-Pacific, typically associated with coral reefs; other previously suspected widely distributed species were generally found to be allopatric, cryptic sibling species related to the tropical Australasian fauna. Introduction Prior to the present work, only 11 species belonging to the (polyphyletic) order Axinellida had been described from the New Caledonian region: five in the family Axinellidae (four of which are apparently endemic and one also known from the Indo-Malay archipelago), one in Desmoxyidae (also known in northern New Zealand), one in Trachycladidae (also known from New Zealand and south-eastern and south-western Australia), two in Agelasidae [now considered t o be a separate order: Hartman 1982 (one of which is apparently endemic)], and two hypercalcified 'sclerosponges' described by Vacelet (1981) which have also been suggested as having possible affinities with the Agelasida (e.g. Vacelet 1981; van Soest 1984). However, these species represent only a small proportion of the axinellid fauna known t o live in this region; furthermore, 6 of these 11 described species are strictly deeper-water taxa, collected from the shelf and slope to the south of New Caledonia. Thus, the shallowwater axinellid sponge fauna is virtually undescribed, although many species have already been collected by ORSTOM. The present paper describes 16 species of shallow-water sponges in the families Axinellidae and Desmoxyidae living in the lagoon and coral reef habitats surrounding the island. These species represent some of the more visible axinellids in New Caledonian shallow-water habitats, although there are still many more taxa yet to be described from this region (LCvi, unpublished data). J. N. A. Hooper and C. LCvi 1396 This is the second of a series of papers on the taxonomy of demosponges in the lagoon and outer reefs of New Caledonia (Hooper and LCvi 1993); a third contribution will examine the biogeographical relationships of the New Caledonian Microcionidae, Raspailiidae and Axinellidae (Hooper and LCvi, unpublished data). These papers represent the results of a collaborative investigation amongst several sponge biologists (C. Battershill, P. Bergquist, J. Fromont, J. Hooper, M. Kelly-Borges, C. LCvi, J. Vacelet, C. Wilkinson), to document the major components of this sponge fauna, arising from a series of workshops jointly funded by ORSTOM and DITAC. The eventual aim of this investigation is to produce a comprehensive taxonomic inventory of the islands' shallow-water sponges, and an authoritative lay field guide to this fauna. Unlike the previous study on Poecilosclerida (Hooper and LCvi 1993) for which there were good contemporary revisions for most of the higher taxa, the axinellids have not been substantially revised, and hence some level of revision was necessary for many of the genera in order to place the New Caledonian species. These revisions, however, are certainly incomplete and relatively cursory due to the unavailability of much of the relevant type material required to undertake this considerable task. Methods Methods used to prepare and examine sponges for both light microscopy and scanning electron microscopy follow the procedures described in Hooper (1991). Measurements are based on examination of 25 random samples of each spicule category for each specimen, and are cited as range (and mean in parentheses). Abbreviations used in the text are as follows: AIMS, Australian Institute of Marine Science, Townsville; AM, Australian Museum, Sydney; BMNH, The Natural History Museum, London; CMMI, Canterbury Museum, Marine Invertebrate collection, New Zealand; CSIRO, Commonwealth Scientific and Industrial Scientific Organisation, Division of Fisheries, Hobart; DITAC, Commonwealth of Australia Department of Industry, Technology and Commerce, Canberra; ICZN, International Commission for Zoological Nomenclature (Anon. 1985); LFM, Merseyside County Museums, Liverpool (formerly Liverpool Free Museum); MMBS, Mukaishima Marine Biological Station, Faculty of Science, Hiroshima University, Onomichi; MNHN, MusCum National dYHistoireNaturelle, Paris; MHNG, MusCum d'Histoire Naturelle de Genkve, Geneva; NCI, U.S. National Cancer Institute shallow-water collection contract (Australian Institute of Marine Science, Townsville; primary collections in AIMS, NTM and QM, duplicates and fragments in USNM); NMNZ, National Museum of Natural History, Wellington; NMV, Museum of Victoria, Melbourne; N.S.W., New South Wales; NTM, Northern Territory Museum of Arts and Sciences, Darwin; ORSTOM, Institut Frangais de Recherche Scientifique pour le Developpement en Cooperation, Centre de Noumea; QFS, Queensland Fisheries Service; Qld, Queensland; QM, Queensland Museum, Brisbane; SM, Strasbourg Museum, France; SMF, NaturMuseum und Forschungsinstitut Senckenberg, Frankfurt; USNM, National Museum of Natural History, Smithsonian Institution, Washinton D.C.; ZMB, Zoologisches Museum fiir Naturkunde an der Humboldt-Universitat zu Berlin, Berlin; ZMC, Zoologisk Museum, Copenhagen. Systematics Order AXINELLIDA Carter Family AXINELLIDAE Ridley & Dendy Definition Axinellid sponges which generally lack microscleres, although several genera (Dragmaxia, Dragmacidon, Dragmatella, Dragmatyle, Rhaphoxya, Raspaigella and Tragosia) may have raphides, occurring singly or grouped into bundles (trichodragmata). Skeleton typically divided into distinct axial and extra-axial components; main skeletal tracts, composed of spongin fibres enclosing megascleres, usually condensed in axis; extra-axial region becoming plumose or plumoreticulate towards surface. However, many exceptions to this pattern shown in this family, and affinities of many species are still contested but poorly understood. Megascleres are monactinal, diactinal, or both, and sinuous spicules (usually strongyles or oxeas) reappear throughout many genera. Axinellidae frequently have branching growth form but funnel-shaped, flabellate, tubular, and massive forms also occur (modified from Wiedenmayer 1989). New Caledonian Axinellids 1397 Remarks This family contains a large and heterogeneous assemblage of taxa, with about 50 nominal genera included, and is in urgent need of an extensive, detailed revision; hence, a comprehensive definition of the family is provided here. The heterogeneity of the family has been indicated by several authors (e.g. Bergquist and Hartman 1969; van Soest et al. 1990), and Hooper and Bergquist (1992) questioned whether the family was in fact monophyletic. Hooper et al. (1992) suggested that the Axinellidae should probably be subdivided further, although they did not propose how this should be done. Many species show a number of similarities with members of the order Halichondrida (skeletal structure, spicule geometry), and indeed a recent revision of that order (van Soest et al. 1990) indicated that the Axinellidae should be placed there. However, this hypothesis is premature and uncorroborated by any other evidence, having no greater or lesser support than the existing arrangement. Thus, the higher systematic relationships of the family are still not clear and, for the time being, the family is retained in a polyphyletic order, Axinellida. Members of this large family occur from polar regions to the tropics, and from tidal habitats to at least 4400 m depth (Hartman 1982). Many species of Axinellidae are known to live in the New Caledonian region (LCvi, unpublished data), but so far only five of these have been published from this region. These are: Ptilocaulis fusiformis LCvi, 1967, from the Bay St Vincent (apparently endemic); Cymbastela cantharella (LCvi, 1983) from outer reef slopes of New Caledonia (endemic); Axinella lifouensis LCvi & LCvi, 1983, from deeperwaters (> 350 m) to the south of the island (endemic); Phakellia columnata (Burton, 1928) also from deeper-waters (>300 m) (also known from the Andaman Sea); and Reniochalina plumosa (LCvi & LCvi, 1983), a new combination here, from deeper-waters (>400 m) (endemic). Genus Cymbastela Hooper & Bergquist Pseudaxinyssa (in part).-Bergquist and Tizard, 1967: 190; Lkvi, 1983: 719. Cymbastela Hooper & Bergquist, 1992: 103 [type species Pseudaxinyssa stipitata Bergquist & Tizard, 1967: 189 (holotype AM Z3101)]. Diagnosis Typically stalked, cup-shaped, thinly lamellate growth form. Ectosome with or without specialised skeleton of smaller oxeas. Choanosomal skeleton with compressed reticulate axial region, in which major tracts run longitudinally through lamellae, and with gradually ascending, diverging, radial, plumose or plumo-reticulate extra-axial region, in which tracts become plumose and project through surface. Spongin fibres well developed, cored by oxeas, frequently with telescoped points. Predominantly autotrophic, most species containing symbiotic cyanobacteria. Oviparous (from Hooper and Bergquist 1992). Remarks This genus was recently erected for atypical species previously referrable to Pseudaxinyssa Burton, including thinly lamellate, frequently cup-shaped, predominately coral reef species. The genus is known only from Australasia, with seven species described including one endemic to New Caledonia, C. cantharella. A second species from this region is described below. Cymbastela cantharella (LCvi) (Figs 1-2) Pseudaxinyssa cantharella Levi, 1983: 719-22, fig. 1, pl. 1. Cymbastela cantharel1a.-Hooper and Bergquist, 1992: 119-20, figs 12-14, table 1 . Material Examined Holotype. MNHN DCL3141: outer reef, SW. coast New Caledonia, 22°20'S.,166013'E.,40 m depth, coll. ORSTOM. J. N. A. Hooper and C. LCvi 1398 New Caledonian material. ORSTOM R1261 (fragment QM G30001.5): stn 198, pinnacle S. of Canyon Central, Chenal des Cinq Milles, 22"30.4'S.,166"45.1'E., 25 m depth, 15.ii.1978, coll. G. Bargibant, SCUBA; ORSTOM R163: stn 116, SW. pointe, Baie de Prony, 22°21.8'S.,166049.9'E., 40 m depth, 30.vi.1976, coll. P. Laboute, SCUBA. Comparative material. Refer to Hooper and Bergquist (1992) for additional material from New Caledonia. Description Colour. Pale orange-brown alive (Munsell 2.5-7.5 YR 8/10), beige in ethanol. Shape. Short, erect cup-shaped or vasiform sponges, up to 150 mm high, 170 mm maximum diameter, with relatively thick lamellae, up to 6 mm diameter, usually with convoluted margins, occasionally with secondary cups or lamellae growing inside primary cup, often with buttresses and exterior secondary projections, and with a short, cylindrical, basal stalk, up to 40 rnm long, 17 mm diameter. Surface. Predominantly smooth, with distinct interior (inhalant, osculiferous) and exterior (smooth but uneven, buttressed) faces of lamellae. Oscules small, up to 2 mm diameter, about 2 mm apart, each surrounded by slightly raised membraneous lip. Texture firm, flexible, slightly compressible. Ectosome. Membraneous, with heavy collagen, through which choanosomal oxeas protrude, individually or in paucispicular plumose bundles, arising from ascending radial tracts in peripheral skeleton. Choanosome. Choanosomal skeleton plumo-reticulate, without axial compression or axial and extra-axial differentiation. Two components of choanosomal skeleton present: longitudinal (radial) spongin fibres run through lamellae, cored by multispicular tracts of oxeas, plumose near periphery; transverse uni- or paucispicular tracts of oxeas interconnecting radial fibres; overall skeleton appears nearly disorganised, almost halichondroid. Mesohyl with heavy collagen. Megascleres. Oxeas short, slender, slightly curved, symmetrical, occasionally asymmetrical, tapering, fusiform, usually with slightly telescoped points. Length 143-(207.1)245 pm, width 2.5-(7.6)-12 pm. Microscleres. Absent. Distribution Known only from the New Caledonia region. Remarks The description given above is mainly condensed from Hooper and Bergquist's (1992) more comprehensive redescription of the species, as the present study includes only one Fig. 1. Cymbastela cantharella (Lkvi): specimen ORSTOM R1261, structural oxeas. New Caledonian Axinellids 1399 previously unpublished specimen from the south-west New Caledonia lagoon. This species has also been illustrated in more detail in this earlier work, and present illustrations are provided for comparative purposes. Cymbastela cantharella differs from other tropical species of the genus by its atypical orange pigmentation in life (most other species are green and mauve), prominent surface sculpturing on the inner (oscular) face of lamellae, having a dense radial-plumose skeleton, with a secondary paucispicular secondary reticulate skeleton, together producing a nearly halichondroid appearance, a radial-plumose ectosomal skeleton, and specific dimensions of its oxeas (refer to Hooper and Bergquist 1992: table 1). The species is most similar to C. stipitata, both having thinly flabellate, irregular, buttressed surface processes, and similar geometry of oxea megascleres, but the two species differ markedly in most other features. Fig. 2. Cymbastela cantharefla (Lkvi): A, specimen ORSTOM R1261 in situ (photo C . Debitus); B, specimen ORSTOM unregistered (stn 163/1) in situ (photo P . Laboute); C, skeleton [specimen QM G300015 (ORSTOM R1261)l (scale= 500 pm); D, peripheral skeleton (scale=200 pm); E, SEM skeletal structure (specimen QM G300004) (scale = 500 pm). 1400 J. N. A. Hooper and C. LCvi Cymbastela concentrica (Lendenfeld) (Figs 3-5, Table 1) Antherochalina concentrica Lendenfeld, 1887: 788, pl. 22, fig. 42. Cymbastela concentrica. -Hooper and Bergquist, 1992: 114-19, figs 9-1 1, table 1. Material Examined Lectotype. known. AM 21993: Port Molle (now Airlee Beach), Qld, 20°13'S.,148049'E., no other details New Caledonian material. QM G301229 (ORSTOM R180): stn 109, Baie des Citrons, Noumea, 22°18~0'S.,1660251'E., 10 m depth, 31.v.1976, coll. P. Laboute, SCUBA; QM G301230, G301264, G301266 (ORSTOM R153): stn 180, S. of sand cay, Ilat l'Areignere, 22°20~0'S.,1660191'E., 12 m depth, 4.v.1977, coll. P. Laboute, SCUBA; ORSTOM R363: stn 150, S. of entrance, Baie St Vincent, 22"02.2'S.,165"59.5'E., 18 m depth, 9.ix. 1976, coll. P. Laboute, SCUBA; ORSTOM 'cfR363': stn 148, NE. point, Ilat Mboa, 22"08.3'S.,166"09.3'E., 13 m depth, 9.ix.1976, coll. P. Laboute, SCUBA; ORSTOM 'cfR363': stn 154, channel between Ilat Puen and RCcif, 21°59~0'S.,165"57l1E., 14 m depth, 15.ix.1976, coll. P. Laboute, SCUBA; QM G301272, G301268, (ORSTOM RHO), ORSTOM R181: stn 110, SE. Il6t RCdika, 22°31~1'S.,166036.0'E., 15-19 m depth, 3-8.vi.1976, coll. P. Laboute, SCUBA; QM G301330: Croisant-Larigriere, Ilat Maitre, off Noumea, 22"20.2'S.,166"225'E., 20 m depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA. Comparative material. QM G301233: Davies Reef, Great Barrier Reef, Qld, 18"501S.,147"3g1E., 22 m depth, coll. C. R. Wilkinson, AIMS, 24.iii.1982, SCUBA; QM G301234: same locality, 15 m depth, coll. S. Seddon, 13.ix.1991, SCUBA; AIMS R.442-PS (NTM 22729): Myrmidon Reef, Great Barrier Reef, Qld, 18°10'S.,147"23'E., 15 m depth, l.i.1985, coll. C. R. Wilkinson, AIMS, SCUBA; NTM 23170: Blue Lagoon, Lizard I., Great Barrier Reef, Qld, 14°40'S.,145028'E., 10-20 m depth, 1.i.1987, coll. A. W. D. Larkum, SCUBA. [Refer to Hooper and Bergquist (1992) for additional material from eastern Australia.] Description Colour. Pale beige, olive-brown or reddish brown alive (Munsell 2.5Y 8/6-2.5YR 4/4); chlorophyll pigments present. Shape. Growth form predominantly vasiform, but varying from more-or-less symmetrical cup-shaped with small basal stalk, to vasiform with symmetrical or asymmetrical lamellae, Table 1. Comparison in spicule dimensions between known specimens of Cymbastela concentrica All measurements given in micrometres, and expressed as minimum-(mean)maximum range of measurement (comparative data from Hooper and Bergquist 1992). N = 2 5 for each specimen Material (Locality) AM 21993 (holotype) (Airlee Beach, 20"s.) NTM 23169 (Lizard I., 14"s.) QM G301233 (Davies Reef, 18"s.) QM G301234 (Davies Reef, 18"s.) ORSTOM R180 (Noumea lagoon, 22's.) ORSTOM R153 (Noumea lagoon, 22's.) QM G300003 (Moreton I., 26"s.) Oxeas Length Width 172-(239.6)-305 2.5-(9.5)-16 103-(145.6)-172 2.5-(3.3-6 138-(165.0)-184 4.0-(5.2)-6.0 92-(118.6)-146 3.5-(5.2)-8.0 107-(126.4)-142 3.0-(4.1)-5.0 96-(115.2)-132 3.0-(4.9)-6.5 67-(86.8)-104 2.5-(3.4)-4.5 New Caledonian Axinellids LATITUDE ( O S ) Fig. 4. Cymbastela concentrica (Lendenfeld), comparison of spicule dimensions with latitudinal distribution of samples: A, mean length; B, mean width. to thickly encrusting plate-like, attached directly to substrate. Size up to 150 mm high, 140 mm maximum width. Lamella thickness variable, ranging from card thin to thick and rubbery, 1.0-3.5 mm thick. Surface. Typically with convoluted, multiple lamellae inside cups or with digitate projections on exterior surface, but some specimens lack any surface ornamentation. Lamellae smooth, even or irregular. Texture flexible, compressible, velvet-like. 1402 J. N. A. Hooper and C. LCvi Ectosome. Membraneous without specialised skeleton, but microscopically villose from protruding spicules from peripheral skeleton, usually forming multispicular plumose brushes. Choanosome. Reticulate to plumo-reticulate mineral and fibre skeleton, with poorly differentiated axial and extra-axial regions. Reticulate skeleton predominant over plumose portion. Fibres in axial region only slightly condensed, forming an open reticulation, cored by uni- or paucispicular tracts of spicules. Extra-axial fibres reticulate, slightly plumose, paucispicular, whereas peripheral skeleton clearly diverges into plumose multispicular spicule bundles. Spongin fibres well developed; collagen abundant in mesohyl. Fig. 5 . Cymbastela concentrica (Lendenfeld): A , peripheral skeleton [specimen QM G301229 (ORSTOM R180)] (scale=200 pm); B, specimen ORSTOM R180 in situ (photo P . Laboute); C , specimen QM G301230 (ORSTOM R153) in situ (photo G. Bargibant); D, specimen ORSTOM 'cfR153' in situ (photo G. Bargibant); E, preserved specimen QM G301229; F, SEM skeletal structure (QM G301230) (scale=200 pm). New Caledonian Axinellids 1403 Megascleres (refer to Table 1 for dimensions). Oxeas vary considerably in size between specimens, usually slender, fusiform, straight or slightly curved, symmetrical, typically with very faintly telescoped points. Microscleres. Absent. Distribution Northern, central and southern Queensland, and south-west New Caledonia, found in the lagoon, inshore fringing reef or platform coral reef fauna, 10-22 m depth. Remarks This is a new locality record for this species in the New Caledonian region. Although C. concentrica has recently been comprehensively redescribed and illustrated (Hooper and Bergquist 1992), the description given above is based on a number of previously undescribed specimens from both the Great Barrier Reef and New Caledonia. The present records, together with material described by Hooper and Bergquist (1992), show that the species is widespread throughout all sections of the Great Barrier Reef, and it is possible that the species will eventually be found to inhabit many other islands and reefs throughout the Coral Sea. This species is remarkable for the heterogeneity in its spicule dimensions, unlike any of the other known species of the genus. Hooper and Bergquist (1992) suggested that this variability may correspond to latitudinal gradients in distribution of the species, although re-examination of spicule dimensions from all known material (Fig. 4) shows no such trend. It is possible that we have two or more sympatric sibling species presently included in C. concentrica but present morphometric characters cannot clearly distinguish these morphotypes. Genus Reniochalina Lendenfeld Reniochalina Lendenfeld, 1888: 82.-Hallmann, 1914b: 346; Wiedenmayer, 1989: 48 [type species Reniochalina stalagmitis Lendenfeld, 1888: 82 (lectotype BMNH 1887.4.27.122)l. Axiamon Hallmann, 1914~:440. -de Laubenfels, 1936: 130 [type species Axiamon folium Hallmann, 1914c: 440 ('syntypes' AM G9004, B5478)I. Diagnosis Arborescent, frondose, ramose or lobate growth forms; typically with tubercular, ridged or closely conulose surface; with plumo-reticulate skeletal structure, mainly in the form of ascending skeletal bundles, and without condensation of axial skeleton or differentiation between axial and extra-axial components; spiculation with interchangeable styles, oxeas and anisoxeas in approximately equal proportions and similar size, without differential distribution within the skeleton (modified from Wiedenmayer 1989). Remarks The above diagnosis is for the nominal genus Axiamon, since Lendenfeld's diagnosis of Reniochalina is grossly inadequate. Furthermore, this genus is best known under its junior synonym of Axiamon, and further comment is required on that synonymy. Hallmann (1914b: 346) noted that the primary description of ~endenfeld'sReniochalina stalagmitis was erroneous; consequently, Reniochalina was considered to be a genus dubium. Hallmann (1914~:440-1) subsequently described a specimen under the name of Axiamon folium (from Western Australia), which became the type species of the genus Axiamon, whilst at the same time he recognised that it was probably identical to Lendenfeld's R. stalagmitis (from the Illawarra region, N.S.W.). De Laubenfels (1936: 47) concurred with Hallmann. However, although Lendenfeld's (1888) definition of R. stalagmitis was not entirely accurate, differing significantly from the actual characteristics of the species (Whitelegge 1902: 283), it is still true that Lendenfeld's published name has priority since it is the earliest available for the species (Wiedenmayer 1989): hence, Axiamon is an objective synonym of Reniochalina. 1404 J . N. A. Hooper and C. Ltvi Prior to the present study, only four nominal species of the genus were known, but of these species only two are valid. Two other species are also referred to the genus in the present study. The genus now contains four valid species, all from the Indo-west Pacific region: R. condylia, sp. nov., from the shallow-water New Caledonian fauna (see below) (Figs 7-8); R. plumosa (LCvi & LCvi, 1983), comb. nov. (holotype MNHN DCL2972), from the deeper-water New Caledonian fauna (Fig. 6C-D); R. sectilis Wiedenmayer, 1989: 49, pl. 4, fig. 1, pl. 23, fig. 1 (holotype NMV F51962), from south-east Australia (not figured here); and the type species R. stalagmitis Lendenfeld, 1888, from north-west and north-east Australia (Fig. 6A-B) [with synonyms R. lamella Lendenfeld, 1888 (holotype AM B5478 from Western Australia), and Axinella echidnaea sensu Ridley (1884; specimen BMNH 1882.2.23.261 from north-west Australia) and Hentschel (1912; specimen SMF1687 from southern Indonesia) not Spongia echidnaea Lamarck, 1814 (Wiedenmayer 1989)]. Recent collections from tropical Australasia also discovered another six undescribed species referable to this genus, which will be described elsewhere in the future. Reniochalina is characterised by its plumo-reticulate skeletal architecture, and in having anisoxeas, sometimes with spinose extremities. The skeletal architecture of R. stalagmitis is certainly atypical of Axinellidae, and the existence of such structures makes the task of satisfactorily diagnosing the family a difficult one, but that feature is certainly not unique in the family (e.g. Phycopsis). Reniochalina should also be compared to Axinosia in skeletal construction, and to Ptilocaulis, from which it differs mainly in lacking axial skeletal condensation, or differentiation between axial and extra-axial skeletons, and also in lacking telescoped endings on spicules. Wiedenmayer (1989) suggested that the variability in spiculation known to occur in Reniochalina (e.g. Hallmann 1914c: 443), and the confused skeletal architecture in some species (e.g. Wiedenmayer 1989), raises doubts about which characters should be treated as most important in the systematics of axinellid genera (spicule geometry or skeletal structure) since these appear to occur in all combinations (cf. Axinosia and Pseudaxinella, for example). It is clear that an evaluation of those characters in the taxonomy of these genera is central to a revision of Axinellidae, but a satisfactory resolution of these systematics will probably not be possible without additional (non-skeletal) evidence. Fig. 6. Reniochalina spp.: A, Reniochalina stalagmitis Lendenfeld, peripheral skeleton (leitotype BMNH 1887.4.27.122) (scale= 500 pn);B, specimen NTM 21107 in situ (photo J. N. A. Hooper); C, Phakellia plumosa Ltvi and LCvi, peripheral skeleton (paratype MNHN DCL2974) (scale =200 pm); D, paratype. New Caledonian Axinellids 1405 If the evidence cited for Reniochalina were to be carried over to other genera, (viz. the unstable nature of axial and extra-axial differentiation, and the presence or absence of certain categories of megascleres), then many of the nominal axinellid genera would have to be merged. Reniochalina condylia, sp. nov. (Figs 7-8) Material Examined Holotype. QM G300020 (ORSTOM R1223): stn 184, SE. of Ilbt Ua, New Caledonia lagoon, 22'42. l'S., 166"49+WE.,16 m depth, 8.vi.1977, coll. G. Bargibant, SCUBA. Description Colour. Pale orange alive (Munsell 7.5YR 8/10), pale grey-brown in ethanol. Shape. Thickly encrusting plate, 260 mm diameter, 10-35 mm thick, with prominent low, conical-bulbous, digitate projections on upper surface; bulbous digits 14-23 mm high, 10-14 mm maximum diameter, each with a single, large osculum on apex, 3-5 mm diameter. Surface. Membraneous (where intact in preserved condition), even, porous upper surface, without ornamentation other than large bulbous digits. Subdermal spicule bundles clearly visible below translucent dermal membrane. Ectosome. Without specialised spiculation; with plumose brushes composed of choanosoma1 megascleres protruding slightly through surface, up to about 80 pm from surface, but presumably these are confined to within the dermal membrane when intact. Mesohyl in peripheral skeleton with heavier, granular collagen as compared with the deeper choanosomal region. Choanosome. Skeletal structure plumo-reticulate, without axial compression or differentiation between axial and extra-axial regions. Skeleton divided into ascending, multispicular, diverging primary skeletal tracts, up to 150 pm diameter, with well-developed plumose brushes at surface, and transverse, paucispicular, secondary spicule tracts, 2040 pm diameter, more-or-less interconnecting primary tracts. Skeletal meshes 150-300 pm diameter, with few loose spicules between tracts. Spongin fibres very lightly invested with Fig. 7. Reniochalina condylia, sp. nov., holotype QM G300020: A, oxeas, styles and modifications; B, section through peripheral skeleton. 1406 J. N. A. Hooper and C. Levi spongin, seen clearly only between major spicule tracts. Choanocyte chambers oval, 80170 pm diameter. Mesohyl with light collagen. Megascleres. Single category of structural megasclere present, varying from oxeas to styles, with various intermediate forms also present. Spicules consist of predominantly slightly curved, slightly asymmetrical oxeas; less frequently styles with evenly rounded bases and slightly curved towards basal end; rarely asymmetrical anisoxeas. Points mostly fusiform, tapering, sharply pointed; occasionally fusiform rounded. Length 208-(259.8)-289 pm, width 10-(12.2)-14 pm. Microscleres. Absent. Distribution Known only from the south-west lagoon of New Caledonia, 16 m depth, on coral rubble substrate. Remarks The present species is most similar to R. stalagmitis in its skeletal structure (plumose spicule bundles) and spicule geometry (oxeas, styles and anisoxeas), although in R. stalagmitis many spicules have spinous extremities, particularly the anisoxeas. Reniochalina condylia, sp. nov., is also very similar to two undescribed species of Reniochalina from western and north-western Australia (Reniochalina provisional species numbers 353, 798), having virtually identical skeletal architecture and spicule geometry, but in both cases these undescribed species have quite different external morphology and surface ornamentation, and there are also differences in the specific dimensions of megascleres (broader and shorter in the two undescribed species). These new species will be contrasted further with other described species in a future revision of Reniochalina. The deeper-water R. plumosa from New Caledonia has a plumo-reticulate skeleton composed mostly of ascending discrete skeletal columns, cored by sharply pointed oxeas incompletely separated into two size categories, 112-(284.21-372 x 9-(11.0)-15 pm (Fig. 6C). Etymology Named for the conical digitate surface projections; Lat. condylus, knob, prominence. Fig. 8. Reniochalina condylia, sp. nov.: A, SEM skeletal structure [holotype QM G300020 (ORSTOM R1223)l (scale=500 pm); B, holotype; C, holotypeh situ (photo P. Laboute). New Caledonian Axinellids 1407 Genus Axinella Schmidt Axinella Schmidt, 1862: 60.-Gray, 1867: 513; Ridley and Dendy, 1887: 178; Dendy, 1905: 193; Dendy, 1922: 114; Vosmaer, 1912: 308,318; Topsent, 1928: 37-8; Bergquist, 1970: 14; Bergquist, 1978: 167, 192; Lbvi, 1973: 605; Pansini, 1983: 79-98 [type species Axinellapolypoides Schmidt, 1862: 62 (holotype possibly SM, 'schizotypes' MNHN DCL1148L, BMNH 1867.7.26.81) (de Laubenfels 1936)l. Chalinissa Lendenfeld, 1887: 771.-Burton, 1927: 502 (type species Isodictya dissirnilis Bowerbank, 1866) (de Laubenfels 1936). Astrospongia Gray, 1867: 514 (objective synonym of Axinella). Diagnosis Variable growth form, ranging from digitate to flabellate; surface typically hispid, conulose; choanosome always with some axial compression o f spongin fibres and spicules; with or without differentiated primary and secondary fibre elements; extra-axial skeleton plumose or plumo-reticulate, diverging single or bundles o f spicules; ectosome without specialised spiculation, although extra-axial megascleres usually protrude through surface; megascleres oxeas, styles, occasionally strongyles, in various combinations; microscleres may include raphides or microraphides, although these do not appear to be widely distributed within the genus [compiled from Vosmaer (1912) and Donadey et al. (1990)l. Remarks Schmidt (1862) characterised A. polypoides in having a peculiar distribution o f oscules, for which Gray (1867) erected Astrospongia, but Vosmaer (1933, 1935a, 1935b) correctly surmised that the character was unlikely to be o f much systematic importance, and in any case he noted that it also occurred in other axinellid genera. Other authors who have dealt with this genus (e.g. Ridley and Dendy 1887; Thiele 1903; Vosmaer 1912, 1935a; Dendy 1922; Babic 1922; Topsent 1934; Vacelet 1961, 1969; Bergquist 1970; Pansini 1983) agree that it is still problematic, with no completely satisfactory or clearly discriminatory diagnosis yet developed. Part o f the problem arises from the existence o f species with predominantly stylote spiculation (e.g. Dendy 1922), whereas typically the genus has both oxeas and styles o f equal size and proportion (e.g. Vosmaer 1935a; Wiedenmayer 1989). Vosmaer (1935a) and Pansini (1983) suggest further that intra-specific variability is known for the type and other nominal Axinella species, whereby both oxeas and styles may be modified to anisoxeote or anisostrongyle forms, in some cases almost completely. Consequently, the placement o f species with a reticulate axially condensed skeleton, and reticulate to plumoreticulate extra-axial skeleton o f oxeas and styles in Axinella is relatively easy (e.g. A . polypoides; Fig. lOC), whereas species such as A . spiculifera (Lamarck), A . profunda Ridley and Dendy, and A . erecta Carter have greatly modified spiculation, and could be easily placed in genera such as Phakellia or Reniochalina. The literature contains records o f 39 species from the Australasian region (including New Zealand, New Caledonia, Papua New Guinea and southern Indonesia), that have been referred to Axinella at one time or another but, o f these, only 18 appear to be correctly placed here. The type material o f many o f these species has not yet been located, so a specific revision o f the Indo-west Pacific fauna in this genus is not yet possible. However, valid species o f Axinella from this region are presently thought to be: Phakellia aruensis Hentschel, 1912 (holotype SMF 953; Fig. 9A-B), from southern Indonesia, north-west Australia around to the Great Barrier Reef (Hentschel 1912; Bergquist et al. 1980; Hooper et al. 1992); Axinella australiensis Bergquist, 1970 (holotype NMNZ Por .18 [not seen]), from New Zealand; Axinella brondstedi Bergquist, 1970 (holotype probably ZMC [not seen]),with synonym Axinella verrucosa sensu Brondsted, 1923 (preocc.) from New Zealand; Acanthella carteri Dendy, 1889, from New Caledonia (LCvi 1979) (Fig. 11-12; see below); Axinella clathrata Dendy, 1897a (holotype NMV G2330 [not seen]), from south-east Australia; Phakellia crassa Carter, 1885 (holotype BMNH 1886.12.15.129), from southeast Australia (Fig. 9C-D); Acanthella euctimena Hentschel, 1912 (holotype SMF 1012; Fig. 9E-F) from southern Indonesia; Axinella Iifouensis LBvi & LBvi, 1983 (holotype 1408 J. N. A. Hooper and C. Levi MNHN DCL2943; Fig. 9G-H), from deeper-waters off New Caledonia; Axinella globula Brondsted, 1924 (holotype probably ZMC [not seen]), from New Zealand (Bergquist 1970); Axinella kirkii Dendy, 1897a (lectotype NMV G2370 [not seen]), from south-east Australia; Axinella meloniformis Carter, 1885 (holotype BMNH 1886.12.15.117; Fig. 91-J), from south-east Australia (considered atypical of the genus [Vosmaer 19121, but skeletal structure is most similar to Axinella of all other genera); Axinella retepora (Lendenfeld, 1887) (lectotype BMNH 1886.8.27.417; Fig. 10A-B) from Torres Strait, Qld (not Port Phillip, Vic., as published); Axinella richardsoni Bergquist, 1970 (holotype NMNZ Por. l9 [not seen]), from New Zealand; Axinella sinclarii (Gray) (holotype unknown) from New Zealand (Dendy 1897b; Bergquist 1970); Axinella torquata Brondsted, 1923 (holotype probably ZMC Fig. 9 . Axinella spp.: A , Phakellia aruensis Hentschel, in situ (specimen NTM 22156) (photo J. N. A. Hooper); B, skeleton (holotype SMF 953) (scale=500 pm); C, Phakellia crassa Carter (holotype BMNH 1886.12.15.129); D, skeleton (scale = 500 pm); E, Acanthella euctimena Hentschel (holotype SMF 1012); F, skeleton (scale = 500 pm); G, Axinella lifouensis Ldvi and L6vi (holotype MNHN DCL2943); H, skeleton (scale=500 pm); I, Axinella meloniformis Carter (holotype BMNH 1886.12.15.117); J, skeleton (scale = 500 pm). New Caledonian Axinellids 1409 [not seen]), from New Zealand (Bergquist 1970); Axinella tricalyciforrnis Bergquist, 1970 (holotype CMMI 3/63 [not seen]), with synonym Axinella larnellata sensu Bergquist, 1961, from New Zealand; and Axinella villosa Carter, 1885 ('syntype' BMNH 1886.12.15.396; Fig. lOD), from south-east Australia (Dendy 1897a). There are another 28 possibly undescribed species collected from tropical Australasia (Hooper, unpublished data), which are also probably most appropriately placed in this genus. These will be described in future revision of the tropical Australasian axinellids. Other Australasian species described in Axinella but better placed elsewhere include: Axinella aurantiaca Lendenfeld (1888, 'syntypes' AM G9171,Z468) from south-east Australia belongs with Bubaris (Bubaridae) (Whitelegge 1889, 1901; Hallmann 1914a; Burton 1928); Axinella arborescens Ridley & Dendy, 1886 (holotype BMNH 1887.5.2.64), from south-east Australia belongs in Hornaxinella (Whitelegge 1889, 1907; Hallmann 1914~;Vacelet et al. 1976); Axinella atropurpurea Carter, 1885 (holotype BMNH 1886.12.15.I), from southeast Australia is a species of Raspailia (Raspailiidae) (Hooper 1991); Axinella axifera Hentschel, 1912 ('syntype' SMF 1666), from southern Indonesia and north-west Australia is a species of Ceratopsion (Raspailiidae) (Hooper 1991); Axinella colvilli Brondsted, 1924 ('syntypes' probably in ZMC), from New Zealand belongs to Ciocalypta (Halichondriidae) (Bergquist 1970); Axinella chalinoides et varr. Carter, 1885 (lectotype BMNH 1886.12.15.401), and Axinella cladoflagellata Carter, 1886a (holotype BMNH 1886.12.15.407), from southeast Australia, are synonymous and belong to Echinoclathria (Microcionidae) (comb. nov., determined from re-examination of the type material); Axinella ? coccinea Carter, 1886a (holotype BMNH 1886.12.15.8), from south-east Australia is an Zotrochota (Desmacididae) (comb. nov.); Axinellaflabellata Carter, 1885 ('syntypes' BMNH 1886.12.15.471, 143), from south-east Australia belongs to Sigrnaxia (Desmacellidae) (Hooper 1984); Axinella frondula Whitelegge, 1907 (holotype AM G4349), from south-east Australia is a species of Raspailia (Hooper 1991); Axinella hispida var. gracilis Lendenfeld, 1888 (holotype AM G9083), and var. tenella Lendenfeld, 1888 (lectotype AM G9074), are species of Raspailia (Hooper 1991); Axinella inflata Lendenfeld, 1888 (holotype AM G9081), from south-east Australia is virtually unrecognisable (but possibly belonging to Dictyodendrillidae; F. Wiedenmayer, Fig. 10. Axinella spp.: A , Chalinopora retepora Lendenfeld (lectotype BMNH 1886.8.27.417); B, skeleton (scale =500 pm); C, Axinella polypoides Schmidt, skeleton (specimen BMNH 1925. 1 1.1.1003 from Naples) (scale = 1 mm); D, Axinella villosa Carter, skeleton ('syntype' BMNH 1886.12.15.396) (scale = 1 mm). 1410 J. N. A. Hooper and C. LCvi personal communication);Axinella labyrinthica Dendy, 1889 (holotype BMNH 1889.1.21.3), from the Indian Ocean and north-west Australia belongs to Teichaxinella (comb. nov.); Axinella obtusa Lendenfeld, 1888 (holotype AM G9082, presently missing), from south-east Australia is unrecognisable;Axinella pilifera Carter, 1885 (holotype unknown), from southeast Australia is virtually unrecognisable; Axinella setacea Carter, 1885 (holotype BMNH 1886.12.15.61) belongs to Raspailia and is a synonym o f R. pinnatifda (Carter) (Hooper 1991); Axinella solida Carter, 1885 (lectotype BMNH 1887.7.11.24), from south-east Australia (Dendy 1897a) is a species o f Rhaphoxya (comb. nov.; Fig. 32C-D); Axinella stelliderma Carter, 1885 (lectotype BMNH 1886.12.15.33), and var. acerata Carter, 1885 ('holotype' BMNH 1886.12.15.63), from south-east Australia (Dendy 1897a) are species o f Raspailia (Hooper 1991);Axinella symbiotica Whitelegge, 1907 (holotype AM G4350), from south-east Australia is the type species o f Axinosia (Hallmann 1914b);Axinella vermiculata Whitelegge, 1907 (holotype AM G4360), from south-east Australia belongs in Teichaxinella (de Laubenfels 1936); Axinella virgultosa var. massa Carter, 1886b (holotype BMNH 1889.6.9.4; Fig. 20F-G), from the central Indian Ocean, south-east Asia and north-west Australia, with synonym Suberites mollis Kieschnick (1898, 1900), belongs to Stylissa (see below). Also, re-examination o f material described as 'Axinella echidnaea' by Ridley (1884) and Hentschel (l912), from north-west Australia and southern Indonesia respectively, shows that they refer to Reniochalina stalagmitis (see above). Axinella carteri (Dendy) (Figs 11-12, Table 2) Acanthella carteri Dendy, 1889: 93-4, pl. 4, fig. 6; Dendy, 1905: 193, pl. 8, fig. 6; Dendy, 1922: 119, pl. 5, fig. 5; Vacelet et al., 1976: 43-4, pl. 2, fig. b; van Soest, 1989: 223-4, fig. 14. Axinella carteri. -Burton, 1959: 258-9; LCvi, 1979: 311. Acanthella aurantiaca Keller, 1889: 396.-Topsent, 1906: 562; Row, 1911: 356; Dendy, 1922: 119; Vacelet and Vasseur, 1971: 80. Material Examined Holotype. BMNH not seen; holotype of A. aurantiaca, ZMB 2921. New Caledonian material. ORSTOM R1056: stn 241, Pointe Nind'Hio, Hienghene, 20'41 .OIS., 164"57.2'E., 10 m depth, 6.ix.1978, coll. P. Laboute, SCUBA; ORSTOM R01: stn 153, barrier reef, Ilat Taenia, Passe de St Vincent, 22'00. 11S.,165"56.llE., 7 m depth, 14.ix.1976, coll. P. Laboute, SCUBA; ORSTOM 'cfRO1': stn 253, barrier reef S. of pass, YatC, 22°11~2'S.,167002~1'E.,25 m depth, 26.ix.1979, coll. G. Bargibant, SCUBA; ORSTOM R1429: stn 332, W. entrance Canal Woodin, Table 2. Comparison in spicule dimensions between published and present records of Axinella carteri All measurements given in micrometres, and expressed as minimum-(mean)maximum range of measurement. N=25 for each specimen MateriaVauthor (Locality) Holotype (Dendy 1889) (Gulf of Manaar) Keller 1889 (Red Sea) Vacelet et al. 1976 (Madagascar) Present study (Comores) N = 1 (Papua New Guinea) N = 1 (Great Barrier Reef) N = 1 (New Caledonia) N = 4 Styles Length Width New Caledonian Axinellids 1411 22°23.1'S.,166046.1'E., 18 m depth, 9.ii.1983, coll. G. Bargibant, SCUBA; NTM 23883 (ORSTOM R170): stn 136, channel between I1Bt Canard and IlBt Maitre, 22°19~2'S.,166021.1'E., 12 m depth, l 6 . i ~198 . 1, coll. J.-L.Menou, SCUBA; NTM 23890: Ile aux Canards, Noumea lagoon, 22Ol9.2'S., 166"26'E., 22 m depth, 26.ix.1990, coll. J. N. A. Hooper, SCUBA; QM (3301323: CroisantLarkgritre, Ilat Maitre, off Noumea, 22°202'S.,166"22.5tE., 20 m depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA. Comparative material. Great Barrier Reef, Qld: QM G300291 (fragment NTM 24027): E. side of Magra I., Cockburn Is, Cape York Region, 11°52.0'S.,143017~O'E.,12.5 m depth, 10.xii.1990, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', SCUBA, (stn JH-90-051). Papua New Guinea: NCI Q66C-4329-V (fragment QM G300355): Cement Mixer Reef, E. of Christiansen Research Institute, Madang, 6°40.0'S.,145049~0'E., 6 m depth, 3.ix.1990, coll. NCI, SCUBA. Western Indian Ocean: QM G301000: Comoros Is, 12"'S.,44"'E., 37 m depth, 1991, coll. G. R. Pettit et al., SCUBA (ref. no. M5351). Description Colour. Bright orange-brown alive (Munsell2.5YR 7/8), pale orange-brown in ethanol. Surface slightly darker than interior of sponge. Shape. Flabellate growth form, 110-400 mm high, with massive, lobate, irregularly planar or globular branching, up to 350 mm wide, and branches consist of relatively thick, flattened planar or buttressed lamellae, 4-11 mm thick, with irregular margins; sponge attached to substrate directly or by small basal stalk, 20-90 mm long, up to 40 mm diameter. Surface. Fleshy, conulose, rough surface; conules irregular, 3-5 mm high, solitary or joined to form meandering surface ridges; texture rubbery, compressible, easily torn. Small oscules, 2-5 mm diameter, occur predominantly on the margins of lamellae, usually between surface conules. Ectosome. Membraneous band of heavy collagen, 100-200 pn wide, slightly more darkly pigmented than choanosomal region, with extra-axial spicules protruding only sparsely. A Fig. 11. Axinella carteri Dendy, specimen QM G300291: A , structural styles; B, section through peripheral skeleton. 1412 J . N. A. Hooper and C. LCvi Choanosome. Axial skeleton condensed into several multispicular bundles, 100-250 pm wide, running more-or-less longitudinally through lamellae, fully cored by long slender styles, bound together by very light spongin fibre, and interconnected at irregular angles by vaguely plumose, ascending, paucispicular extra-axial tracts of styles or individual spicules. Fibre reticulation relatively close meshed, with lacunae forming elongate oval chambers, up to 300 x 70 pm. Collagen in mesohyl relatively light compared with ectosomal region. Megascleres (refer to Table 2 for dimensions). Single category of style present, although variable in thickness, occasionally strongylote; styles relatively long, slender or robust, usually slightly curved symmetrically or towards basal end, sharply pointed, fusiform, with evenly rounded base. Microscleres. Absent. Distribution Red Sea (Keller 1889; Row 1911), Arabian Gulf (Burton 1959), western Indian Ocean: Cargados Carajos, Diego Garcia, Amirante, Salomon, Seychelles Is, Comores, Madagascar (Dendy 1922; Vacelet et al. 1976; present study), Gulf of Manaar, Sri Lanka (Dendy 1889, 1905), Lesser Sumba region, Indonesia (van Soest 1989), Papua New Guinea and the Great Barrier Reef (present study), and New Caledonia (LCvi 1979; present study). This species is a common component of Indo-Pacific coral reefs, usually found in both the lagoon and Fig. 12. Axinella carteri Dendy: A , specimen ORSTOM R170 in situ (photo P . Laboute); B, specimen QM G300291;C, SEM skeleton (scale= 500 pm). New Caledonian Axinellids 1413 outer reef slope; usually associated with living and dead coral. The species appears to be most common in areas of strong current, attached to coral rubble or rock, in sand and sea grass beds. Known depth range is 6-37 m. Remarks None of the material described above had a second category of longer style present, as described in Dendy's (1889) original description (cited as 1200x 11 pm, compared with typical spicules of 400 x 21 pm), although Burton (1959) suggests that this longer spicule is present in few of the many known specimens of A. carteri. Apart from this feature, the species appears to be fairly homogeneous in skeletal structure, spicule morphology and size across its wide geographic range, although spicule dimensions vary slightly between material (Table 2). The species is readily recognisable in the field by its bright orange-brown coloration, fleshy surface and thick irregular lamellae. Axinella carteri is difficult to place with absolute confidence in any single genus, given the relatively unclear boundaries between several structurally similar axinellid genera (e.g. Phakellia, Acanthella, Axinella, Teichaxinella). Vosmaer's (1912) criteria for differentiation of these taxa are not entirely clear cut, because many more species have been described since that early revision, showing that many intermediate (transformation) states are possible between the skeleton plans he initially proposed. Nevertheless, under his scheme for differentiating such genera, the present species falls closest to Axinella in having some axial compression and sparsely diverging, plumose extra-axial spicules. Genus Phakellia Bowerbank Phakellia Bowerbank, 1864: 186.-Bowerbank, 1866: 122; Schmidt, 1866: 15; Gray, 1867: 516; Ridley and Dendy, 1887: 169; Dendy, 1905: 190; Dendy, 1922: 116; Vosmaer, 1912: 310; Topsent, 1928: 37; de Laubenfels, 1936: 130; Bergquist, 1970: 17 [type species Spongia ventilabra Linnaeus, 1767: 1296, which is a junior synonym of Phakellia strigosa (Pallas, 1766) (Vosmaer 1912: 310)l. Querciclona de Laubenfels, 1936: 46 [comb. nov.] (type species Antherochalina quercifolia Keller, 1889: 383; holotype ZMB 429). Diagnosis Compressed flabellate or cup-like growth forms predominant; surface smooth or microconulose; oscules frequently surrounded by subdermal drainage canals; ectosome membraneous without specialised skeleton, often lightly hispid from protruding choanosomal megascleres; choanosomal axial skeleton dense, typically with only interwoven styles, or may include sinuous strongyles and styles, or only strongyles, organised into multispicular ascending and paucispicular transverse tracts, together forming compressed axial reticulation; extra-axial skeleton comparatively sparse, with plumose bundles or single styles or oxeas perpendicular to axis, with or without transverse connecting megascleres; megascleres styles, sinuous strongyles or oxeas; microscleres absent (modified from Vosmaer 1912). Remarks Phakellia is similar to, and often confused with, Acanthella, differing primarily in having a well-differentiated axial and extra-axial skeleton (Vosmaer 1912; Bergquist 1970). Querciclona de Laubenfels (1936: 46) is an obvious synonym of Phakellia, although de Laubenfels initially placed the taxon in Microcionidae. Pararhaphoxya Burton (1934) may also eventually be merged with Phakellia, but is presently maintained separately and distinguished only by the possession of asymmetrical megascleres. However, from examination of nearly 40 species of both Acantheua and Phakellia from the Indo-Pacific faunas, consisting of both described and undescribed taxa, it is tentatively suggested that the structure of axial and extra-axial skeletons is the only feature that consistently differentiates these genera, lending support to Vosmaer's (1912) original proposal. By comparison, the presence or absence of sinuous axial megascleres seems to be much less stable and appears to occur indiscriminantly in species of both genera, and these features must be used with more caution. 1414 J. N. A. Hooper and C. Ldvi In total, 21 species from the Australasian region (including New Zealand, New Caledonia and southern Indonesia) have been described in, or subsequently referred to, Phakellia, but examination of relevant type material suggests that only seven of these are most appropriately placed here. Approximately 20 other, undescribed Phakellia species have also been collected from tropical Australia, but these will be described elsewhere at a later date. Valid Australasian species of Phakellia include: Phakellia carduus (Lamarck, 1814; holotype MNHN LBIM DT533) from north-west and north-east Australia (Ridley 1884; Fig. 13. Phakellia spp.: A , Acanthella cavernosa Dendy (holotype BMNH 1921.11.7.100);B, specimen on deck (NTM 22736); C, skeleton (holotype) (scale = 500 pm); D, peripheral skeleton (scale=200 pm); E, Spongia carduus Lamarck (specimen on deck NTM 21837); F, skeleton (paralectotype MNHN DT3447) (scale=500 pm); G, paralectotype MNHN DT533); H, specimen on deck (QM G300431). New Caledonian Axinellids 1415 Topsent 1930; Bergquist 1970; Hooper, unpublished data) (Fig. 13E-H); Phakellia cavernosa (Dendy, 1922; holotype BMNH 1921.11.7.100) [with synonyms Acanthella stipitata, in part (variety o f Ridley and Dendy, 1887; not Acanthella stipitata Carter, 1881; 'holotype' BMNH 1887.5.2.73), and Burton, 1934 (specimen BMNH l930.8.13.142), and Acanthella klethra Pulitzer-Finali, 1982 (holotype MHNG 46927), from the Indian Ocean, north-west and north-east Australia, and the Great Barrier Reef (Burton 1934; Bergquist 1970; Hooper, unpublished data) (Fig. 13A-D)]; Phakellia columnata (Burton, 1928; holotype BMNH [not found]; 'representative specimen' MNHN DCL2934), known from the Andaman Sea and deeper waters o f f New Caledonia (LCvi and LCvi 1983) (Fig. 14A-B); Phakellia conulosa Dendy, 1922 (holotype BMNH [not found]; 'representative specimen' from the Philippines QM G300315), from the Indian Ocean and north-west Australia (Dendy 1922; Hooper, unpublished data) (Fig. 14C-D); Phakellia dendyi Bergquist (1970;holotype NMNZ Porif .24) from north-west, north-east and south-east Australia, and New Zealand (Dendy 1897b; Bergquist 1970; Hooper, unpublished data) (Fig. 14E-F); Phakelliapulcherrima (Ridley and Dendy, 1886; holotype BMNH 1887.5.2.23) (Figs 14G-H, 15-16) (see below); and Phakellia stipitata (Carter, 1881; holotype BMNH [confirmeddestroyed];neotype QM G300874, here designated) from eastern and south-east Australia (see below) (Fig. 17-18). Other Australasian species described in Phakellia but better placed elsewhere include: Phakellia aruensis Hentschel, 1912 (holotype SMF 953), from southern Indonesia is assigned to Axinella; Phakellia brassicata Carter, 1885 (holotype BMNH 1886.12.15.75), from southeast and north-east Australia belongs to Cribrochalina (Niphatidae, order Haplosclerida); Phakellia crassa Carter, 1885 (holotype BMNH 1886.12.15.129), from south-east Australia (Fig. 9C-D) is an Axinella; Phakellia flabellata Carter, 1885 (holotype BMNH 1886. 12.15.471), and Phakellia villosa Carter, 1886a (lectotype BMNH 1886.12.15.78), both from south-east Australia, are both synonyms o f Teichaxinellaflabellata (Carter, 1885);Phakellia inj7exa Pulitzer-Finali, 1982 (holotype MHNG 46931 [not seen by the authors]), from the southern Great Barrier Reef is an Acanthella; Phakellia jacksoniana Dendy, 1897a (lectotype BMNH 1887.5.2.9), is a synonym o f Clathria (Isociella)jacksoniana (Dendy)(Microcionidae, order Poecilosclerida);Phakellia multiformis Whitelegge, 1907 ('syntype' AM G4358), from south-east Australia belongs to Axinosia; Phakellia papyracea Carter, 1886a (lectotype BMNH 1886.12.15.231), from south-east Australia is a synonym o f Echinoclathria leporina (Lamarck, 1814) (Microcionidae, order Poecilosclerida); Phakellia plumosa LCvi & LCvi, 1983 (holotype MNHN DCL2972), from deeper waters o f f New Caledonia belongs in Reniochalina (Fig. 6C-D); Phakellia ramosa Carter, 1883 (holotype BMNH 1884.4.14.2), from south-east Australia belongs to Sigmaxinella (Desmacellidae, order Poecilosclerida); Phakellia tumida Dendy, 1897a (holotype NMV G2464), from south-east Australia belongs to Pseudaxinella; and Phakellia ventilabrum australiensis Carter, 1886a (holotype BMNH 1886.12.15.422), from south-east Australia is a synonym o f Clathria (Thalysias) cactiformis (Lamarck, 18 14) (Microcionidae, order Poecilosclerida). Phakellia pulcherrima (Ridley & Dendy), comb. nov. (Figs 14G-H, 15-16, Table 3) Acanthella sp. -Ridley, 1884: 463. Acanthella pulcherrima Ridley & Dendy, 1886: 218, 479.-Ridley and Dendy, 1887: 177, pl. 32, fig. 3. Not Acanthella pulcherrima. -Capon and MacLeod, 1988: 979-83. Acanthella pulcherrima calyx Dendy, 1922: 120, pl. 5, fig. 6. Material Examined Holotype. BMNH 1921.11.7.100 (fragment MNHN DCL209L): Prince of Wales Channel, Torres Strait, 14 m depth, HMS 'Challenger'. New Caledonian material. Q M G300019 (ORSTOM R1321): stn 102, mid-channel, Canal Woodin, 22"23.1'S.,166°48.1'E.,28.iv.1976, 33 m depth, coll. P. Laboute, SCUBA; ORSTOM R1052: stn 136, channel between Il6t Canard and Ilat Maitre, 22°19~2'S.,166021.1'E.,20 m depth, 16.iv.1981, coll. P. Laboute, SCUBA. J . N. A. Hooper and C. Ltvi 1416 Description Colour. Pale orange-brown or brown alive (Munsell 5YR 7/10), pale beige in ethanol. Shape. Small club-shaped sponge (75-120 mm long, 58-75 mm maximum breadth), with short cylindrical stalk (12-22 mm long, 7-10 mm wide), enlarged basal holdfast, and several very thin, leaf-like, flattened branches (10-30 mm maximum width, 4-8 mm thick), with even margins. Fig. 14. Phakellia spp.: A , Bubaris columnata Burton (specimen from New Caledonia MNHN DCL2934); B, skeleton (scale= 500 pm); C, Phakellia conulosa Dendy, in situ (specimen from the Philippines, QM G300315) (photo J . N. A. Hooper); D, skeleton (scale=200 pm); E, Phakellia dendyi Bergquist (specimen on deck NTM 21318); F, skeleton (scale=500 pm); G, Acanthella pulcherrima Ridley and Dendy (holotype BMNH 1887.5.2.23); H, skeleton (scale = 500 pm). New Caledonian Axinellids 1417 Table 3. Comparison in spicule dimensions between published and present records of Phakellia pulcherrima All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of measurement. N=25 for each specimen MateriaYauthor (Locality) Holotype (BMNH 1887.5.2.23) Dendy (1922) (Cargados Carajos) Present material (New Caledonia) L W L W L W Extra-axial styles Axial strongyles 294-(331.2)-392 ~ 6 - ( 1 1.O)-15 'Larger' 360-(577 1)-940 x 2-(5.2)-9 Up to 1300 x 8.6 449-(488.4-552 x 2-(4.2)-6 253-(360.3)-413 x 2-(5.4)-9 Fig. 15. Phakellia pulcherrima (Ridley and Dendy), specimen QM G300019: A , extra-axial styles; B, axial strongyles; C , anisoxeas frequent in holotype BMNH 1887.5.2.23; D, section through peripheral skeleton of specimen QM G300019. 1418 J. N. A. Hooper and C. L&i Surface. Evenly conulose; conules rounded or pointed, more-or-less arranged in ridges, up to 5 mm high, running longitudinally along branches, with ridges producing an almost striated pattern. Oscules and pores not seen in preserved material. Texture firm, flexible. Ectosome. Membraneous, fleshy, without speciaiised spicules; points of extra-axial styles protrude 100-250 pm from surface conules, whereas between/conules ectosome is merely fleshy, with more darkly pigmented granular collagen than in choanosomal region. Choanosome. Skeleton divided into distinct axial and extra-axial regions. Axis tightly compressed, occupying only about 30% of branch diameter, running longitudinally through branches, cored by closely reticulate sinuous strongyles, more-or-less interlocked and crisscrossed within axis. Extra-axial skeleton consists of radial tracts of individual, or loosely plumose bundles of styles, embedded in and standing perpendicular to the axis. Axial spicules bound by moderately heavy, close-set spongin fibres, up to 30 pm diameter, with fibre reticulation producing elongate meshes, 20-70 pm diameter. Extra-axial spicules free within mesohyl, not associated with spongin fibres except where embedded into axis. No choanocyte chambers observed in peripheral skeleton. Megascleres (refer to Table 3 for dimensions). Extra-axial styles relatively thick, straight or slightly curved near basal end, with tapering fusiform points and with evenly rounded or slightly constricted bases. Axial strongyles usually sinuous, occasionally completely straight or vermiform, relatively thick, with evenly rounded bases. Microscleres. Absent. Distribution New Caledonian lagoon, Cape York, Torres Strait, Great Barrier Reef, Cargados Carajos, Indian Ocean; 5-54 m depth; coral rubble. Fig. 16. Phakellia pulcherrima (Ridley and Dendy): A , specimen in situ (ORSTOM R1052) (photo P. Laboute); B, specimen QM G300019; C, SEM skeleton (magnified 80 x); D, SEM transverse section (magnified 94 x ) . New Caledonian Axinellids 1419 Remarks The New Caledonian population of P. pulcherrima is a new locality record for the species, and is the first validated record (i.e. confirmed by voucher specimen) since Dendy (1922) rediscovered the species from the western Indian Ocean. A major difference between the New Caledonian specimen and the two previously known records of the species is that many of the extra-axial styles have modified oxeote bases in the older specimens, whereas in the New Caledonian specimen these spicules are invariably styloid. Spicule dimensions also vary slightly, particularly in the maximum lengths of strongyles (Table 3), but considering that these spicules break up easily during histological preparation it is possible that they may actually be larger in New Caledonian material than described here. In other details (growth form, surface sculpturing, skeletal structure), all known specimens are similar and undoubtedly conspecific. This species is contrasted with other Australasian Phakellia in Figs 13-14. Phakellia stipitata (Carter) (Figs 17-18) Acanthella stipitata Carter, 1881: 380, pl. 18, fig. 8. Not Acanthella stipitata.-Ridley and Dendy, 1887: 178; Burton, 1934: 565. Material Examined Neotype. QM G300874: Peel I., Lazarette Gutter, Moreton Bay, Qld, 27"28.9'S.,l53"21.4'E., 8 m depth, 14.iv.1992, coll. J . N. A. Hooper and S. D. Cook, SCUBA. New Caledonian material. QM G 300280 (fragment NTM 23888): Ile aux Canards, Noumea lagoon, 22°19.0'S.,166026.0'E.,22 m depth, 26.ix.1990, coll. J. N. A. Hooper and J . Vacelet, SCUBA, sand, rubble, sea grass beds; ORSTOM R1429 (fragment NTM 23875): stn 332, W. entrance Canal Woodin, 22"23.1'S.,166"46.1'E., 18 m depth, 9.ii.1983, coll. J.-L. Menou, SCUBA. Comparative material. QM G301235: same locality as neotype. Description Colour. Bright orange-brown alive (Munsell 5YR 7/10), orange-yellow in ethanol. Shape. Flabellate sponge, 90-130 mm long, 70-110 mm wide, 40-90 mm thick, with one or more fans aligned face-to-face, composed of irregularly fused and reticulated branches, with excavated wide meshes between reticulations, producing thick, nearly bulbous branching; small basal stalk, up to 30 mm long, 15-20 mm diameter (often detached from specimen), with a broad basal holdfast. Surface. Clathrous, excavated surface, with well-developed, regularly spaced conules, 10-20 mm apart, 5-20 mm high; conules have rounded tips, usually joined together by low ridges, surrounding large excavations through the sponge (4-15 mm diameter) (i.e. producing the surface meshes); remainder of membraneous surface with 'goose-flesh' appearance, and covered with small ostia (< 1 mm diameter). Large oscules on margins of fans, 15-25 mm diameter, slightly raised above surface, and slightly more darkly pigmented than rest of surface. Texture firm, rubbery, difficult to tear, usually requires cutting off substrate. Ectosome. Membraneous, heavily collagenous, darkly pigmented, without special spicules and only points of extra-axial styles barely protruding through surface (these usually only on ends of surface conules). Choanosome. Skeleton clearly differentiated into axial and extra-axial regions. Axis moderately compressed, with short, heavy, reticulated spongin fibres, 20-45 pm diameter, producing oval or elongate meshes, 60-110 pm diameter; fibres only partially cored by styles in uni- or multispicular tracts. Extra-axial skeleton not well formed (as in some other species of Phakellia), consisting of radially arranged styles, as individuals or multispicular brushes, standing perpendicular or at acute angles to axis, ascending to and usually 1420 J . N. A. Hooper and C. Lkvi protruding through ectosome. Styles in axial and extra-axial skeletons do not appear to be differentiated. Choanocyte chambers small, elongate, up to 40 pm diameter. Megascleres. Axial and extra-axial megascleres are exclusively styles, long, slender, straight or slightly curved, with abrupt points, sharp or slightly stepped tips, and with evenly rounded or occasionally oxeote bases. Length (neotype): 172-(345.5)-494 pm; (New Caledonian material): 301-(412.7)-545 pm. Width (neotype): 3-(7-3-12 pm; (New Caledonian material): 3-(8.1)-15 pm. Microscleres. Absent. Distribution Gulf of Manaar (original record), south-east Queensland, and New Caledonia; associated with living and dead coral, coral reef lagoon. Fig. 17. Phakellia stipitata (Carter):A, neotype QM G300874, structural styles and anisoxeas; B, specimen QM G300280 from New Caledonia, styles and anisoxeas; C, section through peripheral skeleton of neotype. New Caledonian Axinellids 1421 Remarks There is no extant type material of this species in the BMNH or LFM (S. Stone, BMNH, personal communication, and personal observation), nor has the species been recorded since it was first described. Records cited by Ridley and Dendy (1887) and Burton (1934) refer to Phakellia cavernosa (Dendy, 1922), and this is confirmed by the presence of sinuous strongyles in their material (BMNH 1887.5.2.73 and BMNH 1930.8.13.142, respectively). However, some distinctive features of the species are obvious from Carter's (1881) original description, notably the characteristic clathrous, excavated gross morphology, and spicule morphology (particularly spicule size, geometry and presence of abrupt points). These agree closely with the present material from Moreton Bay and the Noumea lagoon. New Caledonian specimens have marginally larger styles than does the neotype from Moreton Bay, but this difference is relatively trivial, and the species are similar in all other features. Fig. 18. Phakellia stipitata (Carter): A , specimen in situ (ORSTOM R1429) (photo J.-L. Menou); B, preserved specimen (QM (3300280); C, three specimens on deck (neotype QM G300874 top left, indicated by arrow; other specimens QM G301235); D, peripheral skeleton (neotype QM G300874) (scale=200 pm); E, SEM transverse section (specimen QM G300280) (scale = 200 pm). 1422 J. N. A. Hooper and C . L6vi Genus Stylissa Hallmann Stylissa Hallmann, 1914b: 349 [type species Stylotellaflabelliformis Hentschel, 1912: 355, holotype SMF 15871. Dragmaxia Hallmann, 1916a: 543.-Hallmann, 1916b: 674; de Laubenfels 1936: 130 [type species Spongosorites variabilis Whitelegge, 1907: 513, holotype AM (presently missing)]. Diagnosis Fan, cup-shaped or massive foliose growth forms; surface shaggy, often with small papillae or grooved ridges; ectosome fleshy, without specialised spiculation, but with brushes of peripheral choanosomal styles protruding through surface; choanosomal skeleton disorganised plumo-reticulate, with a slightly condensed axis and slight differentiation between axial and extra-axial skeletons; fibres cored by parallel tracts of styles, of 1 or 2 sizes, more-or-less ascending, and diverging towards the periphery; peripheral styles often slightly larger than those in axis. Microscleres absent or present (raphides, trichodragmata) (modified from Hallmann 1914b). Remarks Whitelegge (1907) characterised Spongosorites variabilis partly by the presence of raphides in the dermal layer, and Hallmann (1916~)erected Dragmaxia to emphasise this feature, which he generally considered to be diagnostically important throughout his taxonomy. Hallmann also noted that these raphides occurred throughout the mesohyl, not necessarily confined to the ectosomal region, as supposed by Whitelegge (1907). In addition, Hallmann (1916~)noted that although both Dragmaxia and Stylissa have comparable architecture and only styles for megascleres, those of the former were differentiated into two size categories, with some evidence of spicule localisation (the larger apparently found closer to the periphery). However, in other Stylissa spicule size may extend over a large size range, and often the peripheral spicules are marginally larger than those in the axial skeleton. Whitelegge's holotype of S. variabilis is missing from AM collections, and these features cannot be accurately verified, but the species is perfectly recognisable from Hallmann's (1916~:544) redescription, which includes comprehensive illustrations. It is concluded from this available evidence that Dragmaxia differs in no substantial respect from Stylissa and the two are here merged, as was suggested by de Laubenfels (1936). Stylissa presently contains only three valid species, all from the Indo-west Pacific region: the type species S. flabelliformis (Hentschel, 1912), previously known only from the Arafura Sea but now known from the Indo-west Pacific region in general (Fig. 19-20); S. variabilis (Whitelegge, 1907), known only from the Crookhaven River, N.S. W. (Hallmann 1916a) (holotype AM, missing); and another widely distributed species, S. massa (Carter, 1886b) [for Axinella virgultosa var. massa Carter (holotype BMNH 1889.6.9.4; Fig. 20F-G) (Burton 1959), with synonyms Suberites mollis Kieschnick, 1898, and Stylotella conulosa Topsent, 1897 (lectotype MHNG C-12/45, paralectotype MNHN LBIM DT1775, here designated)], extending from the south Arabian coast (Burton 1959), the Mergui Archipelago, Andaman Sea (Carter 1886b), Torres Strait, Qld (Kieschnick, 1898, 1900), Ambon, Moluccas (Topsent 1897; Desqueyroux-Faundez 1981), Ternate, Moluccas (Kieschnick 1898, 1900), Java Sea (Lindgren 1897, 1898), and Christmas Island, western Indian Ocean (Kirkpatrick 1900). Stylissa flabelliformis (Hentschel) (Figs 19-20, Table 4) Stylotella flabelllformis Hentschel, 1912: 355-6, pl. 19, fig. 26. Stylissa flabelliformis. -Hallmann, 1914b: 349. Teichaxinella Iabyrinthica.-Hooper et al., 1992: 265. Material Examined Holotype. SMF 1587 (schizotype MNHN DCL2314): Meriri, Aru I., Arafura Sea, 6"S.,134"501E., 6-10 m depth. New Caledonian Axinellids 1423 New Caledonian material. QM G300017 (ORSTOM R1257): stn 166, N. pass Toemo, Port de Goro, 22°20.0'S.,167001.OIE., 18 m depth, 28.x.1976, coll. A. IntCs, SCUBA; QM (3300689 (fragment NTM 23874): Baie des Citrons, Noumea lagoon, 22°18'S.,166025'E., 3 m depth, 25.ix.1990, coll. J. N. A. Hooper, snorkel; ORSTOM R565: stn 113, L'epave du 'Bonhomme', Grand RCcif Mbttre, 22°21~O'S.,166014~0'E.,25 m depth, 21.vi.1976, coll. P. Laboute, SCUBA. Comparative material. Seychelles Is: QM G300068: E. of Curieuse I., 4"15'S.,55"47'E., depth and date of collection unknown, coll. Pettit, G. R. et al., SCUBA (ref. no. M5281). Japan: QM G301236: Yonaguni I., E. of Taiwan, 24"29'N.,123"00'E., 30 m depth, 1992, coll. T. Higa (ref. K-1). Sahul Shelf, W.A.: QM G300183 (fragment NTM 22817): West I. passage, outer reef, Ashmore Reef, 12"15'S.,122"55'E., 16.5 m depth, 28.vii.1986, coll. C. Johnston, SCUBA; NTM 22797, 2798: West I. passage, outer reef edge, Ashmore Reef, 12°14.3'S.,123056.0'E., 15.5 m depth, 27.vii.1986, coll. A. M. Mussig, SCUBA; QM G301081, G301082: Cartier I., outer reef slope, N. side reef, 12"31.4'S., 123"33.3'E., 14 m depth, 06.v.1992, coll. J. N. A. Hooper, SCUBA. Northwest Shelf, W.A.: NTM 23374: 2 nm from shore, N. of Barrow I., 20°38.8'S.,11548.8'E., 22 m depth, 26.viii.1988, coll. D. Low Choy, SCUBA; NTM 2664: NW. of Yampi Sound, 15"27.04'S.,121°49.00'E.,76 m depth, 29.iv.1982, coll. CSIRO R.V. 'Sprightly', dredge; QM G300108 (fragment NTM 21220), NTM 21236: 40 m depth, 26.iv.1983, coll. N. of Bedout I., W. of Port Hedland, 19°28~05'S.,118055~30'E., J. N. A. Hooper, R.V. 'Soela', S02/83, stn B9, trawl; NTM 21471: W. of Port Hedland, 1g056'S., 117"57'E., 40 m depth, O5.xii.1985, coll. Ward, T., trawl; NTM 21807, 21829: W. of Port Hedland, 19°26~09'S.,118054~02'E., 50 m depth, 30.viii.1983, coll. Ward, T. R.V. 'Soela', trawl; NTM 2684: N. of Port Hedland, 19"16'S.,118"50'E., 70 m depth, 4.v.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 22321: NW. of Lacepede Is, 16°31'S.,121028'E., 38-40 m depth, 17.iv.1985, coll. Russell, B. C., pair trawl; NTM 2731: N. of Adele I., Collier Bay, 15°58.03'S.,122039.07'E., 59 m depth, 21.iv.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 22351: NW. of Lacepede Is, 16"34'S.,121°27'E., 40-46 m depth, 17.iv.1985, coll. B. C. Russell, pair trawl; NTM 23021: N. of Amphinome Shoals, 19"19.7'S., 119O08.8'E., 50 m depth, 19.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', trawl. Arafura Sea, N.T.: NTM 2607: Cootamundra Shoals, N. of Melville I., 10"49~07'S.,129"12~09'E., 31 m depth, 6.v.1982, coll. B. Thom, R. Lockyer, SCUBA; NTM 2616: same locality, 10°50.22'S., 129"13.17'E., 22 m depth, 10.v.1982, coll. R. Lockyer, SCUBA; NTM 23081: Parry Shoals, 11°11~41'S.,129043~01'E., 18 m depth, 13.viii.1987, coll. A. M. Mussig, SCUBA. Description Colour. Live coloration dark orange-brown (Munsell 5YR 7-6/10), with a paler membraneous ectosome, and brighter orange interior (5YR 7/12); red-brown on deck; produces an orange exudate upon collection. Shape. Growth form generally thickly flabellate, 120-450 mm long, 70-180 mm wide, with flabellate-digitate branches growing in more than 1 plane, with 140 mm maximum span, with even or uneven digitate margins on branches, up to 30 mm thick, attached to substrate by small thickly cylindrical basal stalk, 20-75 mm long, 11-45 mm diameter. Surface. Characteristically rough, striated, conulose, shaggy surface, with either longitudinal striations in larger specimens, or irregularly conulose, sculptured surface in younger material. Surface soft, fleshy in life, contracting in preserved material to produce harsh texture. Oscules visible on apex of surface ridges and margins of branches, up to 14 mm diameter, with a large membraneous lip surrounding each exhalant pore. Fleshy parts of surface porous, with evenly dispersed ostia, predominantly between conules and ridges. Ectosome. Fleshy, darkly pigmented, heavily collagenous ectosome, with points of styles from peripheral skeleton protruding up to 200 pm from the surface. In vicinity of surface conules peripheral (subectosomal) skeleton extends directly into subdermal region, whereas in fleshy area between conules there are only few spicules present. Choanosome. Skeletal structure plumo-reticulate, although it appears disorganised due to proportionally large size of megascleres in relation to spongin fibres, with only slightly differentiated axial and extra-axial regions. Axial skeleton reticulate, with heavy spongin fibres (80-170 pm diameter), forming rectangular meshes (80-170 pm diameter), cored by multispicular tracts of styles. Extra-axial skeleton vaguely plumo-reticulate, with multispicular ascending tracts of styles interconnected at irregular intervals by uni- or pauci- 1424 J. N. A. Hooper and C. LCvi Table 4. Comparison in spicule dimensions between published and present records of Stylissa flabellifonnis and other members of the genus All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of measurement. N=25 for each specimen Population Styles Length Stylissa flabelliformis (Hentschel) Indonesia (holotype) New Caledonia (N= 2) Seychelles (N= 1) Japan (N= 1) North-west Australia (N= 13) Atypical specimen, Northwest Shelf, W.A. (QM G300108) Stylissa massa (Carter) Mergui Archipelago (holotype) Stylissa variabilis (Hallmann) Southern New South Wales (Hallmann 1916a) Width 348-(447.1)-522 339-(433.5)-516 343-(423 '6)-482 396-(486.5)-582 341-(444.6)-554 272-(333.9)-380 9-(16.9)-23 6-(15.8)-22 13-(19.7)-27 8-(20.5)-28 7-(17.4)-28 7-(20.1)-32 504-(564.4)-648 11-(17.8)-22 350-900 1000-1 500 33-45 18 Fig. 19. Stylissaflabelliformis (Hentschel): A, specimen QM G300017, styles and strongylote spicules; B, schizotype MNHN DCL2314, structural spicules; C, section through peripheral skeleton of New Caledonian specimen QM G300017. New Caledonian Axinellids 1425 spicular transverse tracts. Fibre reticulation in extra-axial region slightly more cavernous than axis, with meshes up to 300 pm diameter. Collagen abundant; choanocyte chambers small, elongate, 70-90 pm diameter. Megascleres (refer to Table 4 for dimensions). Styles of a single size class, although great variability in thickness; styles predominantly robust, slightly curved near basal end, rarely straight, usually with evenly rounded bases, rarely rhabdose, tapering to fusiform points, and occasionally modified to strongyles. Microscleres. Absent. Distribution Indian Ocean and Indo-west Pacific, known from the Seychelles, south-eastern Indonesia, Arafura Sea, Timor Sea, mid-Western Australia, Japan and New Caledonia. Recorded from Fig. 20. Stylissa flabelliforinis (Hentschel): A , fragment of (?) paratype SMF1690 (schizotype MNHN DCL2314); B, specimen from New Caledonia [ORSTOM R1257 (QM G300017)l; C, specimen on deck from NW. Australia (NTM 21807); D, specimen from Ashmore Reef (QM G300183); E, SEM skeleton (scale = 500 pm); F, Stylissa massa (Carter), skeleton of holotype BMNH 1889.6.9.4 (scale = 500 pm); G, holotype. J. N. A. Hooper and C. L6vi 1426 coral reefs, in lagoon, fringing and patch reefs and outer reef slope, sand and coral rubble, 3-76 m depth. Remarks The New Caledonian population of S. flabelliformis is currently the most easterly known record of the species. The species is probably a prominent member of the Indo-west Pacific fauna, although its known distribution is patchy. Certainly, in the eastern Indian Ocean, from eastern Indonesia to the vicinity of Northwest Cape, Western Australia, the species is relatively abundant in the subtidal to about 50 m depth, whereas its relative abundance outside this zone is unknown. In New Caledonia the species is also moderately common within the lagoon, less abundant on the outer reefs, and from the restricted material examined there were no obvious differences detected between any of the regional populations (see also Table 4). However, one specimen trawled from the Northwest Shelf of Western Australia (QM G300108) was atypical in growth form (being much larger, with even margins on the lobate branches, and having distinct longitudinal surface ridges), and in spicule dimensions (having generally shorter and thicker spicules; Table 4), but these differences are not considered to be significant and in other features it agreed closely with all other known material of this species. Stylissa flabelliformis is closely related to its sympatric sibling species, S. massa, which, from re-examination of the holotype, appears to differ only in growth form (the latter being massive; Fig. 20G), skeletal structure (S. massa has an even more disorganised reticulate skeleton than S. flabelliformis; Fig. 20F), and larger spicule dimensions (Table 4). These features appear to be consistent in all the descriptions of S. massa, and the two species are retained here, but morphologically they are certainly closely related. From Hallmann's (1916~:544) comprehensive redescription of S. variabilis, it appears to be similar in growth form to S. flabelliformis, whereas the skeleton is much more organised, plumose (plumoreticulate, resembling Teichaxinella more than Stylissa), styles are larger and more-or-less divisible into two size classes, and raphides are also present. Genus Ptilocaulis Carter Ptilocaulis Carter, 1883: 321.-Carter, 1884: 130; Topsent, 1928: 37, 172; de Laubenfels, 1936: 127; Wiedenmayer, 1977: 152 [type species Ptilocaulis gracilis Carter, 1883: 321 (holotype BMNH 1845.12.30.1) (de Laubenfels, 1936) (Fig. 21D)l. Plicatella Schmidt, 1864: 39 [preocc.]. -de Laubenfels, 1936: 132 [type species Reniera labyrinthica Schmidt, 1864: 391. Diagnosis Typically erect, cylindrical, digitate, clavate to bushy, occasionally lamellate growth form; surface conulose or with numerous elongate and overlapping papilliform projections, often dividing at the apex. Choanosomal skeleton plumo-reticulate, with clearly differentiated axial and extra-axial components; axial portion of skeleton condensed, composed of irregularly anastomosing, close-set spongin fibres and spicules; extra-axial skeleton plumoreticulate or plumose, with heavy fibres cored by ascending multispicular tracts of megascleres, interconnected by paucispicular transverse tracts forming subisodictyal reticulation (s.s.), or without transverse spicule skeleton and simply with meandering, plumose extraaxial spicule tracts; ectosome fleshy, without specialised spiculation, although plumose brushes of spicules may protrude through surface. Megascleres styles, subtylostyles, anisoxeas or strongyles (usually asymmetrical), sometimes including sinuous forms. Microscleres absent (modified from Wiedenmayer 1977). Remarks De Laubenfels (1936: 127) suggested that Ptilocaulis was characterised by having superficial surface processes, a condensed and plumo-reticulate skeleton rich in spongin, and only styles for megascleres, but these features barely define the taxon and in any case are not completely accurate (i.e. spicules usually include styles, anisoxeas, strongyles, and oxeas, New Caledonian Axinellids 1427 some or all with telescoped ends). Wiedenmayer (1977: 152) suggested that longer spicules in the extra-axial skeleton, larger than those found in the choanosome, could also be used as a discriminatory character for the genus, but this feature is not present in the type species nor in many other described species. Wiedenmayer's observations were based on two species that he placed in Ptilocaulis, one of which was Spongia spiculifera Lamarck (1814) from southern Australia, following Topsent (1932), which had clearly differentiated size classes of ectosomal and choanosomal megascleres but was similar in other characters to other members of the genus, but this feature cannot be construed as being typical of Ptilocaulis. Fig. 21. Ptilocaulis spp.: A , Spongia echidnaea Lamarck (holotype MNHN DT640); B, specimen (MNHN DT3347); C, Topsent's spicule preparation of holotype (scale = 200 ~ m )D, ; Ptilocaulis gracilis Carter, skeleton of holotype (BMNH 1845.12.30.1) (scale=500 pm); E, Spongia spiculifera Lamarck (lectotype MNHN DT3345); F, Topsent's spicule preparation of lectotype (scale = 200 pm); G, section of peripheral skeleton (scale = 500 pm). J. N. A. Hooper and C. Ltvi 1428 The presence of anisoxeas and a plumo-reticulate extra-axial skeleton in some species of Ptilocaulis is also similar to Reniochalina, as noted above. Prior to the present study only four species of Ptilocaulis had been described from the Australasian region (including New Caledonia), although several others are known worldwide (e.g. Wiedenmayer 1977). Australasian species include: Spongia echidnaea Lamarck, 1814, probably from southern Australia (holotype MNHN DT640; Fig. 21A-C) (Topsent 1932; not Ridley 1884); Ptilocaulis fusiformis Ltvi, 1967, from New Caledonia (see below; Figs 22-23); Ptilocaulis rigidus Carter, 1883, from southern Australia (?holotype BMNH 1936.5.16.1) (Thiele 1899; Hallmann 1914~;de Laubenfels 1936); and Spongia spiculifera Lamarck, 1814, from Bass Strait, Vic. (holotype MNHN DT638; Fig. 21E-G) (Ridley 1884; Dendy 1922; Topsent 1932, 1933; de Laubenfels 1936; Wiedenmayer 1977). Ptilocaulis fusiformis Ltvi (Figs 22-23, Table 5) Ptilocaulis fusiformis Levi, 1967: 21, pl. 1 , fig. b, text-fig. 4. Material Examined Holotype. MNHN DCL818: Baie St Vincent, Grand Tenia, New Caledonia, 40 m depth, l0.xii. 1962, coll. Mission Singer-Polignac. New Caledonian material. QM G300719 (ORSTOM R1547): stn 503, Pointe des Pins, Canal Woodin, 22"23.4'S.,166"49.6'E.,25-35 m depth, 11.x.1991, coll. G. Bargibant, SCUBA; QM (3301262 (ORSTOM R854): stn 198, pinnacle S. of Canyon Central, Chenal des Cinq Milles, 22"30.4'S., 166"45.11E.,35 m depth, 15.ii.1978, coll. G. Bargibant, SCUBA; QM (3301324, G301327, (3301335, G301341: Croisant-Laregritre, IlBt Maitre, off Noumea, 22"20.2'S.,166"22.5'E., 20 m depth, 13.x. 1992, coll. J. N. A. Hooper, SCUBA. Description Colour. Live coloration pale orange, yellow-brown (Munsell7.5YR 8/10), pale orangebrown in ethanol. Shape. Digitate or arborescent digitate, 42-110 mm long, with cylindrical bifurcate branches, 23-55 mm long, up to 10 mm diameter, tapering towards ends, and with short basal stalk, 15-19 mm long, 7-10 mm diameter, and broad basal holdfast, 12-21 mm diameter. Surface. Highly conulose, with more-or-less evenly distributed conules, up to 5 mm high, usually forming meandering ridges running longitudinally along branches. Conules interconnected by fleshy surface membrane, pierced by small ostia (visible only between conules), about 500 pm diameter. Oscules small, 1.5-2 mm diameter, rarely observed, near apex of branches. Texture firm, flexible. Ectosome. Fleshy, mostly membraneous surface, with sparse plumose brushes of longer, usually sinuous megascleres barely protruding through ectosome, restricted to tips of conules. Ectosomal membrane highly collagenous, more darkly pigmented than choanosomal mesohyl, and with small quantities of detritus embedded. Choanosome. Skeletal structure plumo-reticulate, with clearly differentiated axial and extra-axial regions. Axial skeleton compressed, composed of a heavy spongin fibre skeleton with individual fibres no more than 150 pm long, up to 50 pm diameter, forming a closeset reticulation producing with oval meshes, 30-90 pm diameter, and cored by plumose, paucispicular tracts of mostly shorter anisoxeas and fewer sinuous strongyles. Extra-axial skeleton corresponds with distribution of surface conules. Extra-axial fibres plumo-reticulate, noticeably more cavernous than in axis, with individual fibres extending for up to 300 pm, 25-35 pm diameter, producing elongate meshes, and extra-axial fibre system runs predominantly laterally through branch cross-section. Fibres cored by multispicular plumose tracts of both sinuous strongyles and anisoxeas ascending towards surface. Extra-axid skeletal columns are separated by large cavernous areas (canals, up to 650 pm diameter), New Caledonian Axinellids 1429 covered by an external collagenous layer (up to 550 pm wide, extending for 750-1250 pm between surface conules. Megascleres (refer to Table 5 for dimensions). Two length-classes of megascleres distinguished here, although these clearly intergrade in size and morphology. Longer megascleres are thin, curved or sinuous strongyles, predominantly in extra-axial region of choanosome and at surface, although also coring axial fibres, with asymmetrical (styloid), or symmetrical rounded ends. Shorter megascleres are slightly curved, thin anisoxeas, found predominantly in the axial skeleton, although also dispersed in peripheral skeleton, usually with symmetrical rounded or pointed, usually telescoped ends ('oxeas'), or less often with asymmetrical ends (hastate points and evenly rounded bases; 'styles'). Microscleres. Absent. Distribution Known only from the coral reefs of the New Caledonian lagoon, 25-40 m depth. Remarks Although two size classes of megascleres (longer sinuous strongyles and shorter curved anisoxeas) are differentiated in the above description, these intergrade to some extent and intermediate 'oxeas' and 'anisoxeas' could fit into either category. There was no observed regional localisation of these two categories of spicules to any particular region of the Table 5. Comparison in spicule dimensions between species of Ptilocaulis All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of measurement. N = 25 for each specimen - Species (material) Ptilocaulis echidnaeus (Lamarck) (holotype MNHN DT640) Ptilocaulis epakros, sp. nov. (holotype QM G300016) Ptilocaulis fusiformis Levi (holotype MNHN DCL818) (QM G300719) (holotype MNHN DCL818) (QM G300719) Ptilocaulis gracilis (Carter) (holotype BMNH 1845.12.30.1) Ptilocaulis papillatus, sp, nov. (holotype QM (3300748) Ptilocaulis rigidus Carter (?holotype BMNH 1936.5.16.1) Ptilocaulis spiculifera (Lamarck) (holotype MNHN DT638) - Megascleres Length Width 493-(647.8)-816 18-(22.4-29 Styles anisoxeas, oxeas 424-(448.2)-448 1.5-(1.8)-2 Vestigial strongyles 134-(268.4)-328 2.5-(3.4-5 Styles, styloids 414-(664.0)-900 2.5-(3.5)-6 516-(649.3)-743 2-(2.3-3.5 Sinuous strongyles 224-(267.5)-350 6-(8.2)-11 196-(269.9)-339 2,543.3-6 Styles, anisoxeas, oxeas with telescoped ends 183-(290.6)-562 5-(6.2)-8 Styles, sinuous styles 140-(190.1)-243 1.5-(2.4-4 Vestigial styles, styloid, rarely strongyloid 252-(364.0)-481 13-(17.6)-21 Styles, oxeas, anisoxeas 242-(291.6)-334 11-(16.9-20 Short robust curved styles 11-(12.2)-14 492-(599.6)-704 Long slender, curved styles 1430 J. N. A. Hooper and C. L h i skeleton, and thus we cannot confirm Wiedenmayer's (1977) 'typical' diagnosis of these characters for the genus. Similarly, unlike most other species of Ptilocaulis, it is difficult to classify the spicules in P. fusiformis as either styles, oxeas or strongyles, due to the great range in spicule terminations (abrupt points, telescoped endings, evenly rounded bases, tapering points). LCvi (1967) suggested that the spicules in the holotype were mostly styles, but re-examination of this material confirmed that a diversity of spicule terminations is characteristic of the species, although 'true styles' also occur in low numbers. In its spiculation, the present species is most similar to Rhaphoxya (e.g. R. typica Hallmann, 1916b: 645, text-fig. 17), and thus the species is virtually intermediate between Ptilocaulis and Rhaphoxya. However, its characteristic growth form, surface sculpturing (papillae), well-differentiated axial and extra-axial skeletal structure (including a compressed axial skeleton) indicate that P. fusiformis is most appropriately placed in Ptilocaulis. The species is contrasted further with other Australasian Ptilocaulis below, and in Table 5. Fig. 22. Ptilocaulis fusiforrnis L M , specimen Q M G300719: A, shorter predominantly axial anisoxeas; B, longer predominantly extra-axial strongyles; C, section through peripheral skeleton. New Caledonian Axinellids Fig. 23. Ptilocaulis fusCformis LCvi: A , holotype (MNHN DCL818); B, specimen ; [QM G300719 (ORSTOM R1547)l; C, SEM axial fibres (magnified 4 0 0 ~ ) D, skeleton of holotype (scale= 500 pm); E, SEM skeleton (QM G300719) (magnified 40x). Ptilocaulis epakros, sp. nov. (Figs 24-25, Table 5) Material Examined Holotype. QM G300016 (ORSTOM R1232): stn 247, Baie Kouo, Canal Woodin, New Caledonia lagoon, 22°23.5'S.,166049.2'E., 40 m depth, 29.xi.1978, coll. P. Laboute, SCUBA. Description Colour. Live coloration pale yellow-brown (Munsell 2.5YR 8/8), beige in ethanol. 1432 J. N. A. Hooper and C. Ltvi Shape. Arborescent, bifurcate branching, 200 mm long, 70 mm maximum lateral branch span, with thin, cylindrical branches, 27-60 mm long, 5-17 mm wide including papillae, tapering towards pointed branch tips, with long, unornamented stalk, 75 mm long, 4 mm diameter, and expanded basal attachment, 13 mm diameter. Surface. Heavily ornamented, papillose surface, with long, close-set, sharply pointed, soft papillae, 2-4 mm long, 0-5-1 mm diameter, up to 2 mm apart; tips of papillae bifurcate and/or hispid; bases of adjacent papillae interconnected by membraneous ridges running longitudinally along branches, slightly elevated above surface of sponge. Oscules not observed; minute ostia, about 200 pm diameter, scattered between papillae. Ectosome. Fleshy, membraneous ectosome, without specialised spicules, with sparse detritus embedded in and on surface, with heavy collagenous, aspicular matrix, 150-200 pm wide, lying between papillae (=surface ridges) and on sides of papillae; apex of each papilla has plumose brushes of choanosomal styles, in small multispicular bundles, protruding for short distances, up to 200 pm from surface. Choanosome. Skeleton structure plumo-reticulate, with clearly differentiated axial and extra-axial regions, and a compressed axial skeleton. Compressed region of skeleton occupies only about half (2-3 mm) of branch diameter (3-5 mm), composed of heavy, bulbous spongin fibres, with individual fibres only about 100 pm long, 50-70 pm diameter, together Fig. 24. Ptilocaulis epakros, sp. nov., holotype QM G300016:A, styles and styloids, comprising the majority of spicules; B, sinuous strongyle, relatively rare; C , section through peripheral skeleton. New Caledonian Axinellids 1433 forming a close-set reticulation and producing small oval meshes, 50-90 pm diameter; axial fibres cored by uni-, pauci- or multispicular tracts of thin megascleres, occupying only a small proportion of fibre diameter. Abundant, lightly pigmented collagen in mesohyl of axial skeleton. Extra-axial skeleton extensive, including area immediately surrounding axis of branches (1-2 mm diameter) as well as elongated, slender skeletal columns (=papillae; up to 4 mm long). Extra-axial skeleton composed of primary and secondary fibre systems, differentiated mainly by presence or absence of coring spicules; both fibre systems composed of heavy spongin fibres, with individual fibres up to 300 pm long, 40-60 pm diameter, producing a relatively wide, elongate mesh reticulation, 110-170 pm mesh diameter; ascending extra-axial fibres cored by multispicular, plumose columns of choanosomal styles, with spicule tracts becoming heavier towards peripheral skeleton [eventually terminating in plumose brushes which may or may not protrude through surface (tips of papillae, versus between papillae respectively)]. Many (but not all) transverse, connecting fibres in extraaxial skeleton uni- or aspicular, with long exceedingly slender strongyles (although these were invariably broken in situ, and consequently were not observed in spicule preparations). Collagen abundant and slightly more darkly pigmented in extra-axial region; choanocyte chambers not observed. Megascleres (refer to Table 5 for dimensions). Two categories of spicules present, clearly differentiated in morphology but not obviously localised to any particular region of skeleton. Few transverse, connecting fibres contain a single, long, thin, sinuous strongyle, but these spicules are rare. Majority of spicules are styles or styloids, short or long, slender, straight or slightly curved asymmetrically, with evenly rounded or tapering mucronate bases, and hastate, fusiform or telescoped points. Fig. 25. Ptilocaulis epakros, sp. nov.: A, SEM skeletal structure (scale= 1 mm); B, holotype [QM G300016 (ORSTOM R1232)]; C, holotype in situ (photo P . Laboute). J. N. A. Hooper and C. LCvi Microscleres. Absent. Distribution Known only from the New Caledonian lagoon, inter-reef region, 40 m depth. Remarks This species is obviously a closely related, sibling species of Ptilocaulis fusiformis. In its external morphology, including shape and size of surface papillae, it is easily differentiated from P. fusiformis, whereas there are only subtle differences in the geometry and size of megascleres (Table 5), and in the structure of the skeleton between the two species (Figs 22, 23 cf. Figs 24, 25). In fact, all three sympatric sibling species, P. fusiformis, P. epakros and P. papillosus, are most visibly differentiated only by these features. It is conceivable that all three nominal species are members of a single, extremely polymorphic species, but gross external morphology and more subtle skeletal differences do not presently support this hypothesis, and the three taxa are retained here. Only molecular evidence will support or refute this hypothesis. In its arborescent growth form, with thinly cylindrical bifurcate branches and long, unornamented stalk, and its long, tapering, sharply pointed surface conules, P. epakros shows remarkable superficial resemblance to Thrinacophora funiformis Ridley & Dendy (1887: pl. 24, fig. 1) (Raspailiidae), although obviously in spiculation and skeletal architecture the two species are otherwise quite different. Etymology Named for the long, attenuated, sharply pointed surface papillae, from epakros (Gk), pointed at the end. Ptilocaulis papillatus, sp. nov. (Figs 26-27, Table 5) Material Examined Holotype. QM G300748 (ORSTOM R564): stn 113, L'epave du 'Bonhomme', Grand RCcif Mbkre, 22°21.0'S.,166014~(YE.,25 m depth, 21.vi.1976, coll. P. Laboute, SCUBA. Description Colour. Live coloration unknown, white in ethanol. Shape. Elongate, cylindrical digitate, 120 mm long, 14 mm wide at widest point, 4 mm wide at end of branch, single branching and bifurcate tips at end of branches, with short (unornamented) stalk, 17 mm long, 5 mm diameter, and enlarged basal attachment, 16 mm diameter. Surface. Prominently sculptured surface composed of large discrete papillae, each papilla formed by extra-axial skeletal columns, up to 6 mm long, 2.5 mm diameter, standing perpendicular to axial core, dispersed approximately 1.5-3 mm apart, enlarged, flattened and bifurcate at apex, usually interconnected by translucent dermal membrane, but cavernous below membrane. Ectosome. Translucent dermal membrane, collagenous, aspiculose except at ends of extra-axial skeletal columns (papillae) where tufts of megascleres protrude slightly through surface. Choanosome. Skeletal structure plumo-reticulate, with very well differentiated axial and extra-axial skeletons, and heavily compressed axial skeleton. Spongin fibres dominate both sections of skeleton; in axial region fibres heavy, close-set, up to 150 pm long, 50 pm diameter, bulbous, meandering, producing oval, elongate or irregular meshes, 30-140 pm diameter, with heavy collagenous mesohyl; in extra-axial skeleton fibres less heavily compacted, with individual fibres up to 250 pm long, 30 pm diameter, producing elongated New Caledonian Axinellids Fig. 26. Ptilocaulis papillatus, sp. nov., holotype QM G300748: A , vestigial styles, styloids and strongylote spicules; B, section through peripheral skeleton. Fig. 27. Ptilocaulis papillatus, sp. nov.: A , holotype (QM G300748), SEM skeletal structure (scale = 1 mm); B, SEM axial fibres (scale = 200 pm); C, SEM fibre structure (scale = 100 pm); D, holotype. J. N. A. Hooper and C. LCvi 1436 meshes up to 170 pm diameter, more-or-less directed towards surface, and fibres become thinner and more widely spaced near ectosome. Spicule skeleton nearly vestigial, with very few megascleres in axial region, but becoming increasingly abundant towards surface; extraaxial spicule skeleton plumose, ending with plumose bundles of megascleres protruding slightly through surface, at apex of skeletal columns (papillae). Area between skeletal columns usually cavernous, without collagen, fibres or mineral skeleton, measuring up to 2 mm between adjacent columns. Collagen in extra-axial skeletal columns abundant but lightly pigmented; choanocyte chambers small, oval, 20-40 pm diameter. Megascleres (refer to Table 5 for dimensions). Most spicules vestigial, with blackened axial canals, exclusively styles or styloids, rarely strongylote, usually slightly curved, sometimes sinuous, with evenly rounded or tapering bases and hastate points, not telescoped. Microscleres. Absent. Distribution Known only from the New Caledonian lagoon, in the inter-reef region on sandhubble substrate. Remarks This species is most similar to the sympatric P. fusiformis in its skeletal structure, growth form and spiculation, except that all these characters are greatly exaggerated in P. papillatus. Surface papillae are exceptionally large and bifurcate, and rejoin with adjacent papillae at their apex by a translucent dermal membrane; spongin fibre compression is relatively greater in both axial and extra-axial regions; spicules are virtually vestigial in most of the skeleton, except towards the surface where they become exclusively plumose; and spicules are exclusively styloid, without any apparent differentiation in size (Table 5). These features clearly differentiate this species from P. fusiformis, whereas differences between P. fusiformis and P. epakros are more subtle. Etymology Named for the unusual surface sculpturing, from papillatus (Lat.), bud-like, papillose. Genus Pseudaxinella Schmidt Pseudaxinella Schmidt, 1875: 120.-Thiele, 1903: 378; Bergquist, 1970: 20; LCvi, 1973: 606; Wiedenmayer, 1977: 155 [type species Pseudaxinella sulcata Schmidt, 1875: 1201. Diagnosis Usually massive, subspherical, cushion-shaped, unbranched or lobate growth forms, with finely conulose or tuberculate, corrugated surface. Skeleton typically plumo-reticulate, without axial compression or differentiation between axial and extra-axial regions; skeletal tracts consist of oxeas and styles, often in crowded, ascending tracts. Ectosome fleshy, and megascleres in this region may be thinner than choanosomal spicules, but not definitely so. Megascleres typically include only (anis-)oxeas and styles in equal proportions, but some species also have long flexuous styles or strongyles confined to the surface. Microscleres absent (modified from Wiedenmayer 1989). Remarks The presence, absence or modification of megascleres to long flexuous diacts (strongyles) probably has questionable diagnostic value in the Axinellidae. Vacelet (1969) and Pansini (1983) have shown that the traditional distinction between Axinella and PhakeNia (cf. Vosmaer 1912), based on external form and the presence of flexuous diactines in the axial skeleton, is not reliable. Consequently, the importance of those spicules in diagnosing Pseudaxinella may also be of doubtful systematic value (Wiedenmayer 1989). Nevertheless, in the absence of a reliable, comprehensive revision of the Axinellidae, incorporating all taxa that have sinuous strongylote modifications, it is not possible to evaluate whether or not New Caledonian Axinellids 1437 those megascleres occur consistently or at what level they have systematic value. Consequently, definitions of Pseudaxinella provided by Ridley and Dendy (1887), Vosmaer (1912, 1935a), Babic (1922) and Topsent (1934) must be treated with circumspection. Pseudaxinella was originally erected for species 'like Axinella' (i.e. in spiculation), but lacking axial compression (de Laubenfels 1950); but Wiedenmayer (1989) noted that some species [e.g. P. convexa (Hoshino, 1981), P. decipiens Wiedenmayer, 19891 have nearly confused skeletons, atypical of the genus, although agreeing in all other respects. Thus, in Wiedenmayer's (1989) opinion, skeletal organisation may be a poor diagnostic character for these groups, whereas features such as the absence of a special axial skeleton and external morphology (massive subspherical in Pseudaxinella, thinly flabellate in Axinosia/Teichaxinella) might be more reliable diagnostic features. However, rightly or wrongly, the present scheme of classification for the Axinellidae differentiates all constituent genera at least partly on the basis of their skeletal construction, and without revising this current basis for the classification, which is unrealistic in the present work, we propose to retain this character [i.e. the non-compressed, plumose (plumo-reticulate) spicule skeleton] for now. The genus is discussed further at length by Wiedenrnayer (1989: 48). Prior to the present work, only three species had been recorded from the Indo-west Pacific: Pseudaxinella australis Bergquist, 1970, from northern New Zealand (holotype NMNZ Por.26 [not seen]), also recorded recently from northern Australia (Hooper, unpublished; Fig. 30, Table 6); Pseudaxinella decipiens Wiedenmayer, 1989, from Bass Strait, Vic. (holotype NMV F51961 [not seen]); and Phakellia tumida Dendy, 1897a, from Port Phillip, Vic. (holotype NMV G2464 [not seen]) (Vosmaer 1912). A fourth species is also known from the neighbouring province, Ariake Sea, Japan: Axinella convexa Hoshino, 1981 (holotype MMBS AR-1-11 [not seen]). Pseudaxinella debitusae, sp. nov. (Figs 28-29, Table 6) Material Examined Holotype. QM G300725 (ORSTOM 'cfR806'): stn 124, I1Bt Maitre, 22°20.1'S.,166025. l1E., 25 m depth, 1.x.1991, coll. G. Bargibant, SCUBA. Paratypes. QM G300722 (ORSTOM 'cfR806'): stn 181, Ilat Maitre, 22°20.0'S.,166025.0'E.,10 m depth, 1.x.1991, coll. J.-L. Menou, SCUBA; NTM 23887, QM G300695: Baie des Citrons, off Noumea, 22°20'S.,166027'E., 3 m depth, 25.ix.1990, coll. J . N. A. Hooper, snorkel, stn JH-90-019. Other New Caledonian material. QM G301263 (ORSTOM R1227), ORSTOM 'cfR1221': stn 261, SW. IlBt Nda, Lagon Sud, 21°52.5'S.,166"51.2'E.,33 m depth, 4.xii.1979, coll. P. Laboute, ORSTOM, SCUBA; QM G301319, (3301328, G301332: Croisant-Laregribre, IlBt Maitre, off Noumea, 22"20.2'S., 166'22.5'E., 20 m depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA. Description Colour. Orange to orange-yellow alive (Munsell 10R 6/10-2.5YR 7/8), beige or light brown in ethanol. Shape. Massive, irregularly or regularly subspherical, cushion shaped, 55-80 mm diameter, 32-40 mm maximum height above substrate, loosely attached to large pieces of detritus (e.g. dead coral, pelecypod valve), or occasionally rolling free on the substrate (i.e. 'tumbleweed' effect). Surface. Evenly microconulose, goose-flesh appearance, covered by small conules, 1-2 mm diameter, less than 0.5 mm high, scattered over entire surface, interconnected by semi-translucent dermal membrane. Oscules scattered over 'upper' surface, large in life (5-10 mm diameter), contracted in ethanol (1-2 mm diameter), located in slight depressions on surface and surrounded by slightly raised membraneous lip. Texture soft, compressible, relatively easy to tear. Ectosome. Membraneous, fleshy surface, with tips of choanosomal spicules protruding for short distances, up to 150 pm, from surface in sparse plumose brushes. Heavy, more darkly pigmented collagen clearly marks peripheral region, whereas in choanosomal mesohyl J. N. A. Hooper and C. L h i 1438 collagen is only lightly pigmented. Oval choanocyte chambers in peripheral skeleton, 70190 pm diameter, also clearly outlined by more darkly pigmented, granular spongin. Choanosome. Skeleton plumo-reticulate, without axial compression or any noticeable difference between axial and extra-axial regions. Spongin fibre skeleton reticulate, with predominantly ascending primary fibres, up to 70 pm diameter, interconnected by shorter, thinner secondary fibres, 40-60 pm diameter, together producing oval meshes 70-140 pm diameter. Spicule skeleton plumo-reticulate, with clearly differentiated primary, ascending, multispicular tracts, cored by spicules in plumose bundles, interconnected by secondary, unior paucispicular, more-or-less transverse spicule tracts, and spicule reticulation producing a vaguely subrenieroid reticulate skeleton, although the plumose component is emphasised over the reticulate component. Mesohyl moderately light, lightly pigmented. Megascleres (refer to Table 6 for dimensions). Spicules predominantly oxeas, with rare styles and strongyloxeas also present; all spicules relatively long, slender, usually asymmetrically curved (but not rhabdose), sometimes straight, mostly sharply pointed, fusiform, although telescoped and bifurcate points also observed. Microscleres. Absent. Table 6. Comparison in spicule dimensions between similar species of Pseudaxinella All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of measurement. N = 2 5 for each specimen - Species (Locality) (material) Pseudaxinella australis Bergquist (New Zealand) (holotype; Bergquist 1970: 20) (Great Barrier Reef) (QM G300295, G301089, G300880; Hooper, unpublished) Pseudaxinella debitusae, sp. nov. (New Caledonia) (holotype) (paratypes) Pseudaxinella convexa (Hoshino, 1981: 207) Megascleres Length Width 203-(402)-560 9-(15.0)-22 Styles, slightly rhabdose bases 320-(367)-406 3-(4.0)-6 Thinner ectosomal styles 217-(260)-339 8-(9.6)-10 Centrally curved oxeas 172-(274.0)-312 10-(13.2)-16 Styles, slightly rhabdose bases 252-(289.2)-315 8-(11.8)-15 Centrally curved oxeas 243-(358.5)-503 4-(9.6)-15 223-(369.9)-483 2-(7.8)-12 Predominantly asymmetrical oxeas, rare styles or strongyloxeas 550-(760)-920 10-(18)-26 Oxeas, occasionally styles 330-950 9-20 Sinuous oxeas, strongyloxeas Pseudaxinella decipiens Wiedenmayer (1989: 48) (holotype) Pseudaxinella rosacea (Verill) (de Laubenfels, 1950) Pseudaxinella tumida (Dendy, 1897a: 237) Oxeas, styles, strongyloxeas, anisoxeas 542-770 5-9 Sinuous strongyles 235-400 8-1 1 Styles 300-320 8 Oxeas About 180 About 6 Styles only New Caledonian Axinellids 1439 Distribution Known only from the New Caledonian lagoon, subtidal fringing coral reefs, coral rubble substrate, 3-33 m depth. Remarks This species is closely related to Pseudaxinella australis Bergquist, to which it was initially assigned. However, a detailed comparison between the New Caledonian material described above, the New Zealand holotype of P. australis (Bergquist 1970: 20), and another three specimens of P. australis from northern Australia (QM G300880, Moreton Bay, Qld; G300295, Snake Reef, northern Great Barrier Reef, Qld; G301089, Cartier I., Sahul Shelf, W.A.; Fig. 30, Table 6), showed quite a number of differences. The New Caledonian material was pale orange in life, and this colour is probably truly representative of the population as it was observed in all three specimens from the lagoon. In contrast, both New Zealand and Great Barrier Reef material was consistently bright red in life, and all known material was observed to exude clear mucus, whereas this trait was not observed in New Caledonian specimens. Pseudaxinella debitusae has a plumo-reticulate skeleton, with adjacent plumose, ascending skeletal columns interconnected by a paucispicular subreneiroid skeleton, whereas in P. australis the ascending, plumose spicule tracts are more or less discrete with few interconnections (cf. Figs 28B, 29D-E and 30B). In New Caledonian specimens, the plumose spicule brushes protruding through the ectosome were not noticeably thinner than those in the choanosome, unlike the holotype of P. australis (although these 'ectosomal spicules' were not observed in Great Barrier Reef material - Fig. 28. Pseudaxinella debitusae, sp. nov., paratype QM G300722: A , oxeas, styles and strongyloxea structural spicules; B, section through peripheral skeleton. 1440 J. N. A. Hooper and C. LCvi either). Similarly, there was no apparent localisation of different spicule morphologies in P. debitusae (either oxeas, or the rarer styles or strongyloxeas) to any particular region of the skeleton, whereas the plumose spicule columns of P. australis are usually made up of a central core of oxeas surrounded by plumose brushes of styles (the styles verging on echinating). Bergquist (1970) also noted that, in the holotype, oxeas become more abundant towards the peripheral skeleton, but this was not confirmed in northern Australian material of P. australis. Most megascleres observed in P. debitusae were strictly oxeote, with far fewer 'abnormal terminations' (styloid, strongyloid) than seen in P. australis, in which Fig. 29. Pseudaxinella debitusae, sp. nov.: A , paratypes (NTM 23887, QM G300695); B, paratype (QM (3300722); C, specimen in situ (ORSTOM R1227) (photo P. Laboute); D, peripheral skeleton (QM G300722) (scale=500 pm); E, SEM skeleton (magnified 50 x ) . New Caledonian Axinellids Fig. 30. Pseudaxinella australis Bergquist: A , specimens from northern Australia on deck (QM G300880); B, skeletal structure o f specimen Q M G300295 (scale = 500 oxeas and styles occur in about equal proportions. Bergquist (1970) also notes that the texture of the holotype was firm, compressible and brittle, which was similar for the northern Australian material of P. australis, whereas P. debitusae is distinctly soft and compressible, easily torn, probably reflecting the relatively higher spongin fibre content of New Caledonian sponge. Bergquist (1970) contrasts P. australis further with a similar species from Bermuda, P. rosacea, and other Pseudaxinella described from the Indo-west Pacific are also compared in Table 6 . It is our opinion that these subtle and more gross differences between the New Caledonian species and other species of Pseudaxinella justify the creation of a new taxon for this population, although we recognise that the allopatric populations of northern New Zealand, south-east Queensland, Great Barrier Reef, and the Sahul Shelf are closely allied species of this New Caledonian form. Etymology Named for Dr CCcile Debitus, ORSTOM Noumea, in appreciation for facilitating our access to the vast ORSTOM collections, and for her role in the organisation of the collaborative workshops, leading to the publication of the present series of papers on the New Caledonian sponge fauna. Genus Rhaphoxya Hallmann Rhaphoxya Hallmann, 1916b: 641.-de Laubenfels, 1936: 136; Bergquist, 1970: 18 [type species Rhaphoxya typica Hallmann, 1916b: 643 (holotype AM Z1595)l. [Acnnthellina] Carter, 1885: 365 [nomen oblitum; ICZN 50 year rule].-de Laubenfels, 1936: 139; Bergquist, 1970: 18 [type species Acanthellina rugolineata Carter, 1885: 139, holotype BMNH 1886.12.15.941. Diagnosis Massive growth form; surface with papilliform conules. Skeleton not axially condensed, without axial and extra-axial differentiation, consisting of loose, irregularly reticulate, often meandering tracts of spongin fibres and spicules; spicule tracts may (s.s) or may not protrude through ectosome. Ectosome fleshy, lacking any specialised spiculation. Spicules often sinuous or curved, slender, monactinal and/or diactinal (styles, oxeas and strongyles, 1442 J. N. A. Hooper and C. LCvi of one size category, differing only in the character of their extremities). Microscleres raphides, occurring singly or in bundles (trichodragmata) (modified from Hallmann 1916b). Remarks Under van Soest et al.'s (1990) scheme for the distribution of axinellid genera into four families, based on skeletal architecture (i.e. Axinellidae with axially compressed and extraaxially plumoreticulate skeletons; Desmoxyidae with reticulate skeletons; Dictyonellidae with dendritic skeletons; and Halichondriidae with 'halichondroid' disorganised skeletons), Rhaphoxya would fall under Dictyonellidae, together with other genera with atypical, noncompressed, meandering skeletal structure, such as Acanthella. Although we do not currently subscribe to this division of Axinellidae and sister-groups, it does illustrate the problem in classifying axinellids, especially in relying mostly (or exclusively) on skeletal structure. Rhaphoxya does not fit easily with other axinellid genera that have typical skeletal structure (compressed reticulate axial skeleton and plumoreticulate extra-axial skeleton), but it also shows many similarities to Ptilocaulis, as noted above, which does have a typical Fig. 31. Rhaphoxya spp.: A, Acanthella cactiforrnis Carter, skeleton of lectotype ; lectotype; C, Rhaphoxyafelina Wiedenmayer, (BMNH 1886.12.15.81) (scale =500 ~ m )B, peripheral skeleton of specimen (NCI Q66C-3339-U) (photo NCI); D, specimen; E, Rhaphoxya pallida Dendy, choanosomal skeleton of specimen (QM G300469) (scale = 500 pm); F, specimen from northern Queensland. New Caledonian Axinellids 1443 Fig. 32. Rhaphoxya spp.: A , Acanthella rugolineata Carter (holotype BMNH 1886.12.15.24); B, peripheral skeleton (scale =SO0 pm); C, Axinella solida Carter (holotype BMNH 1887.7.11.24); D, peripheral skeleton (scale = 500 pm). axinellid skeleton. These similarities are especially seen by comparing spiculation of Ptilocaulis fusiformis, P. papillatus and P. epakros with Rhaphoxya systremma, sp. nov. (Figs 22, 24, 26 and 33). Bergquist (1970) suggested that Rhaphoxya was also close to Desmoxya (Desmoxyidae) in spiculation and architecture, but differed in having a more lax, less halichondroid skeleton, and in lacking centrangulate spined microscleres. In fact, Bergquist (1970) previously included Rhaphoxya with the Desmoxyidae, whereas Wiedenmayer (1989) referred it to the Axinellidae, illustrating its closer relationships with other axinellids such as Axinyssa. Prior to the present study, six species were assigned to the genus, all recorded from Australasian waters: Acanthella cactiformis Carter, 1885, from Port Phillip Heads, Vic. (lectotype BMNH 1886.12.15.81; Fig. 31A-B) (Carter 1885; Dendy 1897a; Burton 1934; Bergquist 1970; Wiedenmayer 1989); Rhaphoxya felina Wiedenmayer, 1989, from Bass Strait, Vic. (holotype NMV F51964 [not seen]; specimen NCI Q66C-3339-U; Fig. 31C-D); Rhaphisia pallida Dendy, 1897a: 257, from Bass Strait, Vic., and far north Queensland (holotype NMV G2377 [not seen]; specimen QM G300469; Fig. 31E-F) (Hallmann (1916b: 646); Acanthella rugolineata Carter, 1885: 365, from Port Phillip Heads, Vic. (holotype BMNH 1886.12.15.24; Fig. 32A-B); Axinella solida Carter, 1885, from south-east Australia (lectotype BMNH 1887.7.11.24; Fig. 32C-D) (Dendy 1897a); and Rhaphoxya typica Hallmann, 1916b: 643, also from Port Phillip Bay, Vic., and Tasmania (holotype AM 21595 [not seen]) (Guiler 1950: 9). Rhaphoxya systremma, sp. nov. (Figs 33-34, Table 7) Material Examined Holotype. QM G300013 (ORSTOM R1221): stn 261, SW. Ilbt Nda, Lagon Sud, 21°52.5'S., 166'51.2'E., 33 m depth, 4.xii.1979, coll. P. Laboute, ORSTOM, SCUBA. J. N. A. Hooper and C. L&i 1444 Paratype. QM G300442 (fragment NTM 21500): Euston Reef, NE. of Cairns, Great Barrier Reef, Qld, 16°43.0'S.,146013.8'E., 42.5 m depth, 26.i.1981, coll. QFS, dredge (stn B6, Cairns ground truth survey). Description Colour. Pale or dark orange-brown alive (Munsell 7.5 YR 7/10-5/6), beige (holotype) t o dark brown (paratype) in ethanol. Shape. Spherical o r subspherical, globular growth form, 32-75 m m high, 28-60 m m diameter, consisting of aggregated, globular lamellae ('lacunae' of earlier authors), producing a conglomerated, honeycombed, Echinoclathria-like reticulation with numerous, oval, celllike cavities and large canals excavating entire sponge. Sponges are only loosely attached t o pieces of coral rubble or shell fragments, or occasionally rolling free o n substrate ('tumbleweed' sponges). Surface. Membraneous, gelatinous, irregularly convoluted surface, with prominent, rounded papillae, u p t o 3 m m high, 2 mm diameter, most abundant o n apical surface of sponge; largest papillae near apex ('upper surface') surround 1 or more oscules, 2-4 m m diameter, although oscules also occur in other places on the 'upper surface', such as in ridges between surface papillae. Texture soft, compressible, difficult t o tear. Ectosome. Fleshy, heavily collagenous, darkly pigmented ectosomal region, without spongin fibres or spicules, but with a thick collagen layer (up t o 80-300 pm thick) between surface and beginning of choanosomal spongin fibre skeleton; this collagenous layer is thicker in between the surface ridges and papillae than o n top of these structures; sparse Table 7. Comparison in spicule dimensions between Rhaphoxya species All measurements given in micrometres, and expressed as range of measurement Species (locality) [source of information] Rhaphoxya cactiforrnis (Carter) (Port Phillip Bay) [lectotype BMNH 1886.12.15.811 Rhaphoxya felina Wiedenmayer (Bass Strait) [specimen NCI Q66C-3339-U] Rhaphoxya pallida (Dendy) (Port Phillip Head and Bass Strait) [specimen QM G3004691 Rhaphoxya rugolineata (Carter) (Port Phillip Bay) [holotype BMNH 1886.12.15.241 Rhaphoxya solida (Carter) (Port Phillip Head and Bass Strait) [lectotype BMNH 1887.7.11.241 Megascleres Length Width Raphides 167-432 4-12 Absent Predominantly oxeas with telescoped ends, some styles 192-415 5-12 110-245 Exclusively oxeas 211-430 4-9 110-370 Predominantly oxeas with telescoped ends, some anisoxeas and strongyloxeas 305-5 14 7-12 Absent Predominantly oxeas with telescoped ends, some styles 232-388 4-8 65-146 Predominantly styles with hastate or teiescoped points, some oxeas and anisoxeas Rhaphoxya systrernrna, sp. nov. (New Caledonia and northern Great Barrier Reef) [holotype QM G3000131 [paratype QM G3004421 Rhaphoxya typica Hallmann (Port Phillip Bay and Bass Strait) [Hallmann 1916b: 6461 Absent 248-(304.6)-369 2-(2.8)-4 Absent 201-(299.8)-382 2-(3 3-5 Predominantly vestigial, sinuous strongyles, rarer oxeas with telescoped ends 100-700 2-9 55-400 Predominantly oxeas with telescoped ends, fewer styles or strongyles New Caledonian Axinellids 1445 plumose brushes of choanosomal spicules may also protrude through surface, up to 100 pm, especially on tips of papillae, otherwise entire ectosomal skeleton is collagenous. Sparse deposits of detritus also dispersed over surface and incorporated into ectosomal collagenous layer, but this is not a prominent feature of the skeleton. Choanosome. Skeleton plumose, slightly plumo-reticulate, without axial compression or differentiation between axial and extra-axial skeletons; skeleton is dominated mainly by diverging, meandering, sinuous spongin fibres and spicule tracts; spongin fibre system composed of ascending, primary fibres, with individual fibres up to 350 pm long, 5070 pm diameter, cored by multispicular tracts of choanosomal megascleres, and interconnected by shorter, transverse, secondary fibres, up to 120 pm long, 20-40 pm wide, usually aspicular or sometimes paucispicular; generally the reticulate, connecting, secondary spicule tracts are greatly reduced in proportion to the primary, plumose, ascending skeleton. Fibre reticulation produces elongate, often cavernous meshes, 70-230 pm long, up to 80 pm wide, becoming more cavernous in periphery than in axis, with some scattered, sinuous spicules outside fibres; choanosomal mesohyl with abundant but only lightly pigmented collagen, with only few, small choanocyte chambers seen, up to 30 pm diameter. Megascleres (refer to Table 7 for dimensions). Single category of choanosomal spicule present, relatively homogeneous in size, although terminations of spicules vary from symmetrical, evenly rounded and hastate tapering rounded ends (strongyles), to sharply pointed, telescoped ends (oxeas). Majority of spicules strongylote, sinuous, very slender. Microscleres. Absent. Distribution New Caledonian lagoon and northern Great Barrier Reef, coral reef rubble and inter-reef region, 30-43 m depth. Fig. 33. Rhaphoxya systremma, sp. nov., holotype QM G300013: A, strongyles and oxea structural spicules; B, section through pheripheral skeleton. 1446 J. N. A. Hooper and C. Levi Remarks This is the first record of the genus outside southern Australian waters. Rhaphoxya systremma differs from most other species of the genus in lacking raphide microscleres (four of the six described species have raphides), having exceedingly thin megascleres with predominantly strongylote endings (most other species have predominantly or exclusively styles or oxeas with telescoped ends; Table 7), and having i?i sinuous, virtually non-reticulate skeleton (similar to R. typica). According to their published descriptions, all six previously described species of Rhaphoxya are morphologically very similar and difficult to clearly distinguish on the basis of their published gross characteristics, including growth form, surface features, skeletal architecture and spiculation. However, examination of type material and other representatives of these species shows a number of distinctive, cryptic characters which support their differentiation (Figs 31-34). Rhaphoxya felina has a distinctive, perpendicular ectosomal skeleton, a more confused plumo-reticulate choanosomal skeleton, and megascleres are exclusively oxeas; R. cactiformis has a more confused plumo-reticulate skeleton, heavy collagenous mesohyl, spicules are predominantly oxeas with telescoped ends, and raphides are absent; R. rugolineata has a virtually plumose, diverging spicule skeleton in which the outer layer of spicules on spicule tracts are inserted at acute angles (i.e. nearly echinating), spicules are predominantly oxeas with telescoped ends, and raphides are absent; R. solida has a skeleton similar to that of R. felina, with a perpendicular ectosomal skeleton and compact, confused, plumo-reticulate skeletal tracts, and spicules are predominantly Fig. 34. Rhaphoxya systremma, sp. nov.: A , cross-section through surface papilla (holotype QM G300013) (scale=500 pm); B, holotype; C, specimen in situ (ORSTOM R1221) (photo P. Laboute). New Caledonian Axinellids 1447 asymmetrically curved styles with hastate or telescoped points, with less frequent anisoxeas and oxeas; R. pallida has a meandering plumo-reticulate skeleton similar to that of the present species, with plumose (plumo-reticulate) spicule tracts and a heavy collagenous mesohyl, but spicule tracts are not sinuous and spicules are predominantly oxeas with telescoped ends; R. typica also has a meandering, plumo-reticulate skeleton, similar to that of R. systremma, and its growth form is similar in being massive, subspherical, with papillae on the upper surface, but spicules are substantially larger and consist predominantly of oxeas with telescoped ends, with only few strongyles or styloid modifications (which are predominant in the present species), and raphide microscleres are also abundant. A comparison of spicule dimensions between these species is given in Table 7. Altogether, the present species appears to be most closely related to, and clearly a sibling species of, the southern Australian R. typica. Etymology The species name refers to the subspherical conglomeration of lacunae, forming a honeycombed mass, from the Greek systremma, relating to anything aggregated, consolidated, or twisted together into a round object. Family DESMOXYIDAE Hallmann Definition Axinellid sponges with 'microscleres' in the form of smooth or spined microxeas, often centrangulate or strongly bent centrally, sometimes also with raphides, occurring singly or in bundles (trichodragmata), or acanthose cladotoxa and birotules in one genus. Megascleres monactinal, diactinal or both. Skeleton consisting of widely spaced reticulate bundles of multispicular fibres, with little spongin, with poorly developed or no axial compression, and relatively poorly differentiated extra-axial skeleton (disorganised-plumose). Growth forms encrusting, massive or ramose [compiled from Hartman (1982) and Wiedenmayer (1989)l. Remarks The presence of smooth or spined oxeas is characteristic of, and apomorphic for, desmoxyid genera. These spicules are traditionally classed as microscleres (e.g. Hallmann 1916b, Wiedenmayer 1977), although in comparison to many other taxa they are generally much too large to be considered as such (e.g. Myrmekioderma). Wiedenmayer (1977) suggested that the Desmoxyidae had affinities with the Hemiasterellidae (e.g. Laonaenia Hallmann), and the Hadromerida (e.g. Paratimea Hallmann). However, Topsent (1928) had previously included these genera in the Hemiasterellidae, but he was not followed by subsequent workers. Although some similarities can be drawn between some taxa of both families, the hemiasterellids are restricted to forms with asterose microscleres, whereas the desmoxyids included forms with microxeote spicules. On that basis, several genera currently assigned to Bubaridae (e.g. Rhabdoploca, Bubaropsis) also have inferred affinites with the Desmoxyidae, having acanthose or smooth oxeote megascleres. This family was established by Hallmann (1916b: 673; with synonym Higginsiinae de Laubenfels, 1936: 132), initially for five genera, of which only four were correctly assigned (Desmoxya, Higginsia, Holoxea and Halicnemia). Numerous other genera were subsequently added or associated with the family, although affinities between them were not always completely clear. Of these, the following genera are excluded from the family: Negombo Dendy (type species N. tenuistellata Dendy) and Diacarnus Burton (type species Axos spinipoculum Carter) are probably best assigned to the Latrunculiidae [both suggested as possible synonyms according to Hooper (1986), but this has not been firmly established]; Allantella Hallmann (type species Trachytedania arborea Keller) is probably a hadromerid; Laonaenia Hallmann (type species Hymeraphia verticillata Bowerbank), and Paratimea Hallmann (type species Bubaris constellata Topsent) are aster-bearing taxa, both belonging to Halicnemia Bowerbank according to Topsent (1928), but are probably hemiasterellids or hadromerids (e.g. Hooper 1986). Ommatosa (sensu de Laubenfels 1936; type and only J. N. A. Hooper and C. LCvi 1448 species Axinella rugosa Schmidt) is either a desmoxyid or a bubarid; its placement is problematic, but it probably has closest affinities with Bubaris. Eight genera are considered to be valid and presently included in Desmoxyidae (Acanthoclada Bergquist, Halicnemia Bowerbank, Heteroxya Topsent, Higginsia Higgin, Holoxea Topsent, Microxistyla Topsent, Myrmekioderma Ehlers and Parahigginsia Dendy) but, as van Soest et al. (1990) suggest, there is a pressing need for a complete revision of the family and a re-evaluation of its constituent genera. Only one species of Desmoxyidae has been recorded previously for the New Caledonian region, Parahigginsia phakellioides Dendy, 1924, also known from northern New Zealand. Genus Myrmekioderma Ehlers Myrmekioderrna Ehlers, 1870: 28. - Bergquist, 1965: 177 [type species Alcyonium granulaturn Esper, 1830: 711. Acanthoxifera Dendy, 1905: 156. -Dendy, 1922: 129; Bergquist, 1965: 177 [type species A. ceylonensis Dendy, 1905: 1571. Anacanthaea Row, 1911: 329.-van Soest et al., 1990: 31 [type species A . nivea Row, 1911: 3291. Callistes Schmidt, 1868: 16.-van Soest et al., 1990: 31 [type species C. lacazii Schmidt, 1868: 161. Neoprosypa de Laubenfels, 1954: 189 [type species N. atina de Laubenfels, 1954: 1901. Diagnosis Growth form massive or encrusting; surface hispid, with characteristic canals and grooves forming polygonal tuberculate plates. Choanosome condensed, with confused mass of acanthoxeas and oxeas, strongyles or less frequently styles, forming irregular, ascending, multispicular tracts bound together with sparse collagen. Extra-axial skeleton dense paratangential layer of acanthoxeas, with larger choanosomal styles protruding. Ectosomal skeleton without specialised spiculation, subectosomal acanthoxeas protrude, forming closely adjacent brushes perpendicular to surface. Megascleres long, smooth oxeas, strongyles, or more rarely styles, often sinuous, and centrally flexed or straight acanthoxeas. Microscleres raphides, occurring singly or in bundles (trichodragmata) (modified from Bergquist 1965). Remarks Bergquist (1965) meticulously redescribed the type species from type and recent material, and showed conclusively that the type species of both Acanthoxifera and Neoprosypa were junior synonyms of M. granulata, despite their apparent differences according to their published descriptions. Bergquist (1965) also noted that M, granulata had a wide geographical distribution with a corresponding high degree of skeletal variability, particularly in the presence, absence and size of certain spicule categories. The genus is similar to Anacanthaea and Heteroxya, and also apparently related to Higginsia. Van Soest et al. (1990) revised the higher systematics of Myrmekioderma, placing it in a redefined order Halichondrida and family Halichondriidae, and suggested that it was most closely associated with a genus-group also containing Didiscus (based on a synapomorphy of one or more categories of trichodragmata, the larger sinuously curved). They suggested that within this group, the smaller oxea 'microscleres' varied from an acanthose to a completely smooth condition, and thus had a dubious systematic value. In contrast, Higginsia was retained in Desmoxyidae by van Soest et al. (1990), implying a more distant relationship with M~rmekiodermathan previously recognised, but this opinion was not supported by a chemotaxonomic study (Hooper et al. 1992). Consequently, Myrmekioderma and Higginsia are retained here in the same family. Van Soest et al. (1990) included six species in the genus, of which only two live in the Indo-west Pacific: M. dendyi (Burton, 1959) from the south Arabian coast and Indonesia, M. granulata widespread throughout the Indo-west Pacific (Figs 35-36), M. rea de Laubenfels, 1934, from the vicinity of Puerto Rico, M. spelea (Pulitzer-Finali, 1983) from the Mediterranean, M. styx de Laubenfels, 1953, from the Gulf of Mexico, and M. tulearensis (Vacelet et al., 1976) from south-west Madagascar. New Caledonian Axinellids Myrmekioderma granulata (Esper) (Figs 35-36) Alcyonium granulatum Esper, 1830: 71, pl. 24. Myrmekioderma granu1ata.-Ehlers, 1870: 28; Burton, 1938: 39, pl. 7, fig. 42; de Laubenfels, 1954: 121, fig. 75; Lbvi, 1961: 14, fig. 17; Bergquist, 1965: 177, fig. 27a, b; van Soest et al., 1990: 29, fig. 28; Hooper et al., 1992: 265. Acanthoxifer ceylonensis Dendy, 1905: 157, pl. 9, fig. 5. -Dendy, 1922: 129. Myrmekioderma tylota de Laubenfels, 1954: 119, fig. 74. Neoprosypa atina de Laubenfels, 1954: 190, fig. 127. Acanthoxifer fourmanoiri LCvi, 1956: 5. Material Examined New Caledonian region material. QM G300022 (ORSTOM R1347): stn 427, Caye de I'Observatoire, Chesterfield Is, 21°24.6'S.,158050.3'E., 28 m depth, 26.vii. 1984, coll. G. Bargibant, SCUBA. Comparative material. Darwin region, N.T.: NTM 2196: Dudley Pt, East Pt, 12°25'S.,130049.01'E., 0-0.5 m depth, 13.ix.1981, coll. J. N. A. Hooper, by hand; NTM 22053: same locality, 12'25.01S., 130°48.4'E., 6-10 m depth, 10.v.1984, SCUBA, coll. J. N. A. Hooper; NTM 2214: Lee Pt, 12O19.02'S., 130°53.01'E., 0 m depth, 14.xi.1981, coll. J. N. A. Hooper, by hand; NTM 2430: same locality, 13.xii.1981. Arafura Sea, N.T.: NCI Q66C-0540-A (fragment NTM 23097): Parry Shoals, 1l011.411S., 129"43.011E., 18 m depth, 14.viii.1987, coll. A.-M. Mussig and NCI, SCUBA. Cobourg Peninsula, N.T.: NTM 22508: NW. Table Head, Port Essington, 11°13~5'S.,132010~5'E., 3-5 m depth, 14.ix. 1985, SCUBA, coll. J. N. A. Hooper; NTM 23249: SW. Table Head, Port Essington, 11°13.5'S., 132"105'E., 2-5 m depth, ll.ix.1986, coll. J. N. A. Hooper and C. Johnson, C., SCUBA. Wessel Islands, N.T.: NCI Q66C-4769-Y (fragment QM (3300741): Bay N. side of Cumberland Strait, 11°28.0'S.,131029.0'E., 20 m, 14.xi.1990, coll. J. N. A. Hooper and NCI, SCUBA. Northwest Shelf, W.A.: NTM 23423: 150 m W. of Enderbry I., Dampier Archipelago, Dampier, 20°35.8'S.,1 16'28 .OIE., 14 m depth, 31.viii.1988, coll. Low Choy, D. and NCI, SCUBA; NCI Q66C-1447-0 (fragment NTM 23332): S. reef 500 m from Direction I. Natl Pk, 21°33~1'S.,1150071'E., 6 m, 24.viii.1988, coll. Low Choy, D. and NCI, SCUBA. Sahul Shelf, W.A.: QM G301141: Hibernia Reef, entrance to lagoon, NE. side reef, 11°57.8'S.,123022.3'E., 23 m depth, 10.v.1992, coll. J. N. A. Hooper, SCUBA; QM G301104: Cartier I., outer reef slope, N. side of reef, Australia, 12"31.4'S.,l23"33.3'E., 22 m depth, 7.v.1992, coll. J. N. A. Hooper, SCUBA. Description Colour. Light orange-brown to bright orange exterior alive (Munsell 7.5YR 7/10-5YR 6/10), often with silt covered 'dusty' surface, orange-brown exterior and beige interior in ethanol. Shape. Massive, subcylindrical, vaguely elongate, rounded, bulbous growth form, partially burrowing soft sediments or excavating hard sediments, up to 350 mm long, 200 mm wide, 160 mm thick (although on the Sahul Shelf specimens several metres in diameter were observed). Surface. Pineapple-like, convoluted, crustose surface, with large conules or rounded or polygonal plates, 18-35 mm diameter, slightly raised above surface and separated by shallow but disctinct grooves; apex of sponge with irregularly meandering or discrete series of relatively deep, excavated channels (sieve-plates or porocalyces), up to 60 mm deep, containing large oscules (up to 50 mm diameter) especially near apex of sponge, each oscule with a raised membraneous lip. Exterior surface invariably silt covered, interior soft, mango-like. Texture harsh, firm, spiculose. Ectosome. Distinct, thick, detachable, paratangential crust of smaller (ectosomal) oxeas, 400-850 pm wide, with innermost layer nearly horizontal and outermost layer nearly perpendicular to the surface, together forming a continuous palisade of spicules. Ectosomal crust supported by long, pillar-like tracts of large oxeas, usually widely spaced (450780 pm long, 750-900 pm apart), producing an excavated subdermal region containing large cavities, about 650 pm diameter, with sparse collagen, collagenous fibrils, bundles of raphides and sparsely scattered smaller oxeas. 1450 J. N. A. Hooper and C . LCvi Choanosome. Cavernous, reticulate skeletal structure, with differentiated primary and secondary spongin fibres and spicule tracts; primary, ascending, multispicular fibres (70250 pm diameter) more-or-less regularly spaced, 300-650 pm apart, interconnected by secondary, transverse or oblique, pauci-, uni- or aspicular fibres (30-80 pm diameter). Fibres lightly or heavily invested with spongin, depending on their thickness, and cored only by larger (choanosomal) oxeas. Fibre meshes evenly rectangular, triangular or irregularly oval, 90-260 pm diameter, containing abundant collagen, collagenous fibrils, many scattered smaller oxeas, and bundles of raphides. Choanocyte chambers small, oval, 40-70 pm. Megascleres. Two categories of megascleres of similar morphology, clearly distinguished only by size and distribution in skeleton; both entirely smooth, relatively large, straight or slightly curved at centre, rarely asymmetrical, tapering to sharp fusiform points. Smaller (ectosomal) oxeas: length 319-(535.3)-708, width 4-(9.2)-12. Larger (choanosomal) oxeas: length 644-(688.1)-782, width 13-(17.3)-22. Fig. 35. Myrmekioderma granulata (Esper), specimen Q M G300022: A, larger (choanosomal) oxeas; B, smaller (ectosomal) oxeas; C, bundles of raphides (trichodragmata); D, section through cavernous peripheral skeleton. New Caledonian Axinellids 1451 Microscleres. Raphides rarely seen individually, but more commonly occurring as bundles of hair-like raphides (trichodragmata). Dimensions of bundles up to 140 x 15 pm. Distribution Widely distributed throughout the Indo-west Pacific: Madagascar, Aldabra (LCvi 1956, 1961), Seychelles (Dendy 1922; Hooper, unpublished data); Gulf of Manaar (Dendy 1905; Burton 1938); Indonesia (Esper 1830; Ehlers 1870; van Soest et al. 1990); north-west Australia (Hooper, unpublished data); central western Pacific: Ponape, Truk, Ebon Atoll (de Laubenfels 1954), Palau, Ifaluk (Bergquist 1965); Chesterfield Islands and New Caledonia (present study). Common habitats range from heavily sedimented fringing coral platforms and coral pools, in sand, silt, beach rock and dead coral rubble substrates, to pristine coral reef slopes, often in spurs and grooves; sublittoral depths to approximately 20 m. Fig. 36. Myrmekioderma granulata (Esper): A , specimen in situ (ORSTOM R1347) (photo J.-L. Menou); B, SEM fibre junction (specimen QM G300022) (magnified 100~)C ; , SEM skeletal structure (magnified 1 0 0 ~ ) D, ; peripheral skeleton (scale = 500 pm). J. N. A. Hooper and C. L h i 1452 Remarks There is no doubt that M. granulata, and probably all of its sibling species, fits poorly with the present concept of the axinellids, and possibly the hypothesised relationship with the halichondrids, as proposed by van Soest et al. (1990), may be more appropriate. However, this relationship between the desmoxyids, axinellids and halichondrids is still very poorly understood, and indeed none of these taxa have yet been revised or well investigated. The conservative choice of leaving Myrmekioderma in Desmoxyidae is presently the best alternative and also supported by chemotaxonomic data (Hooper et a[. 1992). The New Caledonian specimen (described above) differs from typical morphotypes of the species in lacking any spination on the smaller (ectosomal) oxeas, and in lacking any modification to the oxea morphology; in lacking acanthose oxeas the species is not clearly a desmoxyid. However, in most other comparative material examined (listed above), the smaller oxeas are clearly acanthose (and also sometimes modified to strongyles, anisoxeas or styles). Bergquist (1965) provides a comprehensive comparison in spicule dimensions between all nominal species and regional populations of M. granulata, and a detailed discussion of the species' morphometric variability and relationships. Genus Higginsia Higgin Higginsia Higgin, 1877: 291.-Hallmann, 1916b: 655; Dendy, 1922: 126; Topsent, 1928: 39; de Laubenfels, 1936: 132; Burton, 1959: 255; Wiedenmayer, 1977: 156 [type species Higginsia coralloides Higgin, 1877: 291 (Hallmann 1916b: 656), possible junior synonym of Spongia strigilata Lamarck, 1814: 4501. Dendropsis Ridley & Dendy, 1886: 483.-Ridley and Dendy, 1887: 191; Hallmann, 1916b: 693; de Laubenfels, 1936: 132; LCvi, 1973: 606 [type species Dendropsis bidentifera Ridley and Dendy, 1886: 4831. Desmoxya Hallmann, 1916b: 649.-de Laubenfels, 1936: 132; van Soest et al., 1990: 18 [type species Higginsia lunata Carter, 1885: 3581. Diagnosis Growth forms erect, lamellate, massive, vasiform or lobate; surface conulose, papillose, often silt covered or membraneous. Skeletal structure ranges from halichondroid with a partially compressed, reticulate axis, and an irregularly plumo-reticulate extra-axial region (Higginsia), a compressed axis and a radial, non-plumose extra-axial region (Dendropsis), to a lax plumose or plumo-reticulate axial and extra-axial region, without axial compression or regional differentiation of the skeleton (Desmoxya); spongin fibres usually poorly developed although heavy collagen forms mesohyl, usually with numerous megascleres and microscleres scattered between main skeletal tracts; all skeletal tracts cored by monactinal and/or diactinal megascleres. Ectosome without specialised spiculation, but with extra-axial spicule tracts (1 or 2 categories of megascleres) protruding through surface. Megascleres oxeas, strongyles and/or styles of 1-3 sizes. Microscleres include spined, centrangulate curved or straight microxeas, and sometimes also raphides occurring singly or in bundles (trichodragmata) (modified from Wiedenmayer 1977). Remarks The nominal genera Higginsia, Dendropsis and Desmoxya differ essentially in skeletal construction (reticulate or plumo-reticulate; with a compressed axis and radial extra-axis; and plumose-halichondroid, with meandering, occasionally reticulate skeletal tracts respectively). Most species lack definite axial compression of the skeleton (except Dendropsis bidentifera), having instead a halichondroid, vaguely reticulate axis, and in most species there is often some differentiation of axial and extra-axial skeletons (except Desmoxya lunata), suggesting some sort of affinity with the concept of Axinellida. All three nominal genera share the apomorphy of spined microxeas, but we are not completely convinced that the major differences in their skeletal structures can be ignored, particularly in the case of Dendropsis which has a nearly 'classical axinellid' architecture, in recognising these taxa merely as synonyms of Higginsia (e.g. Hallmann 1916b; van Soest et al. 1990). New Caledonian Axinellids 1453 It is possible that resurrection of Desmoxya is required to accommodate Higginsialike species that lack any evidence of axial compression (as seen in both H. lunata and H. anfractuosa, sp. nov.), having instead a simply halichondroid, meandering Rhaphoxyalike reticulation of choanosomal tracts. These species also differ from Higginsia sensu strict0 only in having one size category of megascleres, having raphides in addition to spined microxeas, as well as the more lax skeletal architecture. However, intermediate forms of architecture between Higginsia and Desmoxya are present in some species of Higginsia (e.g. H. massalis), and this condition is interpreted here as being merely a reduced form of the skeleton that is typical for Higginsia (van Soest et al. 1990). Hallmann (1916b: 655-9) provides a comprehensive review of the genus and many of the species it contained at that time, but many more species have since been included in the Fig. 37. Higginsia spp.: A , Higginsia lunata Carter (holotype BMNH 1886.12.15.138); B, peripheral skeleton from BMNH slide (scale = 500 pm); C, Dendropsis bidentlfera Ridley and Dendy (holotype BMNH 1887.5.2.59); D , skeleton (scale=500 pm); E, Higginsia coralloides var. rnassalis Carter (holotype of variety, BMNH 1886.12.15.122); F, skeleton (scale=500 pm); G, Higginsia coralloides (sensu Carter 1885) ('representative specimen' BMNH 1886.12.15.79); H, skeleton (scale=500 pm). 1454 J. N. A. Hooper and C. Ltvi genus, and all are In need of detailed revision. Few type specimens have yet been located and seen, so it is not possible to undertake a review of these species at the present time. Species thought to be valid are: H. bidentifera (Ridley & Dendy) from the Cape of Good Hope, South Africa (holotype BMNH 1887.5.2.59; Fig. 37C-D); H. coralloides Higgin from the Caribbean (holotype BMNH [not seen]), with varieties H. c. liberiensis Higgin, 1877, and H. c. arcuata Higgin, 1877 [possible synonym of H. strigilata (Lamarck); Wiedenmayer 19771; H. higgini Dendy, 1922, from the western Indian Ocean (Okharnandal, Diego Garcia, Providence, Egmont Reef, and south Arabian coast; Dendy 1922; Burton 1959) (holotype BMNH [not seen]); H. lunata Carter from Port Phillip Heads (Dendy 1897a; Hallmann 1916b) (holotype BMNH 1886.12.15.138, schizotype MNHN LBIM DCL283; Fig. 37A-B); H. massalis Carter, 1885, from Port Phillip Heads, Vic., and Ambon, Indonesia (Dendy 1897a; Topsent 1897; Hallmann 1916b) (holotype BMNH 1886.12.15.122; Fig. 37E-F), with Fig. 38. Higginsia spp.: A , Higginsia mixta Hentschel ('representative specimen' NTM 22236); B, skeleton (scale = 500 pm); C, Higginsia scabra Whitelegge (specimen on deck NCI Q66C-0523-5) (photo NCI); D, skeleton of specimen NTM 22801 (scale = 500 pm); E, Spongia strigilata Lamarck (holotype MNHN DT637); F, skeleton (scale = 500 pm). New Caledonian Axinellids 1455 synonym 'H. coralloides7o f Carter (1885)from Port Phillip Heads ('representativespecimen' BMNH 1886.12.15.79; Fig. 37G-H), and 'H. strigilata' o f Desqueyroux-Faundez (1981); H. mixta Hentschel, 1912, from southern Indonesia (holotype SMF 968 [not seen]) and Palau (Bergquist 1965), also known from north-west and north-east Australia (Hooper, unpublished data; 'representative specimen' NTM 22236; Fig. 38A-B); H. natalensis Carter, 1885, from the Cape o f Good Hope, South Africa (holotype BMNH [not seen]); H. papillosa Thiele, 1905, from Calbuco, Chile (holotype probably ZMB [not seen]); H. petrosioides Dendy, 1922, from the Seychelles and Indonesia (holotype BMNH [not seen]); H. pumila (Keller, 1889) from the Red Sea (holotype ZMB 442 [not seen]); H. robusta Burton, 1959, from the Gulf o f Aden (holotype BMNH 1936.3.4.342 [not seen]);H. scabra Whitelegge, 1907, from Port Jackson, N.S.W. (Hallmann 1916b), and north-west and north-east Australia (Hooper, unpublished data) (holotype AM 2480; 'representative specimen NCI Q66C-0523-J; Fig. 38C-D); H. strigilata (Lamarck) from an uncertain locality (Turgot collection), possibly originating from the West Indies (holotype MNHN DT637; Fig. 38E-F) (Topsent 1932; Wiedenmayer 1977); and H. thielei Topsent, 1904, from the Azores (holotype possibly Monaco). O f these species, only five have been recorded from the Indo-west Pacific: H. lunata, H. massalis, H. mixta, H. petrosioides and H. scabra. Higginsia anfractuosa, sp. nov. (Figs 39-40, Table 8 ) Material Examined Holotype. QM G300723 (ORSTOM 'cfR806'): stn 181, E. reef flat, Il6t Maitre, New Caledonia lagoon, 22°20.1'S.,166025.0'E., 1.5 m depth, 2.vi.1977, coll. G. Bargibant, by hand. Description Colour. Pale orange alive (Munsell 7.5YR 8/8), olive-brown in ethanol. Shape. Erect, globular, cylindrical digit, tapering towards base and apex, 62 mm long, 24 mm diameter at base, 32 mm widest diameter, attached directly to substrate (with embedded detritus in basal end), without stalk or other processes. Surface. Evenly distributed, rounded Cliona-like papillae, up to 2.5 mm diameter, only raised slightly above surface, each with a terminal apical oscule (now closed), 1.0-1.5 mm Table 8. Comparison in spicule dimensions between Higginsia anfractuosa, sp. nov., and H. lunata All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of measurement. N = 2 5 for each specimen Spicules Species H. anfractuosa, sp. nov. (holotype QM G300723) Megascleres Microscles Acanthoxeas Raphides I Raphides I1 L W L W L W L W H. lunata Carter (holotype BMNH1886.12.15.138) 238-(298.2)-318 2.5-(3.3)-4.5 (Vestigial, predominantly strongyles) (Robust, predominantly styles, fewer strongyles) 106-(129.3)-173 2.5-(2.9)-3.5 (Straight, evenly spined) 176-(252.9)-286 1.5-(2.1)-2.5 91-(99.8)-112 0.5-(1 .I)-2.0 30-(38.1)-48 2.0-(2.8)-4.0 (Curved, terminally spined) 448-(540.8)-620 0.5-(1.3)-2.0 60-(132.3)-210 0.5-(1.3)-2 1456 J . N. A. Hooper and C. L h i diameter. Surface with distinct (non-detachable) dermis, more darkly pigmented than choanosomal region, and has overall goose-flesh appearance. Texture rubbery, compressible. Ectosome. Heavy collagenous ectosomal layer, 100-250 pm wide, darkly pigmented, containing scattered megascleres and microscleres, predominantly paratangential to the surface. Around oscules spicules ordered into diverging rays, presumably supporting surface papillae and providing support for oscule contractile mechanism (Fig. 40C). Choanosome. Skeleton plumose-halichondroid, with predominantly ascending skeletal tracts, meandering and rejoining throughout skeleton; fibre meshes irregular, elliptical-oval, more cavernous in interior than peripheral skeleton; no axial compression and no differentiation between axial and extra-axial skeletons; spongin fibres only lightly invested with spongin, 30-90 pm diameter, with poorly differentiated primary and secondary elements; fibres fully cored with megascleres, dispersed in plumose-diverging tracts. Mesohyl contains heavy collagen and abundant loose megascleres and rnicroscleres. Choanocyte chambers 20-85 pm diameter. Megascleres (refer to Table 8 for dimensions). Vestigial megascleres, predominantly strongyles, occasionally styloid or strongyloxeas, straight, slender, with symmetrical, rounded ends, or slightly tapering hastate points. Microscleres (refer to Table 8 for dimensions). Acanthoxeas long, slender usually straight, occasionally asymmetrical, evenly spined, granular spination, with tapering, sharply pointed ends. Two size categories of raphides present, both straight, thin, tapering to sharp fusiform points, the larger category nearly the same length as megascleres, differentiated only by Fig. 39. A-D, Higginsia anfractuosa, sp. nov., holotype QM G300723: A , vestigial strongyles, strongyloxeas and styloids; B, two sizes of raphide microscleres; C, acanthoxeas; D, section through peripheral skeleton. E, Higginsia lunata Carter, holotype B M N H 1886.12.15.138, curved acanthoxeas. New Caledonian Axinellids Fig. 40. Higginsia anfractuosa, sp. nov.: A , holotype (QM G300723); B, peripheral skeleton (scale = 500 am); C, oscule and supporting spicules radiating around oscule (scale = 200 pm); D, SEM fibre characteristics (scale = 200 pm). their terminations and relative thickness. Raphides dispersed individually in skeleton, no trichodragmata observed. Distribution Known only from the type locality, coral rubble. Remarks This species is a closely related, sibling species of Higginsia lunata Carter (Carter 1885; Dendy 1897a) from Port Phillip, Vic., meticulously redescribed by Hallmann (1916b: 650). The two species differ substantially only in the size of megascleres and microscleres (Table 8), and geometry of acanthoxeas (curved, terminally spined, slightly centrangulate in H. lunata; straight, evenly spined, not swollen at centre in the present species; Fig. 40C, E ) . Growth form, papillose surface features and skeletal structure are comparable between the two species, and both are atypical of all other Higginsia in having a lax, plumosehalichondroid, meandering skeleton. Nevertheless, these differences between the New Caledonian and Port Phillip populations are greater than normally attributed to interspecific variability, substantially more than cryptic differences, and the New Caledonian population is considered to represent a separate species. Etymology Named for the winding, meandering skeletal columns; from anfractuosus (L.), very winding, sinuous. Higginsia tanekea, sp. nov. (Figs 41-42, Table 9) Material Examined Holotype. QM G300024 (ORSTOM R1298): stn 305, N. Ile Paaba, New Caledonia lagoon, 19"55.3'S.,161°37.3'E., 27 m depth, 24.vi.1981, coll. G . Bargibant, SCUBA. 1458 J. N. A. Hooper and C. Lkvi Description Colour. Pale orange alive (Munsell 7.5YR 7/10), beige in ethanol. Shape. Massive, irregularly bulbous, subspherical, subcylindrical sponge, 210 mm long, 80 mm wide, 55 mm thick, without stalk or other processes, loosely attached directly to substrate, with embedded detritus on 'ventral surface', or unattached and rolling free on substrate ('tumbleweed' sponge). Surface. Slightly bulbous surface, with low, rounded ridges, distinct skin-like, detachable dermis and irregularly dispersed microconules, up to 2 mm diameter, conical or elongate and irregular in shape, not raised more than 2 mm from surface, interconnected by shallow canals and grooves. Surface smooth, not hispid. Texture soft, compressible, relatively fragile, easily torn. Internal consistency porous, cavernous, Echinoclathria-like. Oscules not observed. Ectosome. Collagenous detachable dermis, 100-350 bm wide, containing darkly pigmented collagen, sparsely dispersed thinner ectosomal oxeas forming paratangential tracts, interdispersed with crust of acanthoxeas mostly erect on surface, mostly confined to peripheral skeleton. Choanosomal megascleres not protruding beyond surface. Subdermal region cavernous, directly below skin-like collagenous layer, with sparse tracts of choanosoma1 megascleres supporting dermal layer. Choanosome. Skeleton halichondroid-reticulate, with vaguely ascending spongin fibres and skeletal tracts forming wide-meshed reticulation. Spongin fibres light, more-or-less Fig. 41. Higginsia tanekea, sp. nov., holotype QM G300024: A , choanosomal oxeas and styloids; B, ectosomal oxeas; C, acanthoxeas; D, section through peripheral skeleton. New Caledonian Axinellids 1459 divided into primary, ascending, multispicular fibres, 60-160 pm diameter, and shorter, interconnecting, predominantly transverse, uni- or paucispicular, secondary fibres, 30-75 pm diameter; fibre reticulation forming cavernous, oval or elongate meshes, 350-1200 pm diameter, wider in peripheral skeleton than in deeper regions of choanosome; all fibres cored by larger oxeas, forming irregular, slightly confused, sometimes meandering tracts, with spicules usually occupying most of fibre diameter; numerous larger choanosomal and Fig. 42. Higginsia tanekea, sp. nov.: A , holotype [QM G3000241, peripheral skeleton (scale=500 pm); B, specimen in situ (ORSTOM 'cfR1298') (photo P. Laboute); C, holotype in situ (ORSTOM R1298, on right) (photo P. Laboute); D, SEM skeletal structure (scale =200 ~ m ) . 1460 J. N. A. Hooper and C. LCvi smaller ectosomal oxeas also dispersed between fibre skeleton, but few acanthoxeas seen in choanosome. Fibre meshes contain abundant collagen, more lightly pigmented than in peripheral skeleton, and small oval or eliptical choanocyte chambers, 25-70 ym diameter in peripheral region. Megascleres (refer to Table 9 for dimensions). Choanosomal megascleres predominantly oxeas, rarely styloid, usually long, slender, symmetrically curved, with fusiform, sharply pointed ends. Ectosomal oxeas with same morphology, slightly shorter and substantially thinner. Microscleres (refer to Table 9 for dimensions). Acanthoxeas variable in length, mostly relatively long, slender, usually with slight, angular, central curvature, occasionally straight or with acute bend, sharply pointed tips, with evenly dispersed, large spines. Distribution Known only from the New Caledonia lagoon, in Halimeda beds in the inter-reef region, 27 m depth. Remarks The relationship of Higginsia tanekea to other members of the genus is uncertain. It is not obviously closely related to any other species in the combination of its morphological characters (skeletal structure, spicule size or spicule geometry), although individually most of these features can be found in at least one other species of the genus. In fact, the only apomorphic character in H. tanekea appears to be the possession of unusually large, perpendicular spines on acanthoxeas; a plesiomorphic feature is the lack of long, extra-axial styles which are present in most other species. Higginsia tanekea shows some similarities, in some of its features, with several other species, but these can be differentiated as follows: H, scabra has long slender extra-axial styles, lacks smaller, slender ectosomal oxeas, and has an axinellid skeleton with distinctive plumo-reticulate, diverging architecture, and differentiated axial and extra-axial regions (depicted by Hallmann 1916b: pl. 41, figs 1-3); H. massalis has a similar skeletal architecture and has similarities in the geometry of some of its spicules, but it has shorter, more robust choanosomal oxeas (often asymmetrical and with telescoped points), long extra-axial styles, and shorter acanthoxeas; H. mixta has much thicker choanosomal oxeas (some styloid, some with telescoped points), also with long, slender extra-axial styles, longer ectosomal oxeas, and has an axinellid skeletal structure; H. robusta has much thicker choanosomal spicules (varying from styles to oxeas), long, slender extra-axial styles are present, acanthoxeas substantially shorter, and it too has an axinellid skeleton; H. pumila is poorly known but, from Keller's (1891) description, it differs in having choanosomal spicules consisting of thick styles, and the extra-axial spicules are long, slender styles. Spicule dimensions of these species are compared in Table 9. Etymology Named for the long, slender, sharp spines on acanthoxea microscleres; tanekes (Gk), long-pointed. Higginsia massalis Carter (Figs 43-44, Table 9) Higginsia coralloides.-Carter, 1885: 357. Not Higginsia coralloides Higgin, 1877: 291, pl. 14, figs 1-5. Higginsia coralloides Higgin var. massalis Carter, 1885: 357.-Dendy, 1896: 243; Hallmann, 1916b: 656, 659-65, pl. 29, fig. 6, pl. 38, figs 6-7, pl. 39, figs 1-2, pl. 40, figs 1-4. Material Examined Holotype. BMNH 1886.12.15.122: Port Phillip Heads, Vic., coll. J. B. Wilson. New Caledonian material. QM G300023 (ORSTOM R1222): stn 124, Ili3t Maitre, 22"20.11S., 166"25.11E.,20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM unregistered: Fosse aux New Caledonian Axinellids 1461 Canards, 22"19.2'S.,l66"26'E., 20-25 m depth, date of collection unknown, coll. P. Laboute, SCUBA; QM G301321: Croisant-Larkgritre, IlBt Maitre, off Noumea, 22°20.2'S.,166022 S'E., 20 m depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA. Comparative material. BMNH 1886.12.15.79: Port Phillip Head, Vic., 22 m depth, coll. J. B. Wilson. Description Colour. Pale orange alive (Munsell 5YR 8/6), grey-brown in ethanol. Shape. Massive, elongate, irregularly subspherical, 73 mm long, 46 mm diameter, without stalk or other processes, attached directly but loosely to the substrate with embedded detritus in 'ventral surface', or rolling freely on the substrate ('tumbleweed' sponge). Table 9. Comparison in spicule dimensions between Higginsia tanekea, sp. nov., and related species All measurements given in micrometres, and expressed as range of measurement Spicule Species (locality; source of data) Choanosomal Extra-axial Ectosomal megascleres megascleres megascleres Higginsia tanekea, sp. nov. (New Caledonia; holotype QM G300024) L 628-(824.3)-993 Absent 392-(512.9)-622 W 4-(10.2)-14 3-(4.3-7 (Slender oxeas, (Oxeas) rarely styloid) H. mixta (Hentschel) (Aru I.; Hentschel 1912) 775-1175 L 624-744 2240 4-5 W 28-3 1 20-3 1 (Robust oxeas) (Long styles) (Oxeas) H. mixta (Hentschel) (Palau; Bergquist 1965) 650-912 L 1025-1150 1900-3125 5-7 W 16-21 14-18 H. mixta (Hentschel) (NW. Australia; unpublished data, NTM collection) L 570-(871.3)-1250 850-(1260.0)-2015 Absent W 25-(38.6)-50 4-(9.8)-16 H. scabra (Port Jackson, N.S.W.; Hallmann 1912) L 550-770 950-1 100 Absent W 8-35 15-25 (Robust oxeas) (Long styles) H. scabra Whitelegge (NW. Australia; unpublished data, NTM collection) L 550-(608.3)-675 222-(327.3)-417 Absent W 18-(33.4)-42 5-(7.3)-11 H. massalis Carter (Port Phillip, Vic.; holotype BMNH 1886.12.15.122) 321-(568.4)-704 492-(1129.8)-1750 L 535-(599.8)-706 2-(6.4)-11 W 10-(13.4)-17 5-(9.0)-14 (Robust oxeas, (Long styles, (Oxeas, occasionless often rarely strongyles ally bidentate) styloid) or oxeas) H. massalis Carter (Port Phillip, Vic.; specimen BMNH 1886.12.15.79) 265-(468.8)-669 L 464-(610.1)-692 412-(867.0)-1221 4-(6.4)-9 7-(10.6)-15 W 8-(12.8)-16 H. massalis Carter (Port Phillip, Vic. ; Hallmann 1916b) Up to 900 200-400 L 560-700 4-5 9 W 14-18 H. massalis Carter (New Caledonia; specimen QM G300023) 512-(716.8)-843 L 841-(898.8)-936 632-(1484'3)-2121 2-(5. 1)-8 6-(8.2)-10 W 12-(14.6)-18 (Robust oxeas, (Long styles, (Oxeas) never styloid) never oxeas) Acanthoxea microscleres 71-(111 .@-I43 1.5-(2.9)-4.5 88-152 3-5 62-200 2-5 70-(108.3)-175 3-(6.1)-10 60-130 Up to 5 72-(110.0)-154 1-(4.1)-6 72-(97'8)-124 2-(3.7)-5 71-(84.9-96 2.5-(3.8)-5 40- 130 4-5 74-(96.4-137 2-(3.1)-4.5 J. N. A. Hooper and C. Livi 1462 Surface. Uneven, irregular, lumpy surface, with a distinct skin-like, detachable dermis, covered with evenly dispersed, irregularly shaped microconules, up to 3 mm diameter, raised no more than 2 mm from surface, forming meandering, crenellated ridges and valleys. Oscules, 3-6 mm diameter, slightly raised above surface, with membraneous lip; smaller ostia, less than 1 mm diameter, situated between ridges. Texture soft, compressible, easily torn; internal consistency compact, only slightly cavernous, spiculose, friable, with low spongin fibre component. Ectosome. Minutely hispid surface produced by sparsely dispersed, long extra-axial styles, protruding through a highly collagenous, darkly pigmented dermal layer, 70-220 pm wide. Ectosomal layer contains sparse, paratangential tracts of smaller, thinner oxeas and a thick, paratangential crust of acanthoxeas; acanthoxeas mainly confined to dermal skeleton. Subdermal region slightly cavernous, with elongate canals, 350-900 pm diameter. Fig. 43. Higginsia rnassalis Carter, specimen Q M G300023: A , choanosomal oxeas; B, extra-axial styles; C, 'ectosomal' oxeas; D, acanthoxeas; E, section through peripheral skeleton. New Caledonian Axinellids 1463 Choanosome. Skeleton plumo-reticulate, verging on disorganised-halichondroid, without well developed axial compression of skeleton, with only poorly differentiated axial and extra-axial regions, with extra-axial spicule tracts slightly more plumose than more reticulate choanosomal spicule tracts, and containing long extra-axial styles in peripheral region, usually perpendicular to surface; skeleton consists of differentiated primary, more-or-less ascending, multispicular tracts, 70-110 pm diameter, interconnected by shorter, uni- or paucispicular, secondary, mostly transverse tracts, 20-70 pm diameter, cored exclusively by long choanosomal oxeas, with some thinner 'ectosomal' oxeas interdispersed; fibre reticulation produces elongate meshes, 110-300 pm diameter, but often partially obscured by choanosomal spicules dispersed between major skeletal tracts; spongin fibre system poorly developed, spicules appear to be cemented together primarily by granular collagen; meshes contain abundant collagen, with numerous choanosomal oxeas, only few acanthoxea microscleres, and very small quantities of detritus (arenaceous); choanocyte chambers small, oval to elongate, 25-45 pm diameter. Fig. 44. Higginsia massalis Carter: A, specimen in situ [ORSTOM R1222 (specimen QM G300023)l (photo P. Laboute); B, peripheral skeleton (scale=500 pm); C, same specimen preserved; D, SEM peripheral skeleton (scale = 500 pm). 1464 J. N. A. Hooper and C . Ltvi Megascleres (refer to Table 9 for dimensions). Choanosomal megascleres exclusively oxeas, robust, relatively long, straight or slightly curved at centre, usually symmetrical, with fusiform, tapering, sharp, rarely tel~scopedpoints. Extra-axial styles variable in length, usually very long, slender, slightly curved, sometimes straight or sinuous, with evenly rounded bases, tapering to sharp or slightly telescoped points. 'Ectosomal' oxeas very long, slender, usually slightly curved, sometimes greatly curved or sinuous, sharply pointed; these spicules not confined exclusively to ectosomal region, but also found in smaller quantities in choanosomal skeletal tracts. Microscleres (refer to Table 9 for dimensions). Acanthoxeas relatively long, slender, with slight angular curvature at centre, tapering to sharp points, evenly covered with small spines; spines possibly larger at centre of spicule than on ends of spicule. Distribution Southern Australia and New Caledonia lagoon, inter-reef region on sand and coral rubble substrate, 10-22 m depth. Remarks There are several differences between the New Caledonian specimen of H. massalis, described above, and the holotype from southern Australia. Apart from minor differences in spicule sizes (Table 9), the points of choanosomal oxeas are often telescoped in the holotype, whereas they are rarely so in the present material, being more commonly sharply pointed; ectosomal oxeas in the holotype often have slightly rounded or bidentate terminations, whereas those in the New Caledonian specimen appear to be exclusively sharply pointed; extra-axial styles in the present material are always styloid, whereas those in the holotype are occasionally oxeote; and skeletal architecture is a little more disorganised in the New Caledonian specimen, whereas it is more regularly plumo-reticulate in the holotype. According to Hallmann (1916b: 659), the skeletons of several non-typical specimens from the Port Phillip region are composed of a series of thin lamellae, separated at the periphery but united further towards the choanosome, producing a shaggy peripheral skeletal structure (or 'skeleton-sponge' of Hallmann 1916b). This latter feature was not obvious in the New Caledonian specimen, nor was it apparent in the holotype. These apparently minor differences may eventually prove to be indicative of two cryptic sibling species, possibly justifying the erection of a new taxon for the New Caledonian population. However, examination of a second specimen from Port Phillip Heads, originally assigned to the West Indies species H. coralloides by Carter (1885), showed it to be virtually intermediate in most of these characters, making it nearly impossible to clearly delineate the populations into separate species (without other, non-morphological characters to support such a proposal). Hallmann's (1916b) measurements of spicules in his material differ slightly from those observed in both the holotype and Carter's (1885) specimen, especially the size of ectosomal spicules, and the corrected measurements for this material are presented in Table 9. Species related to H. massalis have already been discussed above and are also compared in Table 9. Discussion An earlier study of five families of poeciloscleridan sponges from the New Caledonian region (Hooper and LCvi 1993) listed 28 species, 18 of which were apparently endemic to this fauna. Although it was noted that there was a greater proportion of indigenous poecilosclerid sponge species (71 %) living exclusively in deeper-waters of the shelf and slope of New Caledonia, agreeing with the preliminary biogeographic model suggested by LCvi (1979), the level of endemism for the shallow-water poecilosclerid fauna in the lagoon and outer reefs was also found to be significantly higher (62%) than previously predicted by LCvi. The present study described or redescribed 16 axinellid species from the shallow-waters in the lagoon and outer reefs surrounding New Caledonia, bringing the total number of New Caledonian Axinellids 1465 known axinellid species in this region to 25, of which 7 are new species and 7 are new locality records. The non-endemic, shallow-water axinellid fauna appears to be most closely related to the north-east Australian (Solanderian) and Indo-Malay fauna, usually representing the easternmost extent of their known distributions in the Indo-west Pacific. The shallow-water fauna also includes two authenticated 'widely distributed' species: Axinella carteri (Dendy) (distributed throughout coral reefs of the western and eastern Indian Ocean, the Indo-west Pacific and the western Pacific rim), and Astrosclera willeyana Lister [found in the western Indian Ocean (RCunion, Madagascar and Mozambique; Vacelet and Vasseur 1965, 1971; Vacelet 1967; Vacelet et al. 1976), eastern Indian Ocean (Christmas I.; Kirkpatrick 1910; Ashmore, Cartier and Hibernia Reefs, Sahul Shelf; Hooper 1992; southern Indonesia; van Soest 1990), central western Pacific Ocean (Guam; Hartman in Vacelet 1967), south-western Pacific Ocean (Great Barrier Reef; Ayling 1982; Loyalty and Tuvalu Is; Lister 1900; New Caledonia; Vacelet 1981), and central southern Pacific (French Polynesia; Vacelet 1977)], whereas other previously suspected 'widely distributed' species were generally found to be separate, closely related, cryptic sibling species. By comparison, the previous study on the poecilosclerid fauna indicated that its affinities lay mainly with both the northern and southern Australian shallow-water fauna, with surprisingly fewer similarities with the northern New Zealand fauna, plus a relatively higher endemic element (Hooper and LCvi 1993). Similar to the Poecilosclerida, however, many of the new axinellids described here were recognisable as sister-species of known, mainly tropical Australasian axinellids. These observed cryptic differences between many of the New Caledonian and Australasian sister-species are considered to be real, representing specific fixed differences in the genotype, rather than phenotypic variability of single, widely distributed species. This point of view contrasts with earlier studies in the western Pacific region (e.g. Burton 1934), and generally has empirical support (biochemical, genetic and detailed morphometric analyses of both sympatric and allopatric species; exhaustive comparison of relevant voucher material cited in these earlier studies, with less reliance on the published literature; Hooper et al. 1990, 1992; van Soest et al. 1991; van Soest and Hooper 1993). Of the four families and 25 species of axinellids known to live in the vicinity of New Caledonia, 12 have not been recorded elsewhere (48% endemism). Most of this endemism can be accounted for by the deeper water fauna (three of five species), compared with only nine of the 20 species being shallow-water species. The proportion of endemic species for all four axinellid families is significantly lower than equivalent figures obtained for five families of Poecilosclerida (71% deeper-water, 63% shallow-water; Hooper and LCvi 1993), or for the entire described sponge fauna in this region (72% deeper-water, and 40% shallowwater). However, considering only the family Axinellidae so far collected and described for the region, the proportion of endemic species is high for both the shallow- and deeper-water species. Further comments on the biogeographic affinities of the New Caledonian Axinellidae will be discussed in a separate contribution. A summary of the New Caledonian axinellid fauna is presented below [S, predominantly or exclusively shallow-water species (0-100 m depth); D, predominantly or exclusively deeperwater species (100-500 m depth); species marked with an asterisk are thought to be endemic to the region]. Axinellidae (15 species have been described for this region, nine of which are apparently endemic): Cymbastela cantharella (LCvi, 1983) [S]* C. concentrica (Lendenfeld, 1887) [S] Reniochalina plumosa (LCvi & LCvi, 1983) [Dl* R. condylia, sp. nov. [S]* Axinella lifouensis LCvi & LCvi, 1983 [Dl* A . carteri (Dendy, 1889) [S] Phakellia columnata (Burton, 1928) [Dl P. pulcherrima (Ridley & Dendy, 1886) [S] P. stipitata (Carter, 1881) [S] J. N. A. Hooper and C. Ltvi Stylissa flabelliformis (Hentschel, 1912) [S] Ptilocaulis fusiformis LCvi, 1967 [S]* P. epakros, sp. nov. IS]* P. papillatus, sp. nov. [S]* Pseudaxinella debitusae, sp . nov. [S]* Rhaphoxya systremma, sp. nov. IS]* Desmoxyidae (five species known for the region, only two of which are possibly endemic): Myrmekioderma granulata (Esper, 1830) [S] Higginsia anfractuosa, sp. nov. [S]* H. tanekea, sp. nov. [S] Higginsia massalis Carter, 1885 [S] Parahigginsia phakellioides Dendy, 1924 [Dl Trachycladidae (one species known for the region, also recorded from south-eastern and south-western Australia): Trachycladus digitatus Lendenfeld, 1887 [S] Agelasidae [now in the order Agelasida; two species known for the region, one apparently endemic, and two hypercalcified 'sclerosponges' are also included here (following van Soest 1984 and others)]: Agelas mauritiana (Carter, 1883) [S] A . novaecaledoniae Levi & LCvi, 1983 [Dl* Astrosclera willeyana Lister, 1900 [S] Stromatospongia micronesica Hartman and Goreau, 1976 [S] Acknowledgements We are extremely grateful to Dr CCcile Debitus, ORSTOM, Noumea, for assisting with the acquisition of specimens and in situ photographs of live material, which have greatly facilitated the preparation of this publication. We also gratefully acknowledge funding provided by both ORSTOM Noumea and DITAC Canberra, which enabled the authors to participate in a series of workshops at ORSTOM Noumea on the taxonomy of New Caledonian shallow-water sponges. This publication is one of several recent contributions on the shallow-water fauna of the Noumea lagoon, as a prelude to the publication of a forthcoming popular book on the subject, and we acknowledge the assistance and interaction of our colleagues Chris Battershill, Patricia Bergquist, Jane Fromont, Michelle Kelly-Borges, Jean Vacelet and Clive Wilkinson. For collection of specimens and photographs we thank Pierre Laboute, Georges Bargibant, Jean-Louis Menou, and other members of ORSTOM Noumea for the hospitality, helpful assistance and cooperation during this project. References Anon. (1985). 'International Code of Zoological Nomenclature.' 3rd edn. pp. 1-338. (International Trust for Zoological Nomenclature, H. Charlesworth and Co.: Huddersfield.) 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