Poecilosclerida from the New Caledonia lagoon (Porifera: Demospongiae).

John N . A . Hooper

Twenty species of shallow-water poecilosclerid demosponges are described from the lagoon and reefs of New Caledonia, including thirteen new species and six new locality records, belonging to five families and eight genera {Myxillidae (1, Acarnus), Desmacellidae (1, Neofibularia), Crellidae (1, Crella), Microcionidae 113, Clathria (Clathriopsamma), C. (Clathria), C. (Thalysias), Echinochalina (Echinochalina), E. (Protophlitaspongia)] and Raspailiidae [4 species, Raspailia (Raspailia), Ceratopsion, Aulospongus]}. The shallow-water fauna contained a mixture of endemic species (65%) and those with affinities to both northern and southern Australia, in contrast to the deeper-water fauna which was previously found to have high endemism (70%) and ancestral affinities to northern New Zealand. The non-endemic shallow-water fauna represents the easternmost extent of species' distributions in the Indo-west Pacific, and the endemic component was also usually immediately recognisable as (transformed) sister-species from Australian tropical and temperate provinces.

Znvertebr. Taxon., 1993, 7, 1221-302 Poecilosclerida (Porifera :Demospongiae) from the New Caledonia Lagoon John N. A . ~ o o ~ e and r * Claude ~ 6 v i ~ Queensland Museum, P.O. Box 3300, South Brisbane, Qld 4101, Australia. Laboratoire de Biologie des InvertBbrts marins et Malacologie, Museum national d'Histoire naturelle, 57, rue Cuvier, 75005 Paris, France. A Abstract Twenty species of shallow-water poecilosclerid demosponges are described from the lagoon and reefs of New Caledonia, including thirteen new species and six new locality records, belonging to five families and eight genera {Myxillidae (1, Acarnus), Desmacellidae (1, Neofibularia), Crellidae (1, Crella), Microcionidae 113, Clathria (Clathriopsamma), C. (Clathria), C. (Thalysias), Echinochalina (Echinochalina), E. (Protophlitaspongia)] and Raspailiidae [4 species, Raspailia (Raspailia), Ceratopsion, Aulospongus]}. The shallow-water fauna contained a mixture of endemic species (65%) and those with affinities to both northern and southern Australia, in contrast to the deeper-water fauna which was previously found to have high endemism (70%) and ancestral affinities to northern New Zealand. The non-endemic shallow-water fauna represents the easternmost extent of species' distributions in the Indo-west Pacific, and the endemic component was also usually immediately recognisable as (transformed) sister-species from Australian tropical and temperate provinces. Introduction Only about 170 species of Porifera have been described from the New Caledonian region, although many hundreds of other species have already been collected by ORSTOM, and the fauna is thought to consist of about 600 species. Furthermore, most of the published studies have concentrated on the deeper-water fauna (Demospongiae, including the polyphyletic Lithistida; LCvi and LCvi 1983a, 1983b, 1988; Uvi 1991), or have dealt exclusively with Hexactinellida (LCvi and L h i 1982), hypercalcified 'sclerosponges' (Vacelet 1981), or Calcarea (Vacelet 1981). Prior to the present study, 76 species of deeper-water (> 150 m depth) Demospongiae had been described from the New Caledonian region, including the Norfolk Rise, whereas only 68 species were known from shallow-waters (LCvi 1967, 1983; LCvi and LCvi 1978; Vacelet 1981; Desqueyroux-Faundez 1984, 1987; Hooper and Bergquist 1992). The present study concerns only the shallow coastal demosponge fauna of New Caledonia, from lagoon and coral reef habitats surrounding the island. This first paper describes a number of species of the order Poecilosclerida; a second paper will deal with the order Axinellida (Hooper and LCvi 1993); the biogeographical relationships of the New Caledonian shallow-water microcionid and raspailiid fauna, with respect to other provinces within the Indo-west Pacific, will be discussed in a separate contribution. These papers represent the results of a collaborative investigation amongst several 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. 1222 J. N. A. Hooper and C. LCvi Methods Methods used to prepare and examine sponges for light microscopy and electron microscopy are described in Hooper (1991b). Spicule measurements are based on examination of 25 random samples of each spicule category for each specimen, and 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, Natural History Museum, London; CSIRO, Commonwealth Scientific and Industrial Scientific Organisation, Division of Fisheries, Hobart; DITAC, Commonwealth of Australia Department of Industry, Technology and Commerce, Canberra; LFM, Merseyside County Museums, Liverpool; MNHN, Museum National d'Histoire Naturelle, Paris; NCI, US National Cancer Institute shallow-water collection contract (Australian Institute of Marine Science, Townsville; primary colle&ions in National Museum of Natural History, Washington, duplicates and fragments in NTM and QM); NMB, Naturhistorisches Museums zu Basel; NMV, Museum of Victoria, Melbourne; N.S.W., New South Wales; N.T., Northern Territory; NTM, Northern Territory Museum of Arts and Sciences, Darwin; ORSTOM, Institut Francais de Recherche Scientifique pour le Developpement en Cooperation, Centre de Noumea; QFS, Queensland Fisheries Service; Qld, Queensland; QM, Queensland Museum, Brisbane; S.A., South Australia; SMF, Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt; Tas., Tasmania; USNM, United States National Museum, Smithosonian Institution, Washington; Vic., Victoria; W.A., Western Australia; ZMA, Institut voor Taxonomische Zoologic, Zoologisch Museum, Universiteit van Amsterdam; ZMB, Zoologisches Museum fiir Naturkunde an der Humboldt-Universitat zu Berlin, Berlin; ZMH, Zoologisches Institut und Zoologisches Museum der Universitat Hamburg, Hamburg. Systematics Order POECILOSCLERIDA Topsent Family MYXILLIDAE Topsent Definition (emended) Poecilosclerida with a renieroid, regularly reticulate o r (plumo-) reticulate choanosomal skeleton, composed of tracts of smooth or acanthose monactinal megascleres (styles, smooth or spined, rarely oxeote or strongylote), with or without echinating spicules; ectosomal tangential skeleton composed of diactinal megascleres, typically being smooth tylotes with microspined bases. Microscleres commonly arcuate isochelae, but may also include anchorate, unguiferous, birotulate or palmate isochelae, anisochelae, sigmas, bipocilli, forceps, and toxas (modified from van Soest 1984~). Remarks Only two other species of Myxillidae have so far been described for the New Caledonian region, both from deeper waters (410-525 m depth)-Lissodendoryx bifacialis LCvi and LCvi, 1983b and L. stylophora LCvi & LCvi, 1983b. Genus Acarnus Gray Acarnus Gray, 1867: 544 [type species Acarnus innominatus Gray, 1867: 5441 [full synonymy and discussion given by Hooper (1987: 72), Hiemstra and Hooper (1991: 433) and van Soest et al. (1991: 49)]. Diagnosis Myxillidae with ectosomal tylotes, choanosomal megascleres include styles or subtylostyles, completely smooth or with basal spines, echinating cladotylotes of 1 o r 2 sizes, with o r without echinating acanthostyles of 1 o r 2 sizes; microscleres include palmate isochelae and toxas, the latter with up to 3 categories including a thick, evenly curved form with recurved ends ('oxhorns') (from Hiemstra and Hooper 1991). Remarks All species of Acarnus share the apomorphic feature of cladotylotes echinating fibres, and some species also have echinating acanthostyles, a symplesiomorphic character for the Poecilosclerida (also found in the Microcionidae, Raspailiidae, Crellidae and Anchinoidae). The presence of acanthostyles was previously used t o differentiate the nominal genus Acanthacarnus LCvi, 1952, from species lacking acanthostyles (i.e. Acarnus s.s.), but the New Caledonian Poecilosclerids 1223 character is considered to be valid only at the infrageneric level (Hooper 1987; van Soest et al. 1991). There is still some debate about the correct family placement of Acarnus, and this stems mainly from the different emphasis given by various authors to the importance of chelae geometry at the family (or even generic) level. Acarnus is included in the Myxillidae, following the scheme of de Laubenfels (1936) and repeated by Bakus (1966), van Soest (1984a) and Hooper (1987), because it has a specialised tangential layer of ectosomal tylote megascleres bearing spined bases, the primary apomorphy for the family, as well as a more-or-less renieroid reticulate skeleton shared by many (but not all) members of the family. The alternative point of view is that the genus is not a 'typical myxillid' because it has palmate isochelae and toxa microscleres, and might better be included in the Microcionidae (e.g. Burton 1959; LCvi 1973; Vacelet et al. 1976). Certainly, many of the nominal myxillid genera have arcuate isochelae, but this feature cannot be considered to be apomorphic for the family (being also found in many other families of PoeciloscleridaCrellidae, Anchinoidae, Coelosphaeridae, Microcionidae, Hymedesmiidae), nor is it found in all genera of Myxillidae: other myxillids have only anchorate isochelae (e.g. Ectyomyxilla Lundbeck, Plocamissa Burton), birotulate isochelae (e.g. Iotrochota Ridley, Iotrochopsamma de Laubenfels, Sigmarotula Bergquist & Fromont), palmate anisochelae (e.g. Acanthorhabdus Burton), unguiferous isochelae (some Myxilla Schmidt), or bidentate sigmoid isochelae (Plumocolumetta de Laubenfels), and some genera have various combinations of these chelae spicules, such as arcuate and unguiferous isochelae (e.g. Anomomyxilla Burton), anchorate and arcuate isochelae (e.g. Forcepia Carter), anchorate and unguiferous isochelae (e.g. Fibulia Carter, Stelotrochota Bakus), palmate anisochelae and birotulate isochelae (e.g. Iophon Gray), or unguiferous and birotulate isochelae (e.g. Iotaota de Laubenfels). It should also be noted that a similar range of chelae diversity is also found in the Microcionidae, and the relative importance of these microscleres, at different levels of classification, has been discussed by Hooper (1991a). Apart from Acarnus, only one other genus (Allocia Hallmann), presently included in Myxillidae, has palmate isochelae, and both these genera have also been previously included in the Microcionidae [by Hallmann (1920) and Burton (1959) respectively], on the basis of their possession of palmate isochelae and toxa microscleres. However, in the senior author's opinion, this interpretation ignores the importance of the tangential ectosomal skeleton of tylote megascleres as the primary apomorphy of the Myxillidae, and it also over-emphasises the importance of chelae microsclere geometry as a family character. Hooper (1991a) suggested that, from a phylogenetic perspective, the significant difference between the Microcionidae and other poecilosclerid families, including Myxillidae, concerned the geometry, derivation and distribution of megascleres within the choanosomal and ectosomal skeletons, whereas the presence or absence of particular skeletal structures or particular forms of microscleres was of lesser importance at the family level. It was also demonstrated that extraordinary levels of homoplasy occurred within the classification when both these sets of features (i.e. the geometry, origin and distibution of megascleres and chelae microsclere geometry) were used in combination and given equal importance in the classification. It was also suggested that because the conjunctive use of these characters produced high levels of homoplasy, then a parsimonious classification based on rnicrosclere geometry (e.g. de Laubenfels 1936) was probably mutually exclusive from one based on structural megascleres, as proposed in part by van Soest (1984a) and developed further by Hooper (1991a). The implication of accepting this latter scheme is that modifications to the geometry of chelae microscleres, which is interpreted by the senior author as being one of grade and not one of clade, must have occurred independently in several poecilosclerid families. This debate is not yet resolved, and for now Acarnus is retained in the Myxillidae following van Soest (1984) and Hooper (1987). Acarnus caledoniensis, sp. nov. (Figs 1-2, Table 1) Material Examined Holotype. QM G300724 (ORSTOM R806): stn 124, Il6t Maitre, 22°20~1'S.,166025~1'E., 23 m depth, 15.vii.1976, coll. P. Laboute, SCUBA. 1224 J . N. A. Hooper and C. Ltvi Paratype. QM G300749 (ORSTOM R806): same locality. Other New Caledonian material. ORSTOM R499: stn 124, Ilbt Maitre, 22°20.1'S.,166025.l'E., 25 m depth, 1.x.1991, coll. G. Bargibant, SCUBA; ORSTOM R522, R528: stn 147, ENE. Ilbt N'Due, 22°12~0'S.,166016~(YE., 22 m depth, 8.ix.1976, coll. P. Laboute, SCUBA; ORSTOM R617: stn 160, W. reef flat, Ilbt Maitre, 22°20.0'S.,166023l'E., 5 m depth, 13.x.1976, coll. A. Intts, SCUBA. Description Colour. Dark red 'fistules' (Munsell 5R 4/10), red-orange body (5R 6/6) alive, siltcovered in situ, light brown in ethanol. Shape. Small spherical, subspherical or massive sponges, up to 90 mm diameter and 45 mm high, not usually firmly attached to substrate but able to roll around freely over sand and rubble. Surface. Covered with silt and evenly distributed microconules, up to 3 mm high, only seen in live state. Several prominent oscules on upper surface, each 8-15 mm diameter, 8-18 mm high, each surrounded by a prominent membraneous lip ('fistule'); 'fistules' and microconules usually collapse upon preservation; surface simply optically smooth with partially translucent ectosomal membrane when preserved. 'Under-side' of sponge usually retains embedded detritus (but this may not be a permanent feature as the sponge appears to be able to roll over the substrate). Ectosome. Skeleton at surface microscopically hispid, with paratangential layer of tylotes concentrated in peripheral region, many protruding through surface. Isochelae abundant and collagenous spongin moderately heavy in peripheral skeleton; collagen granular and brown pigmented. Choanosome. Main skeleton plumose, slightly reticulate, vaguely halichondroid in arrangement due to preponderance of echinating spicules interconnecting adjacent skeletal columns. Skeletal tracts composed of basally spined subtylostyles, containing few (5-10) spicules abreast, 30-80 pm wide; adjacent tracts run more-or-less parallel to each other, ascending to surface, with numerous plumose styles protruding through tracts at acute angles, but only few cross-connections between them. Two size classes of both echinating acanthostyles and cladotylotes present, all relatively abundant, embedded at acute angles to skeletal tracts, together producing misleading impression that skeleton is reticulate. Spicules not visibly enclosed within spongin fibres, although collagenous spongin abundant around skeletal tract. Numerous microscleres dispersed between spicule tracts, with radial columns of toxas particularly abundant. Choanocyte chambers oval, 40-70 pm diameter; chambers frequently vaguely outlined by plumose tracts of both toxas and isochelae. Megascleres. Choanosomal principal subtylostyles long, slender, slightly curved, tapering to sharp points, with swollen tylote bases, usually microspined; some longer, more slender, sinuous subtylostyles also present usually with smooth tylote bases: length holotype 321(354.3)-423 prn [paratype 266-(343.1)-394 pm), width 4-(7.5)-10 pm (4-(7.4)-11 pm]. Ectosomal tylotes long, slender, straight or very slightly curved asymmetrically, with prominently swollen, invariably microspined, tylote bases: length holotype 304-(330.1)365 pm [paratype 299-(337.8)-383 pn), width 3-(5.0)-6 pm (3-(5.2)-7 pm] . Cladotylotes of 2 sizes: larger forms (type I) relatively long, slender, slightly curved, with recurved spines concentrated in middle of shaft, and shaft thicker at base than at apex; main cladome at apex with 4 sharply pointed long clads, minor cladome at base with 4 small clads: length holotype 122-(165.7)-192 pm [paratype 166-(184.9)-206 pm], width 1 .5(2.3)-3 pm 12.543 ~5)-4 pm], width at base 2.5-(3.9-5 pm [4-(4.4-6 pm]. Smaller cladotylotes (type 11) with similar morphology, more slender, usually straight, with granular spines over shaft: length holotype 68-(I 11.2)-152 pm [paratype 82-(111. @-I40 pm], width 1.5-(1.3-2 pm [I-5-(2.6)-3 pm], width at base 2-(2.4)-3 a5 pm [2-5-(3-4)-4 pm]. Several examples of 'intermediate'-sized cladotylotes, 150-170 pm long, also occur, and the recognition of the two size categories defined above is based on a bimodal distribution of measured spicules. Acanthostyles of 2 sizes, divided on the basis of a bimodal distribution of spicule New Caledonian Poecilosclerids 1225 measurements; both size categories cylindrical, straight, fusiform, with slightly swollen base, with fewer spines in centre of shaft than at ends; larger acanthostyles relatively rare: length holotype 95-(111.7)-129 pm, [paratype 89-(113.3)-138 pm], width 2-(3.3)-4 pm, [2-(3 8)-4 pm]. Smaller acanthostyles very abundant: length holotype 69-(76.3)-84 pm [paratype 69-(77.2)-82 pm], width 2-(3.1)-4 pm [l 5-(2.8)-4 pm]. Microscleres. Only 2 varieties of toxas present. Oxhorn (type I) toxas absent. Smaller centrally curved (type 11) toxas very thin, with angular central curvature and recurved arms: length holotype 52-(93 -2)-141 pm [paratype 48-(78.0)-117 pm], width 0.5-(0.9)-1.2 pm [O-5-(0.9)-1.4 pm]. Larger accolada (111) toxas very long, slender, mostly straight or very slightly reflexed arms, with angular symmetrical or asymmetrical central curvature: length holotype 185-(269.6)-386 pm [paratype 119-(243.4-416 pm], width 0.4-(1.1)-1.8 pm [0.5-(1 3-3 pm]. Some intermediate examples between the type I1 and I11 toxa geometries were also observed, but generally these were smaller, probably developmental forms of one or the other type of toxon, and were not included in measurements given above. Palmate isochelae large, homogenous: length holotype 14-(18.4)-22 pm [paratype 17(19.5)-22 pm]. Fig. 1. Acarnus caledoniensis, sp. nov.: A , holotype QM G300724, choanosomal principal subtylostyles (enlarged spined base); B, ectosomal tylote; C, cladotylotes (type I); D, cladotylotes (type 11); E, acanthostyles; F, larger accolada (type 111) toxas; G, smaller centrally curved (type 11) toxas; H, palmate isochelae; I, fibre characteristics; J, section through peripheral skeleton. 1226 J. N. A. Hooper and C . Ldvi Distribution Known so far only from the south-west coast of the Noumea lagoon, in Halimeda beds, on sand and coral rubble substrate, rolling across the sea bed or only loosely attached to benthos, 5-25 m depth. Remarks Acarnus caledoniensis, sp. nov. is the 22nd member of the genus so far described, and ninth member belonging to the 'souriei' species-complex. It is also the most easterly record of this group in the Indo-Pacific (cf. van Soest et al. 1991: fig. 8). The species is most closely related to A. bicladotylota and A. tener, which together comprise a subgroup within the 'souriei' species-complex distributed on the western rim of the Pacific Ocean, from Japan to the Indo-Malay archipelago and western Micronesia. It also shows similarities to A. peruanus from the eastern Pacific. Other members of the 'souriei' species-complex include: A. levii (Vacelet, 1960) (from the Mediterranean), A. primigenius Hiemstra & Hooper, 1991 (from Indonesia), A . nicoleae van Soest, Hooper & Heimstra, 1991 (from the Caribbean), A. radovani (Boury-Esnault, 1973) (from Brazil), and A. souriei (LCvi, 1952) (restricted to populations living in west Africa-Senegal, Cape Verde and Gulf of Guinea). It has been shown elsewhere that these 'souriei'-like species have small but consistent differences from the west African A. souriei (s.s.). In general these morphological differences correlate with regional species distributions, supporting both the decision to recognise them as valid species, and the notion that they may have evolved from a previously widespread 'souriei'-like ancestor (van Soest et al. 1991). Therefore, as would be expected by its transitional distribution, Acarnus caledoniensis has several character states of intermediate condition between the western (A. bicladotylota, A. tener) and eastern (A. peruanus) Pacific subgroups. Fig. 2. Acarnus caledoniensis, sp. nov.: A , holotype QM G300724 (ORSTOM R806) in situ (photo P . Laboute); B, SEM of cladotylotes (magnified 600 x); C, SEM of skeletal structure (magnified 60x); D, SEM of echinating acanthostyles (magnified 600 x). New Caledonian Poecilosclerids Table 1. Comparison in spicule dimensions between Acarnus The new species is compared t o other related species in the 'sourieri' species complex. Measurements are given in micrornetres; comparative data from van Soest et al. (1991) A. caledoniensis A. souriei A. bicladotylota A. tener A. peruanus A. levii Massive Massive Encrusting 260-340 4-10 244-371 9-12 315-480 6-9 Smooth Spined Spined 180-330 2.5-5 21 1-282 3-4 260-460 3-4.5 121-141 4-5 180-210 4.5-6 66-75 3 80-140 2-4 Few Few Thick Thick 82-97 4 55-115 3-4.5 Absent Absent Absent 20-140 Subspherical Encrusting Shape Encrusting 266-423 4-11 160-350 3-10 195-460 5-12 Spined Rugose Spined 299-383 3-7 210-400 3-5 143-404 3-6 80-180 4-6 80-2 10 4-7 Absent 38-95 1-6 Styles Base of style Tylotes Cladotylotes I 130-190 6 Cladotylotes I1 Absent Spines on shaft of cladotylotes Few Many Thick Thick Thick 100-165 4-6 70-120 1-5 63-90 2-3 Absent 50-80 48-75 Many Many Basal part of shaft of cladotylotes Thin Acanthostyles I 70-130 4-6 Acanthostyles I1 Absent Toxas I ('oxhorn') Absent 70-1 10 Toxas I1 (deeply curved) 50-150 30-212 160-200 130-250 11-15 12-20 35-40 Toxas I11 ('accolada') Absent 188-21 1 Absent 12-18 13-17 13-16 Isochelae Acarnus caledoniensis is dark orange, and has a massive, subspherical growth form with large raised oscules on the upper surface (Fig. 24). It has a plumose, slightly reticulate skeleton, lacks oxhorn (type I) toxas, has much larger type I11 toxas with an upper range of 416 pm long, has two sizes of cladotylotes, two sizes of acanthostyles, and has generally 1228 J. N. A. Hooper and C. Ltvi larger styles and tylotes than its sibling species (refer to Table 1). By comparison, Acarnus souriei s.s. is red, invariably encrusting, has a plumose skeleton, has all three forms of toxas, type I11 toxas have an upper range of only 200 pm long, has only one size class of cladotylotes but two sizes of acanthostyles, and styles and tylotes are generally smaller. Acarnus peruanus is a thickly encrusting mat, has a plumoreticulate skeleton, also lacks oxhorn (type I) toxas, type I11 toxas have an upper size range of only 211 pm long, cladotylotes are smaller and consist of only one size class, there is only one size class of acanthostyles, tylotes are shorter and thinner, and acanthostyles are relatively rare as compared to those of A. caledoniensis. Acarnus bicladotylota is red-orange to blood red, encrusting with embedded detritus, without raised oscules, has a plumose skeleton, has all three forms of toxas, and type I11 toxas have an upper range of only 250 pm long, it also has two size classes of cladotylotes, of similar size to those in A. caledoniensis, but it has only one size of acanthostyles. Acarnus tener is red, massive, with a plumose skeleton, has all three forms of toxas, type I11 toxas have an upper range of only 230 pm long, and has only one category of cladotylotes and acanthostyles, similar in size to those found in A. caledoniensis. The new species should also be contrasted with A. levii-both which show most similarities in their cladotylote geometry and spination, but A. Ievii is bright red to maroon-red, encrusting, has a plumose skeleton, lacks accolada (type 111) toxa, has only one size of acanthostyles, although it also has two size classes of cladotylotes. In the present contribution, it is not appropriate to represent an emended worldwide phylogenetic classification of Acarnus (van Soest et al. 1991) to include the New Caledonian species, but it is possible to hypothesise on where A. caledoniensis fits into the proposed phylogenetic scheme depicted by those authors (van Soest et al. 1991: fig. 6 ) . As already noted, A. caledoniensis is most closely related to the 'souriei' group of species. It belongs to the subgroup of 'souriei' containing A. radovani, A. souriei, A. tener and A. nicoleae, based on similarities in skeletal structure (being plumose, an apomorphy for the 'souriei' group), and cladotylote morphology [with none or only minor thickening of the shaft below the clad; and the presence of hook-shaped spines along the shaft on both size categories of cladotylotes (both pleisomorphic conditions for the 'souriei' group)]. The new species branches from (and is more primitive than) this subgroup, in having microspined bases on styles (which is the ancestral condition; character I l a of van Soest et al. 1991), but it stands out from those species by its growth form (being subspherical and functioning like a 'tumbleweed', a new apomorphy for the genus), and the independent loss of type I 'oxhorn' toxas, which is the fourth occurrence of this (homoplastic) synapomorphy in the genus (also found in A. peruanus, A. deweerdtae, and A. tenuis, all belonging to separate species-gr oups). Etymology Named for the type locality. Family DESMACELLIDAE Ridley and Dendy Definition Sponges with choanosomal axial skeleton ranging from axially condensed, radial, reticulate, plumo-reticulate or halichondroid, produced by variably compressed reticulation of spicules enclosed in spongin fibres; extra-axial skeleton plumose or halichondroid. Megascleres typically monactinal styles or tylostyles, sometimes modified to oxeas or strongyles, occasionally with flexuous interstitial strongyles. Microscleres sigmas, toxas, microxeas, raphides (usually in trichodragmata), spheres or commas in all combinations (after Hooper 1984). Remarks There is only one other published record of Desmacellidae from the New Caledonian region, the hypercalcified sponge Merlia normani Kirkpatrick, 1908, published as M. deficiens Vacelet, 1980 (Vacelet 1981). New Caledonian Poecilosclerids 1229 Genus Neofibularia Hechtel Neofibularia Hechtel, 1965: 22 [type species Amphimedon nolitangere Duchassaing and Michelotti, 1864: 821 [full synonymy and discussion given by Hartman (1967: 3), van Soest (1984a: 77) and Hooper (1984: 33)]. Diagnosis Desmacellidae with an irregular isodictyal reticulate choanosomal skeleton, well-defined spongin fibres, membraneous or tangential ectosomal skeleton, typically smooth surface, megascleres mostly diactinal (strongyles, oxeas), sometimes styles; microscleres sigmas, raphides and microxeas (commas in one species); usually toxic (from Hooper et al. 1991). Remarks This definition is based on knowledge of only three species prior to the present work [i.e. N. nolitangere (Duchassaing & Michelotti, 1864), N. irata Wilkinson, 1978, and N. mordens Hartman, 19671, and from which the genus is supposedly clearly distinguished from the allied Biemna. On paper, Neofibularia has a reticulate skeleton (cf. plumoreticulate in most Biemna), diactinal (cf. monactinal) megascleres, and all species are toxic (v. no toxicity) (Hartman 1967: 6). It also apparently differs in having a well-developed spongin fibre skeleton (cf. poorly developed) and species may produce copious amounts of clear mucus (Bergquist and Fromont 1988: 30). Although one species of Biemna is also known to be toxic and have a renieroid reticulate skeleton, B. saucia Hooper et al., 1991, this species has probably retained ancestral characters no longer present in most other Biemna species (i.e. a primitive Biemna). Neofibularia hartmani, sp . nov. (Figs 3-4, Table 2) Material Examined Holotype. QM G300281 (ORSTOM R234; fragment NTM 23891): stn 106, N. entrance, RCcif des Cinq Milles, SW. New Caledonia lagoon, 22'29.3'S., 166"44.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA. Fig. 3. Neofibularia hartrnani, sp. nov.: A , holotype QM G300281, choanosomal strongyles; B, smaller microoxeas; C, larger microoxeas; D, raphides; E, large sigmas; F, small sigmas; G, section through peripheral skeleton. 1230 J. N. A. Hooper and C. LCvi Paratype. NTM 23892 (ORSTOM R1300): stn 192, E. coast IlBt Maitre, 22"20.01S.,16624.OrE., 1 m depth, 22.ix.1977, coll. P. Laboute, by hand. Other New Caledonian material. ORSTOM R97: stn 172, SE. Passe de Upe, RCcif de ]'Adventure, Ile des Pins, 22'31. 1'S.,167°30.0'E., 15 m depth, 7.i.1977, coll. A. Intes, SCUBA; ORSTOM R1244: stn 268, sand isles, RCcif Croissant, 22°20.2'S.,166022.5'E., 12 m depth, 25.ii.1980, coll. P. Laboute, SCUBA. Description Colour. Pale yellow-brown or khaki-brown alive (Munsell 2.5Y 8/10), often partially silt-covered, dark brown in ethanol. Shape. Thickly lobate, massive, bulbous-encrusting sponge with lobes/bulbs fused to adjacent bulbs, 20-55 mm diameter, up to about 40 mm high, spreading about 120 mm across dead coral substrate. Surface. Well developed sculpturing, consisting of close-set, interconnected microconules, forming web-like striations on surface. Large oscules situated on apex of bulbs, 2-8 mm diameter, with slightly raised rims, contracting to smaller diameter when preserved. Ectosome. Membraneous, lightly arenaceous, with abundant heavily pigmented spongin, but without specialised spicule skeleton. Irregular plumose tufts of choanosomal strongyles protrude through surface, paratangential or erect on surface, more-or-less corresponding to position of surface conules; ectosome rarely intact in preserved material. Choanosome. Skeletal architecture reticulate, with differentiated primary (multispicular) and secondary (uni- and paucispicular) spongin fibres. Primary fibres usually fully cored with choanosomal strongyles, 50-90 pm diameter; secondary fibres with fewer spicules Fig. 4. Neofibularia hartmani, sp. nov.: A, holotyue OM G300281 (ORSTOM R234) in situ (photo P. Laboute); B, SEM of fibre node (magnified 4 0 0 ~ ) ;C, SEM of skeletal structure (magnified 200 x); D, SEM of sigma microscleres (magnified 1000 x ). New Caledonian Poecilosclerids 1231 packed abreast, 15-40 pm diameter. Fibre reticulation produces elongate, oblong or oval meshes, 35-140 pm diameter, with moderate quantities of collagen and foreign particles, and abundant microscleres. Choanocyte chambers not observed, obscured by foreign debris, microscleres and loose megascleres. Megascleres (refer to Table 2 for dimensions). Choanosomal strongyles long, thick or thin, slightly curved, symmetrical or asymmetrical. Table 2. Comparison in spicule dimensions between species of Neofibularia Measurements are given in micrometres. Spicule numbering convention follows Wilkinson (1978). Comparative material of N. irata consists of previously unpublished specimens from the Great Barrier Reef (NTM 21113-14) N,hartmani, sp. nov. N. irata Wilkinson N.nolitangere (Hartman) N. mordens Hartman Source of measurements Present study Present study 218-(257.3)-280 3-(5.3-9 (strongyles) 128-(163.4)-179 1.5-(2.1)-3.0 (strongyles) 28-(38.0)-51 0.5-(0.9)-1.2 24-(31.2)-37 0.4-(0.5)-0.8 69-(79.1)-94 0.8-(1.6)-2.0 54-(58.3-67 0.8-(1.1)-1.5 Absent Absent 79-(86.5)-115 0.2-(0.6)-0.8 60-(67.4)-84 0.2-(0.3)-0.3 13-(22.4)-36 0.8-(1.1)-1.5 14-(31.8)-41 0.5-(0.9)-1.5 50-(67.3-85 1.8-(2.4)-3.5 Absent Hartman (1967) Hartman (1967) Choanosomal megascleres 167-(326)-433 3.1-(9.9)-24.4 (strongyles and styles) 235-(314-357 4.4-(10.2)-13.4 (styles) Absent Absent Microxeas I Microxeas I1 Absent Microxeas 111 98-(122)-150 1.8-(2.9)-5.2 Raphides 82-(105)-126 0.6-(1.1)-2.0 Absent Sigmas I 14-(19)-24 0.9-(1 .+Is6 Sigmas I1 Absent Sigmas I11 Absent Absent Absent Absent Absent Commas 5-(7.9)-12 Absent J. N. A. Hooper and C. L6vi 1232 Microscleres. Microxeas clearly divided into two sizes, both relatively long, stout, widest at midsection. Raphides abundant, long, straight, hair-like. Sigmas incompletely divided into two sizes, both with c- and s-shaped examples, relatively thick, evenly curved at centre. Distribution Known only from the southern New Caledonian lagoon, from the reef flat and shallow subtidal, on dead coral substrate, 1-15 m depth. Remarks This species is most similar to N. irata in the diversity and geometry of its spicules, but otherwise the two species differ substantially in most features: spicule dimensions (Table 2), growth form (Fig. 4A; N. irata is a flat spreading sponge with erect, leaf-like, folded lamellae), surface features (N. irata has a smooth unornamented surface), and skeletal structure (N. irata has a fragile, flattened, honeycomb-like fibre reticulation, cored by whispy, hair-like tracts of thin strongyles). It is also suspected that N. hartmani produces a toxic reaction in humans upon contact with skin (Dr C. Debitus, personal communication), as is typical for the genus, but this has yet to be substantiated experimentally. Etymology This species is named for Professor Willard Hartman, Peabody Museum of Natural History, Yale University, for his pioneering work on Neofibularia, and his other eminent contributions to the biology of the Porifera. Family CRELLIDAE Hentschel Definition Poecilosclerida with a thick ectosomal skeleton, usually a crust of tangentially placed, densely packed acanthoxea or acanthostyles, a regularly reticulate choanosomal skeleton of smooth diactinal megascleres grouped into vague bundles, sometimes reduced to a basal layer of acanthostyles erect on substrate in encrusting forms, and spined megascleres may also be distributed within choanosome, between tracts of smooth diactines. Microscleres may include arcuate isochelae, sigmas, less often arcuate anisochelae (modified from van Soest 1984~). Remarks There is only a single previous record of the family Crellidae from the New Caledonian region, Yvesia acanthosclera LCvi & LCvi, 1983b, from deeper waters (425-430 m depth). Genus Crella Schmidt Crella Gray, 1867: 521 [type species Cribrella elegans Schmidt, 1862: 701 [full synonymy and discussion of the literature given by Bergquist and Fromont (1988: 76)]. Diagnosis Crellidae with an ectosomal skeleton composed of erect or tangential acanthoxeas in densely packed tracts; primary choanosomal skeleton with plumose, plumo-reticulate or reticulate spongin fibres cored by diactinal megascleres (oxeas or tornotes), with or without echinating acanthostyles; secondary choanosomal skeleton a more-or-less even renieroid reticulation overlaying primary skeleton; in encrusting forms skeleton reduced to a basal layer of fibres lying on substrate with erect acanthostyles embedded in spongin; microscleres usually include arcuate isochelae, sometimes arcuate anisochelae, and sigmas. Remarks The genus Crella has not been substantially revised in the contemporary literature, although descriptions of several species have been published since the 1980s (e.g. van Soest New Caledonian Poecilosclerids 1233 1984a; Bergquist and Fromont 1988). These works (and Hallmann 1912) should be consulted for a more detailed overview of the genus. Crella spinulata (Hentschel) (Figs 5-6, Table 3) Grayella spinulata Hentschel, 1911: 340, text-fig. 29. Crella spinu1ata.-Burton, 1934: 553; Hooper et al., 1992: 263. Material Examined Holotype. ZMH: presently unknown. New Caledonian material. QM G300014 (ORSTOM R1228): stn 261, SW. Il6t Nda, Lagon Sud, 21°52.5'S,166"51~2'E., 33 m depth, 4.xii.1979, coll. P. Laboute, SCUBA; NTM 23871 (fragment QM G300686): Baie des Citrons, Noumea lagoon, 22°18.0'S.,166025.2'E., 3 m depth, 25.ix.1990, coll. M. Kelly-Borges, snorkel. Comparative material. Houtman-Abrolhos Islands, W.A.: NTM 22916, 2917 (fragment QM 300194): N. of North Wallabi I., 28°17.9'S.,113047.8'E., 39 m depth, ll.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademic Oparin', beam trawl; QM G300221 (fragment NTM 23216): N. edge of Pelsart I., 28°47.2'S.,113058.5'E., 22 m depth, 10.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', SCUBA; NCI Q66C-4228-M (fragment QM G300687): NW. side of Goss Passage, Wallabi Group, 28929'S.,113°37'E., 18 m depth, 13.ix.1990, coll. NCI, SCUBA; NCI Q66C-4629-W (fragment QM G300502): top of Assail Bank, between North I. and Wallaby Group of Is, 28'211S., 113"42'E., 15 m depth, 16.ix.1990, coll. NCI, SCUBA. Northwest Shelf, W.A.: NTM 2645: W. of Eighty Mile Beach, 19°33.1'S.,119005.1'E., 35 m depth, 04.v.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 2674: N. of Port Hedland, 19"16'S.,118"50'E., 70 m depth, 04.v.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 2746: N. of Eighty Mile Beach, 19°16'S.,118050'E., 75 m depth, 04.v.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 2744: N. of Adele I., Collier Bay, 15'58.3'S., 122"39.7'E., 59 m depth, 21.iv.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 21771: W. of Port Hedland, 19°03.3'S.,l18049.9'E., 82 m depth, 29.viii. 1983, coll. Trevor Ward, R.V. 'Soela', trawl; NTM 22295: NW. of Lacepede Is, 16°31'S.,121028'E., 38-40 m depth, 17.iv.1985, coll. Barry Russell, pair trawl; NTM 23038: N. of Amphinome Shoals, 19°19.7'S.,119008.8'E., 50 m depth, 19.vii.1987, coll. J. N. A. Hooper, USSR R.V. 'Akademik Oparin', beam trawl. Darwin Harbour, N.T.: NTM 22695: East Point Aquatic Life Reserve, 12°24.5'S.,130048~O'E.,9-12 m depth, 03.iv. 1986, coll. J. N. A. Hooper, A.-M. Mussig, C. Hood, SCUBA. Arafura Sea: QM G301206: Flattop Bank, NE. Joseph Bonaparte Gulf, 12°16'S.,129015'E., 17.v.1992, 32 m depth, coll. J. N. A. Hooper, dredge. Great Barrier Reef, Qld: NTM 21513: E. of Murdock I., Howick Group, 14°36'S.,145003'E., 14 m depth, 18.ix.1979, coll. Qld Fish. Service; NTM 21575: E. of Coquet I., Howick Group, l4"32.2'S., 145"07.5'E., 22 m depth, date unknown, coll. Qld Fish. Service; NTM 21568: Stanley Reef, outer slope and lagoon, 19°16'S.,148006'E, 13 m depth, 15.x.1980, coll. Alice Kay; NTM 23514: Dingo Reef, NE. edge of reef front, Whitsunday Is Group, 19°07.2'S,148028.7'E., 12 m depth, 20.xi.1988, coll. NCI, SCUBA. Description Colour. Red-orange alive (Munsell 2.5R 5/10), orange-brown in ethanol. Shape. (QM G300014): Groups of solitary, conical or cylindrical digits, up to 60 mm high, 20 mm basal diameter, 9 mm diameter at apex, erect on substrate, with common basal attachment on or below substrate. Digits evenly cylindrical or slightly flattened. (NTM 23871): Bulbous encrusting in sand sediment. Surface. Slightly conulose, granular surface, with visible ectosomal membrane. Oscules prominent in live material, evenly dispersed over sides and apex of digits, about 1-2 mm diameter, slightly raised above surface 2-3 mm high. Ectosome. Thick crust of acanthoxeas lie tangential to surface, 100-350 pm thick, with plumose brushes of smooth oxeas, arising from major choanosomal spicule tracts, standing perpendicular to surface crust and protruding slightly through it, but more-or-less obscured by tangential surface crust. Moderately light granular collagen permeates surface crust. 1234 J. N. A. Hooper and C . Lhi Choanosome. Skeleton composed of 2 distinct structures. Main skeleton a meandering reticulation of thick, well-formed spongin fibres fully cored by multispicular tracts of smooth oxeas, 50-85 pm diameter, eventually culminating in erect plumose brushes of smooth oxeas at surface. Fibre meshes relatively cavernous, 140-220 pm diameter, oval to eliptical, with light collagen visible around eliptical choanocyte chambers, 60-130 long, 30-50 pm wide. Secondary skeleton renieroid, more-or-less regularly reticulate, overlaying primary reticulate fibre skeleton, each element of renieroid reticulation composed of 2 or more acanthoxeas joined at nodes by collagenous spongin. Secondary skeleton ceases about 200-250 pm from surface crust. Most megascleres confined to one or the other skeletal structures, few spicules scattered within mesohyl. Megascleres (refer to Table 3 for dimensions). Choanosomal fibres cored by smooth oxeas, long, thin, slightly curved, usually symmetrical, occasionally modified to styloids, usually with hastate points. Ectosomal and choanosomal acanthoxeas identical in morphology and size, long, thin, symmetrically curved, with small granular spines evenly dispersed over spicule and with sharp fusiform points. Microscleres. Isochelae arcuate, moderately large, uncommon; thinner, poorly silicified examples also seen, with rudimentary development of alae, presumably juvenile forms of large chelae. Distribution Tropical Australasia: Shark Bay, W.A. (Hentschel 1911); Houtman-Abrolhos Islands, W.A.; Northwest Shelf, W.A.; Timor Sea, N.T.; Darwin Harbour, N.T.; Penguin Channel, Snake Reef, Stanley Reef, Howick Islands and Whitsunday Islands, Great Barrier Reef, Qld (6-55 m depth) (Burton 1934; present study); and New Caledonia (3-33m depth; present study). Partially burrowing into soft sediments (sand, gravel, shell-grit, coral rubble), or bioeroding dead coral substrates, rarely found on live coral substrate. Fig. 5 . Crella spinulata (Hentschel): A, specimen QM G300014, choanosomal oxeas; B, ectosomal and choanosomal acanthoxeas; C,arcuate isochela; D, section through peripheral skeleton. New Caledonian Poecilosclerids 1235 Remarks The holotype of this species is presently unknown, possibly extant in the ZMH. Nevertheless, the species is perfectly recognisable from Hentschel's (1911) description. In addition, abundant fresh material has been recollected from the south-, mid- and north-west coast of Western Australia (including from areas close to the type locality in Shark Bay), and also Fig. 6. Crella spinulata (Hentschel): A , specimen QM G300014 (ORSTOM R1228) in situ (photo P. Laboute); B, specimen ORSTOM R177 in situ (photo P. Laboute); C, specimen QM G300014 (ORSTOM R1228), SEM of peripheral skeleton (magnified 150x); D, SEM of acanthoxeas (magnified 1000 x ). Table 3. Comparison in spicule dimensions between specimens of CreNa spinulata (Hentschel, 1911) Measurements are given in micrometres; n, number of specimens examined - - - - New Caledonia Holotype Shark Bay, W.A. Present study (n = 2) Hentschel 1911 - West coast Australia, W.A. East coast Australia, Qld Present study (n = 14) Present study (n = 4) Source L W 256-(244.0)-33 1 1.8-(2.9)-4.0 Choanosornal smooth oxeas 223-(236.8)-271 264-328 3.0-(5.8)-7.0 3-5 L W 148-(152.4-157 1.5-(1.8)-3.0 120-152 4-5 228-(288.5)-355 2.5-(4.9-7.0 Acanthoxeas 95-(119.3)-147 1 .O-(4.3)-7.0 122-(143 .O)-162 2.0-(4.1)-6.0 12-(19.0)-28 16-(21.2)-25 Arcuate isochelae L 18-(20.2)-22 16-20 1236 J. N. A. Hooper and C . Lkvi from the north-east coast of Queensland. Some of this material has already been analysed in an earlier chemotaxonomic study (Hooper et al. 1992). There is little doubt, on paper at least, that all this material is conspecific with Hentschel's species, but it is also possible that presently undetectable sibling species make up C. spinulata, given the great variation in growth forms possible. A digitate branching morphology is typical, but encrusting and massive specimens are also known. The two representatives of the Noumea population, described above, represent new locality records for the species and differ from most other specimens collected from Australian waters in having only acanthoxeas, i.e. in many specimens of this species, and indeed in many species of Crella, acanthoxeas frequently co-exist in equal proportions with acanthostyles, or sometimes only acanthostyles are present (e.g. Bergquist and Fromont 1988). This feature may eventually be found to correlate with other differences (e.g. growth forms), and hence be important in differentiating possible sibling species, but for the present it not possible to clearly differentiate New Caledonian and Australian specimens on the basis of morphometric features. Family MICROCIONIDAE Carter Definition (emended) Poecilosclerida with ectosomal skeleton composed of styles or anisoxeas (exceptionally oxeas), in erect bundles, forming continuous crust, lying tangential or sparsely dispersed on surface. Choanosomal (or axial) skeleton with well-developed spongin fibres forming hymedesmoid, microcionid, plumose, plumo-reticulate, reticulate or axially condensed tracts. Subectosomal (or extra-axial) skeleton rarely well developed (cf. Raspailiidae), although subectosomal (auxiliary) spicules may be dispersed between fibres, sometimes forming ectosomal structures, and occasionally producing distinct extra-axial tracts (Axociella). Spongin fibres cored by smooth or partially spined large styles, occasionally modified to quasidiactinal forms (Echinochalina) and sometimes replaced partly or entirely by detritus (Clathriopsamma), and with wholly or partially spined small styles (acanthostyles), or modified forms such as acanthoxeas, acanthostrongyles or smooth styles embedded perpendicular to (i.e. echinating) fibres. Microscleres include diverse forms of isochelae, toxas and occasionally raphides or microxeas. Remarks The Microcionidae is a vast and diverse group of the Porifera. Worldwide there are 522 nominal species, of which probably 432 species are valid, and in the Australian fauna (i.e. including coastal, shelf, oceanic island and subantarctic island species) there are 140 valid species (or about 32% of the world's microcionids). In the past, reliable species identifications of Microcionidae have amounted to little more than educated guesswork, considering that the literature is vast, scattered, and many species are neither adequately illustrated nor well recognisable from their published descriptions. Furthermore, species seem to exhibit virtually every combination of characters, and virtually every character has dozens of character states, making the clear delineation of even the genera difficult (e.g. Hooper 1991~).Towards a solution, a major revision of species and higher taxa within the family is currently in progress, which will make the Australian fauna, at least, more accessible. Prior to the present work, only two published records of Microcionidae existed for the New Caledonian region, Antho novizelanica (Ridley, 1881) (LCvi and LCvi 1983b) from deeper water (250-350 m depth; also found in New Zealand), and a shallow-water species originally assigned to Clathria australiensis (Carter) from southern Australia (LCvi 1967), but which is described here as a new species (Clathria rugosa, sp. nov.). The present study adds another 12 species to the shallow lagoon and coral reef fauna, bringing the total number of microcionid species known for the region to 14, of which 10 are suspected endemics. New Caledonian Poecilosclerids Genus Clathria Schmidt Restricted synonymy Clathria Schmidt, 1862: 57 [type species Clathria compressa Schmidt, 1862: 581. Thalysias Duchassaing and Michelotti, 1864: 82 [Spongiajuniperina Lamarck, 1814: 4441. Clathriopsamma Lendenfeld, 1888: 227 [type species Clathriopsamma reticulata Lendenfeld, 1888: 2271. Axociella Hallmann, 1920: 779 [Esperiopsis cylindrica Ridley and Dendy, 1886: 3401. Dendrocia Hallmann, 1920: 767 [Clathriapyramida Lendenfeld, 1888: 2221. Isociella Hallmann, 1920: 784 [Phakellia jacksoniana Dendy, 1897: 2361. Diagnosis (emended) Microcionidae with 1 (subgenus Clathria) or 2 (subgenus Thalysias) categories of auxiliary styles or subtylostyles, sometimes absent entirely, forming peripheral skeleton; choanosomal spicules usually enclosed within spongin fibres, sometimes simply with nodal spongin, consisting of principal styles or subtylostyles usually well differentiated from auxiliary styles; sometimes fibres cored by undifferentiated auxiliary spicules (subgenus Dendrocia) or simply cored by detritus with or without megascleres (subgenus Clathriopsamma); echinating acanthostyles usually present, sometimes absent (subgenera Axociella, Isociella), occasionally with vestigial spination or completely smooth; choanosomal skeleton varies from leptoclathriid, microcionid, plumose, plumo-reticulate, renieroid (subgenus Isociella), to reticulate, with (subgenus Axociella) or without structural differences between axial and extra-axial skeletons; microscleres present or absent, typically consisting of palmate isochelae but sometimes including other diverse forms of isochelae, and smooth or spiny toxas. Remarks Within Clathria (with five subgenera recognised here) there are 383 described species worldwide, of which 328 appear to be valid; there are also another 9 recognisable but as yet unnamed species in the literature, as well as many other unrecognisable species. As noted above, only a single record of the genus has previously been published for the New Caledonia region, Clathriopsamma australiensis (Carter) (LCvi 1967), although this population is now considered to represent a new sibling species of the Australian taxon. The emended definition of Clathria presented here allows the inclusion of species with isochelae other than the usual palmate form. Various nominal genera have been created for species otherwise similar to Clathria, but with modified chelae, such as cleistochelae [e.g. Wetmoreus de Laubenfels (no material seen)], arcuate isochelae [e.g. Quizciona de Laubenfels (BMNH 1901.12.26.13), Paradoryx Hallmann (AM G3046)], anchorate isochelae [e.g. Cionanchora de Laubenfels (MNHN DT939), Folitispa de Laubenfels (BMNH 1921.11.7.69)], and bidentate sigmoid isochelae [e.g. Damoseni de Laubenfels (SMF969)l. Under some interpretations (e.g. Burton 1959), these genera should be assigned to different families, whereas Hooper (1991a) has suggested that their importance at the suprageneric level is questionable (see also Remarks above for Myxillidae and Acarnus). The use of the subgeneric classification, rather than the retention of these taxa at the generic level, is based on a phylogenetic analysis of the Microcionidae (Hooper 1991a). The development of this system, to apply to a geographically wider fauna, is presently in development by the senior author. Clathria (Clathriopsamma) rugosa, sp. nov. (Figs 7-8, Tables 4-5) Clathria australiensis. - Ldvi, 1967: 22, fig. 6, pl. 2, fig. D. Not Wilsonella australiensis Carter, 1885: 366. Material Examined Holotype. QM G300278 (fragment NTM 23880): Ile aux Canards, Noumea lagoon, New Caledonia, 22°19~0'S.,166026~O'E., 22 m depth, 26.ix.1990, coll. J. N. A. Hooper, SCUBA, in sand, rubble, sea grass beds. 1238 J. N. A. Hooper and C. LCvi Paratype. QM G300696 (fragment NTM 23889): Baie des Citrons, Noumea, New Caledonia, 22"18'S.,l66"25'E., 3 m depth, 15.ix.1990, coll. M. Kelly-Borges, snorkel, fringing coral reef. Other New Caledonian material. Nominotypical variety (flabellate-digitate, without astrorhizae): ORSTOM R235: stn 182, NE. IlBt KouarC, 22°46.1'S.,1660471'E., 24 m depth, 7.vi.1977, coll. P. Laboute, SCUBA; ORSTOM R194: stn 185, Pt Eacho, Lifou (ChCpCnCe), 20°47.0'S.,167007.1'E., 12 m depth, 19.vi.1977, coll. P. Laboute, SCUBA; ORSTOM R547: stn 150, S. of entrance, Baie St Vincent, 22"02.2'S.,165"59.5'E., 18 m depth, 9 . k 1976, coll. P. Laboute, SCUBA; ORSTOM R276: stn 193, E. coast, Rtcif Tabu, 22°29.1'S.,166026.1'E., 30 m depth, 13.x.1977, coll. G. Bargibant, SCUBA; ORSTOM R558: stn 148, NE. pt, IlBt Mboa, 22°08.3'S.,166009.3'E., 13 m depth, 9.ix.1976, coll. P. Laboute, SCUBA; ORSTOM R771: stn 206, Banc de la Torche, Ile des Pins, 22051.11S., 167040.01E.,36 m depth, 8.iii.1978, coll. P. Laboute, SCUBA; ORSTOM R1288: stn 302, ENE. lagon Belep, Lagon Nord de la Grande terre, 19°39~0'S.,161"33.OIE., 38 m depth, 22.vi.1981, coll. J.-L. Menou, SCUBA; ORSTOM R620: stn 124, IlBt Maitre, 22°20.1'S.,166025.1'E., 20 m depth, 15.vii. 1976, coll. P. Laboute, SCUBA; ORSTOM R466: stn 109, Baie des Citrons, Noumea, 22018.01S., 166"25.11E., 10 m depth, 31.v.1976, coll. P. Laboute, SCUBA. Variety with astrorhizae, flabellatedigitate: QM G301265 (ORSTOM R157) ('representative specimen'): stn 124, I16t Maitre, 22"20,11S., 166"25.11E., 20 m depth, 30.xi.1977, coll. G. Bargibant, SCUBA; ORSTOM R586: stn 147, midchannel between Ton Du (Tiart) and IlBt Ie, 22°121'S.,166016~5'E., 24 m depth, 8.ix.1976, coll. P. Laboute, SCUBA: ORSTOM R260: stn 114, NE. IlBt TarCti, Banc Gail, 22°221'S.,166039~0'E., 35 m depth, 29.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R263: stn 110, SE. IlBt RCdika, 22'31. 1'S.,166"36.01E., 19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R428: stn 124, IlBt Maitre, 22°20.1'S.,1660251'E., 20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM R404: stn 114, NE. Ilat Tartti, Banc Gail, 22°22~1'S.,166039~0'E.,35 m depth, 29.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R498: stn 124, IlBt Maitre, 22°20.1'S.,166025~1'E., 20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM R626: stn 162, W. (lee) of Ilat KiC (Havannha), 22°221'S.,167004~WE.,15 m depth, 26.x.1976, coll. A. Intis, SCUBA; ORSTOM R1069: stn 249, NW. I16t Vua, Lagon Sud, 2lo33.4'S. 166"43.5'E., 35 m depth, 10.xii.1978, coll. P. Laboute, SCUBA; ORSTOM R1304: stn 315, NE. IlBt Dii-Ami, Ile des Pins, 21°35.5'S.,167017.4'E., 33 m depth, 30.vii.1981, coll. P. Laboute, SCUBA. Variety with fistules, massive: QM G301261 (ORSTOM R1418) ('representative specimen'): stn 453, Nokanhoui, Ile des Pins, 22"43.3'S.,167O35.2'E., 40 m depth, 27.xi.1983, coll. J.-L. Menou, SCUBA; QM G300870 (fragment NTM 23881, ORSTOM cfR53): stn 158, Baie de Saint-Vincent, between Passe de St Vincent and Hugon, 22°03.0'S.,165059.0'E., 10 m depth, 16.ix.1976, coll. P. Laboute, SCUBA. Description Colour. Bright red-orange alive (Munsell 5R 5/10), rarely dark orange (10R 6/10); light grey-brown in ethanol. Shape. Flabellate, palmate-digitate or simply digitate sponges, occasionally massive, subspherical, varying considerably in size (70-340 mm long, 45-140 mm maximum width, 10-20 mm thickness), and varying in development of digits, typically consisting of several short or long cylindrical digits (15-140 mm long), arising from margins of a fan, or at least with an enlarged point of attachment for digits above basal stalk; usually with short cylindrical stalk (30-50 mm long) often attached to coral rubble or dead molluscs. Surface. Nominotypical variety. Optically smooth, even, membraneous surface when alive, with few microconules on margins of digits, none on lateral surfaces of fans; ectosomal membrane destroyed upon preservation, surface beneath membrane very porous, with numerous ostia approximately 0.5-1 mm diameter. Oscules, 2-3 mm diameter, surrounded by symmetrical membraneous lips which usually collapse upon preservation, slightly raised (up to 2 mm) from surface, without any obvious subectosomal canals (astrorhizae); oscules located mainly on margins or less frequently scattered over lateral surfaces in live material, but these are not easily visible after preservation. Variety with astrorhizae. Oscules raised above surface with prominent subectosomal canals (astrorhizae) converging towards central pore, and surrounded by slightly raised membraneous lips, producing Variety with fistules. Oscules on upper and lateral prominent subectosomal sculpturing. surfaces (of subspherical massive growth forms), raised above surface on relatively large fistules, up to 12 mm long, 4 mm diameter, without subectosomal canals (astrorhizae) or other surface sculpturing. However, fistules temporary, membraneous, collapsing upon preservation. New Caledonian Poecilosclerids 1239 Ectosome. Single category of auxiliary spicules (subectosomal subtylostyles) forms irregular paratangential brushes on surface. Surface mineral skeleton not dense, usually consisting of bundles, occasionally individual spicules embedded in or dispersed between relatively close-set, horny spongin fibres; fibres marginally smaller but more compacted (80-140 pm mesh size) in peripheral skeleton than in axis of choanosome; small quantities of detritus (mostly sand grains) embedded in peripheral skeleton, but less abundant than in axis; light brown collagen and numerous microscleres abundantly scattered in periphery. Plumose tracts of choanosomal styles, in paucispicular tracts, terminate just below auxiliary spicules in the peripheral skeleton, and do not protrude through the sponge surface. Choanosome. Skeleton reticulate, spongin fibres heavy, relatively widely spaced (230400 pm mesh size); spongin fibres divisible into primary and secondary components; primary fibres (80-115 pm diameter) sparsely cored by uni- or paucispicular tracts of choanosomal principal styles, usually ascending towards peripheral region; primary fibres heavily echinated by acanthostyles, dispersed relatively evenly over all fibres; many primary fibres also contain sand grains, but these do not usually occupy entire fibre diameter; secondary fibres (20-45 pm diameter) usually uncored, without detritus, but heavily echinated by acanthostyles; collagen moderately heavy, containing abundant microscleres; choanocyte chambers oval, 50-80 pm diameter. Megascleres (refer to Tables 4-5 for dimensions). Choanosomal principal spicules short, straight styles with slightly subtylote bases, rounded or abruptly pointed tips, usually Clathria (Clathriopsarnrna) rugosa, sp. nov.: A , paratype QM G300696, subectosomal auxiliary subtylostyles (quasidiactinal); B, choanosomal principal style and acanthose style; C, echinating acanthostyles; D, toxas; E, palmate isochelae; F, section through peripheral skeleton; 6, fibres characteristic; H, fibre node. Fig. 7. 1240 J. N. A. Hooper and C. L h i completely smooth, occasionally with sparse spines over shaft and both ends; similar in morphology to auxiliary spicules, differentiated mainly by being slightly shorter and thicker. Subectosomal auxiliary spicules quasidiactinal modifications of styles, long, straight with slightly subtylote microspined bases, rounded microspined points, completely smooth shaft. Echinating acanthostyles short, cylindrical, slightly subtylote bases, rounded, slightly swollen tips, evenly spined; spines small, granular. Microscleres (refer to Tables 4-5 for dimensions). Palmate isochelae small. Toxas with consistent morphology, variable in size, slightly curved at centre, recurved and sharply pointed at tips. Fig. 8. Clathria (Clathriopsamma) rugosa, sp. nov.: A , paratype QM G300696, SEM of skeletal structure (magnified 1 0 0 ~ ) ;B, SEM of fibre characteristics (magnified 600x); C, nominotypical variety, specimen in situ (ORSTOM R53) (photo P. Laboute); D, specimen in situ (ORSTOM R450) (photo P. Laboute); E, variety with astrorhizae, specimen in situ (ORSTOM R626) (photo P. Laboute); F, variety with fistules, specimen in situ (ORSTOM R771) (photo P. Laboute). New Caledonian Poecilosclerids 1241 Table 4. Comparison in spicule dimensions between Clathria rugosa, C. australiensis and C. bulbosa Clathria rugosa is compared with the allopatric populations of its sibling species C. australiensis (Carter), as well as the sympatric C. bulbosa, sp. nov. Measurements are given in micrometres C. rugosa, sp. nov. C. bulbosa, sp. nov. C. australiensis* (New Caledonia) (New Caledonia) (Vic.) 134-159 4.5-8 Principal 124-180 2-4 Auxiliary 162-206 2.5-4 Spined 184-207 1.5-3 Smooth 55-74 2-4 32-92 0.8-2.5 14-19 A Choanosomal styles 92-152 2.5-4 Principal C. australiensisB (N.S.W.) C, australiensisC (W-4.) 92-146 1.5-4 Principal 106-173 2.5-5 Principal Subectosomal auxiliary styles 92-155 94-141 1.5-4 2-3.5 Spined Spined Echinating acanthostyles 45-77 2-4.5 54-78 2-4.5 Toxas 28-89 0.5-1 32-74 0.5-1 Isochelae 11-18 12-16 98-172 2-5 Spined Lectotype BMNH 1886.12.15.43. Holotype of Clathiopsamma lobosa Lendenfeld (a synonym of C. australiensis), AM G9053 Holotype of Thorects ramsayi Lendenfeld (a synonym of C. australiensis), AM G8820. Table 5. Comparison in spicule dimensions between specimens (varieties) of Clathria rugosa Measurements are given in micrometres Holotype nominotypical (QM G300278) Paratype variety (QM G300696) 'Fistulose' variety (QM G300870) 'Astrorhizae' variety (QM G301265) Choanosomal principal styles L W 134-(147.8)-159 4.5-(5.8)-8 L W 162-(188.2)-206 2-(2.8)-4 135-(140.0)-146 4.545.9-7 117-(132.6)-156 2.5-(4.4)-6 134-(145.6)-159 4-(4.9)-6 Subectosomal auxiliary styles 180-(184.8)-194 2.5-(3.3)-4.5 144-(172.0)- 192 2.5-(3.1)-4 173-(182.0)-204 2.5-(3.2)-4 Echinating acanthostyles L W 66-(74.2)-91 4.5-(4.9)-6 58-(67.2)-77 4-(5.5)-7 59-(65.8)-72 3-(4.2)-5 55-(65.9-75 3.5-(4.2)-5 L W 24-(75.2)-122 0.8-(1.8)-2.5 Toxas 35-(68.3)-84 0.8-(1.9)-3 32-(58.2)-84 0.8-(1.9)-3.0 32-(63.2)-117 0.8-(1.5)-2.5 L 13-(15.1)-17 Isochelae 12-(14.0)-16 12-(14.2)-18 13-(14.3)-16 1242 J. N. A. Hooper and C. Ltvi Distribution Known so far only from the southern and south-west regions of the New Caledonian lagoon, on dead coral, coral rubble, Halimeda beds and sand substrates; depth range 3-38 m. Remarks There are three distinct populations of this species in the New Caledonian lagoon, as described above. As far as is known, these populations are sympatric, appear to occupy similar ecological niches, and two are virtually identical in their external morphology, coloration, skeletal structure, spicule geometry and spicule size (Table 9, whereas the third, fistulose variety differs more substantially in being subspherical (Fig. 8F) rather than flabellate (Fig. 8C, E) or digitate (Fig. 80). However, these varieties are apparently easily differentiated in the field by their surface sculpturing: the nominotypical variety has oval, symmetrical, slightly raised oscules; the variety with astrorhizae has prominent subectosomal canals (astrorhizae) converging towards the centre of each oscule; the variety with fistules has oscules raised relatively further above the surface, on short membraneous stalks. SCUBA divers report that these features are distinctive in situ (J.-L. Menou, personal communication), although they are virtually undetectable after preservation. These three populations are not formally differentiated here (at the subspecies level) because no other conventional morphological characters can be found to distinguish them. Nevertheless, the described material of each variety is separated above to emphasise these apparently prominent field characteristics, but only non-morphological evidence will eventually determine whether or not their formal separation is warranted. Clathria rugosa is a member of the subgenus Clathriopsamma in having its mineral spicule skeleton supplemented by detritus, mostly sand grains (Fig. 8B). However, this feature may have a fairly high level of homoplasy for the Demospongiae, probably independently acquired in several other Poecilosclerida (e.g. Desmacididae) and other demosponges (e.g. Dysideidae), and must be re-evaluated in a phylogenetic context. Spongin fibres on the ectosome of C. rugosa are more closely compacted than fibres within the choanosome, and this feature is reminiscent of the typical Callyspongia condition (Haplosclerida), but it is undoubtedly one of convergence. Clathria rugosa is also similar to a small group of other Clathria species- C. (Thalysias) major, C. (Dendrocia) pyramida and C. (Clathriopsamma) australiensis-in having microspines on both the bases and tips of auxiliary spicules (Fig. 7 A ) . These spicules at first appear to be diactinal megascleres, and as such suggest affinities with the Myxillidae, but they are in fact modified styles (i.e. they are always asymmetrical), and the feature is interpreted here as being one of convergence with the Myxillidae. Clathria rugosa and C. australiensis are obviously closely related, showing many similarities in their skeletal architecture, spiculation and growth form, but there are several significant differences between them that justify the creation of a new species for the New Caledonian populations. Moreover, it is possible that the widely distributed, temperate Australian C. australiensis itself consists of several heterogenous populations, as each geographic morph differs slightly from the 'original' population from Port Phillip, Vic. There is presently not enough contemporary material (i.e. in good condition, with accurate locality data), nor any available non-morphological, corrobative evidence to divide these populations into sibling species. Clathria rugosa has field characteristics notably different from C. australiensis (s.s.): the former with flabellate, flabellate-digitate, digitate or rarely subspherical growth forms, bright red-orange live coloration, prominent oscules, with or without subectosomal canals (astrorhizae) or small fistules; the latter predominantly lobate, club-shaped or bulbous, pale orange alive, and oscules without ornamentation (as far as is known). Similarly, there are other cryptic morphological differences seen in preserved material that distinguish the two species. Clathria rugosa has auxiliary spicules with blunt or rounded points, and terminal spines are perched only on the extremities of these 'points', whereas in C. australiensis there is a prominent terminal spine/point, and spines occur at least part of the way along the shaft of the spicule. In this feature C. rugosa resembles most closely C. major. Other New Caledonian Poecilosclerids 1243 differences are that in C. australiensis acanthostyles are pointed and tapering, toxas are very thin, strongly curved at the centre and not abundant, and spicule dimensions vary slightly (Table 4). Clathria rugosa is also similar to another species, described below from the Chesterfield Islands (approximately midway between the Great Barrier Reef and New Caledonia), C. bulbosa, sp. nov. (assigned to the subgenus Clathria in lacking detritus in the skeleton), in its skeletal structure and spicule geometry, although there are several important but cryptic differences between the two species which indicate that the two are not conspecific (described below and Table 4). Etymology The name rugosa refers to the roughened, rugose texture. Clathria (Clathriopsamma)litos, sp. nov. (Figs 9-10) Material Examined Holotype. Q M G301269 (ORSTOM R190): stn 109, Baie des Citrons, Noumea, New Caledonia, 22"18.3'S.,l66O25.8'E., 15 m depth, 17.v.1977, coll. P. Laboute, SCUBA. Description Colour. Pale orange-brown alive (Munsell 10R 7/8), beige in ethanol. Shape. Clump of branching, cylindrical digits (clump 240 mm wide, 115 mm maximum height, 90 mm maximum thickness; individual digits 25-65 mm long, 8-16 mm diameter), each at least partially fused to adjacent digits, attached to a common base, forming branching, non-anastomosing growth form. Digits more-or-less regularly cylindrical, usually bifurcating at least once, and often expanded at apical end, with terminal oscule. Surface. Highly porous, both when alive and when preserved, with wide fibre meshes interconnected on surface by ectosomal membrane stretched across adjacent fibres, usually detached in preserved state, leaving an insubstantial, wide-meshed, honeycombed-reticulate appearance. Large oscules on apex of each digit, 6-13 mm diameter, surrounded by membraneous lip. Each oscule with single, prominent (but non-asterose) subectosomal canals (astrorhizae) running longitudinally from oscule to fused portion of digit. In life subectosomal canals covered by membraneous dermis, collapsing after preservation, leaving large longitudinal furrow on lateral side of each digit, up to 4 mm wide, 5 rnm deep. Surface shaggy, microconulose, with fibre endings protruding through ectosomal membrane. Texture very soft, fragile, insubstantial, easily torn, slightly arenaceous. Ectosome. Membraneous, with tips of primary spongin fibres protruding for up to 300 pm through ectosomal membrane, 300-700 pm apart. No spicules associated with ectosomal skeleton, but moderate quantities of detritus, mostly sand grains, scattered on both exterior and interior of peripheral spongin fibres. Choanosome. Skeletal structure irregularly reticulate. Spicule skeleton absent (except for microscleres), but spongin fibre system well developed, differentiated into primary and secondary fibres. Primary fibres predominantly ascending, 50-70 pm diameter, interconnected by thinner secondary, mostly transverse fibres, 15-35 pm diameter. All fibres lightly laminated but without central pith, and sparse detritus also incorporated into some primary fibres (secondaries completely clear). Fibre meshes cavernous, oval or elongate, variable in size, 130-550 pm diameter. Heavy collagen dispersed within fibre meshes, pigmented light brown, granular, containing sparse microscleres and abundant detritus (sand and foreign spicules). Choanocyte chambers oval, 20-40 pm diameter. Megascleres. Absent. Microscleres. Arcuate isochelae, large, thick, with well developed alae. Length 16(19.9)-24 pm. Toxas uncommon, long, thick, slender, with generous central curvature, long straight (unreflexed arms). Length 114-(164.7)-196 pm, width 1.5-(2.1)-2.5 pm. 1244 J. N. A. Hooper and C. Ltvi Distribution Known only from the Noumea region of the lagoon, New Caledonia, coral rubble substrate, 15 m depth. Remarks This species is included with Clathria (Clathriopsamma) on a speculative basis, since it lacks structural megascleres that would otherwise confirm its affinities. It is assigned to the Microcionidae in having toxa and chelae microscleres [and resemblance in its digitate growth form, surface sculpturing and reticulate spongin fibre skeleton to other microcionids, such as Echinochalina, Clathria (Zsociella)],and it is presently included in the species-group Clathriopsamma in having detritus incorporated into spongin fibres (Fig. 10D-E). The 'keratose'4ke nature of this species makes its inclusion with the Microcionidae obscure; that prompted a more thorough than usual investigation of whether or not the recorded isochelae and toxa microscleres were native to the sponge or acquired. Examination of many fragments, taken from several different places on the sponge, consistently turned up these microscleres, albeit they were never more than 'uncommon', supporting the present placement with Poecilosclerida. Fig. 9. Clathria (Clathriopsamma) litos, sp. nov.: A , holotype QM G301269, toxas; B, arcuate isochelae; C , section through peripheral skeleton. New Caledonian Poecilosclerids 1245 Thus, the affinities of Clathria litos must be inferred by the geometry of these isochelae microscleres, having arcuate isochelae instead of the usual palmate sort found in the majority of microcionids. In fact, several species-groups within the Microcionidae (see below), and throughout Poecilosclerida in general (e.g. Ectyodoryx in the Myxillidae), have been created predominantly or exclusively on this basis. Nominal microcionid genera, defined by possession of arcuate isochelae, include (with the type material of the type species indicated, seen by the senior author): Anaata de Laubenfels [type species Aaata spongigartina de Laubenfels (USNM 21428)], Anthoarcuata Bakus [type species A. graceae Bakus (USNM 36284)], Paradoryx Hallmann [type species Clathria dura Whitelegge (AM G3046) from N.S.W. and Tas.], Qasimella Thomas [no material seen], Quizciona de Laubenfels [type species Microciona heterospiculata Brondsted (BMNH 19Ol.l2.26.13)], and Wetmoreus de Laubenfels [type species Microciona novaezealandica Brondsted (no material seen)]. Many other Australian microcionid species are also positively known to have arcuate isochelae, including two species of Dendrocia Hallmann [D. myxilloides (Dendy) (NMV G2376), and D. pyramida (Lendenfeld) (AM GgON), both from south-east Australia], and a number of species of Clathria [C. scabida (Carter) (BMNH 1887.7.11.9) and C. echinonematissima (Carter) (no extant type material), both from Vic.; Fig. 10. Clathria (Clathriopsamrna) litos, sp. nov.: A, holotype QM G301269 (ORSTOM R190); B, holotype in situ (photo P. Laboute); C, fibre structure (scale=200 pm); D, SEM of fibres and debris incorporated (magnified 200x); E, SEM of skeletal structure (magnified 6 0 ~ ) . 1246 J. N. A. Hooper and C. L6vi C. grisea (Hentschel) (ZMB 4435) from the Houtman-Abrolhos Islands, W.A.; C. curvichela (Hallmann) (AM Z59) from the Gulf St Vincent, S.A., C. elegantula Ridley and Dendy (BMNH 1887.5.2.91) from south-east and south-west Australia, C. oxyphila (Hallmann) (AM Z51) from Tas. and S.A.]. By comparison, other obvious sister-species have palmate isochelae but do not differ substantially in their other morphological and/or skeletal characters, indicating that they also clearly belong to these genera [e.g. an undescribed species of Dendrocia from the Houtman-Abrolhos Islands, W.A. (NTM Z3218), and most species assigned to the genus Clathria]. This material cited above has been comprehensively re-examined and it is confirmed that all nominal taxa are undeniably microcionids. Consequently, the importance of microsclere geometry at supraspecific levels of classification must be questioned. Indeed, Hooper (1991a) indicated that the existing generic classification, which was partially based on chelae geometry (after de Laubenfels 1936), and reiterated most recently by Bergquist and Fromont (1988), requires substantial revision because of high levels of homoplasy due to these and other features (the characters reappearing in many places, inferring that they have evolved independently many times). The present interpretation is that the distinction between palmate versus arcuate isochelae is certainly not one of priority at the family level of classification (e.g. Microcionidae versus Myxillidae; Burton 1959); and, furthermore, the differentiation of genera on the basis of this feature must also be questioned [encompassing many of de Laubenfels' (1936) genera]. These questions, and the high levels of homoplasy in the classification resulting from their continuous use, have already been posed by Hooper (1991a) and this topic must now be investigated more seriously (see also comments noted above for Myxillidae and Acarnus). In any case, apart from the possession of arcuate isochelae, Clathria litos shows no close similarities to any of the above mentioned species. In external morphology it resembles to some extent the tropical Australian Clathria (Isociella) eccentrica (Burton), and also C. (C.) menoui, sp. nov. (see below), but this similarity probably merely reflects the porous, reticulate nature of the surface skeleton. The precise affinities of this species are uncertain. Etymology From the Greek litos (plain, simple), referring to the depauperate, virtually vestigial nature of the spicule skeleton in this species. Clathria (Thalysias) vulpina (Lamarck) (Figs 11-12, Table 6) Restricted synonymy Spongia vulpina Lamarck, 1814: 449. Halichondria frondifera Bowerbank, 1875: 288-89. Clathria diche1a.-Vacelet et al., 1976: 71, pl. 3, fig. b. Clathria corallitincta Dendy, 1889: 85, pl. 4, fig. 8. Rhaphidophlus seriatus Thiele, 1899: 14, pl. 1, fig. 6, pl. 5, fig. 7. Clathria reinwardti var. palmata. -Ridley, 1884: 447. Material Examined Holotype. MNHN DT639: ? Australia, Peron and Lesueur collection (dry). New Caledonian material. QM G300279 (fragment NTM 23882): Baie des Citrons, off Noumea, 22°18~0'S.,166025.0'E., 3 m depth, 25.ix.1990, coll. M. Kelly-Borges, snorkel, fringing coral reef; QM G300692 (fragment NTM 23878): stn 106, N. entrance, RBcif des Cinq Milles, SW. New Caledonia lagoon, 22°293'S.,166044.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA; ORSTOM R183: stn 124, Ilbt Maitre, 22°20.1'S.,166025.1'E., 20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM R264, R471: stn 110, SE. Ilbt Redika, 22O3l. 11S.,166"36.0'E., 19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R588: stn 147, mid-channel between Ton Du (Tiari) and Ilbt Ie, 22°121'S.,166016~5'E., 24 m depth, 8.ix.1976, coll. P. Laboute, SCUBA; ORSTOM R275: stn 113, L'Cpave du 'Bonhomme', Grand Rtcif de Mbhre, 22'21 ~O'S.,166°14.0'E., 25 m depth, 21.vi.1976, coll. P. Laboute, SCUBA. New Caledonian Poecilosclerids 1247 Comparative material. Sri Lanka: BMNH 1889.1.21.17: Gulf of Manaar, Ceylon, 80501N., 79"401E., coll. E. Thurston. Micronesia: USNM 23149: Agana Bay, NW. coast of Guam, 14"N., 154OE., 2 m depth, 22.ix.1949, coll. unknown; ZMA POR769: Siboga stn 257. Philippines: USNM 21256, 21257: precise locality unknown. Indonesia: BMNH 1877.5.21.1351-2: Gaspar Straits, off Belitung I., 3°10'S.,107015'E., coll, unknown; SMF 1699 (fragment MNHN LBIM DCL2352): Straits of Dobo, Aru I., 6°S.,134050'E., 40 m depth, 20.iii.1908, coll. H. Merton, dredge; SMF 1673 (fragment MNHN LBIM DCL2230): same locality, 16 m depth, 20.iii.1908; NMB 16, 17 (fragments BMNH 1908.9.24.165-166, ZMB 2897): Kema, off Minahassa, Celebes (Sulawesi), 2°50'S.,123030'E., 30 m depth, 1895, coll. P. and F. Sarasin, dredge. Darwin region, N.T.: NTM 2974, 978: Dudley Pt, East P t Marine Reserve, 12'24.5-25'S.,13O048-48+4'E., 10-12 m depth, 13.ix.1982, coll. J. N. A. Hooper, SCUBA; NTM 21092, 1101: same locality, 6-7 m depth, 22.xii.1982; NTM 22691: same locality, 9-12 m depth, 3.iv.1986; NTM 23194: same locality, 9 m depth, 16.ix.1987; NTM 21976, 1981: Weed Reef, 12°29.2'S.,130037.6-47. l1E., 8 m depth, 1l.v.1984, coll. J. N. A. Hooper, SCUBA; NTM 22097: same locality, 23.viii.1984; NTM 22188: same locality, 16.xi.1984. NTM 2437: Lee Pt Reef, 12"19~2'S.,130"53~1'E.,intertidal, 13.xii.1981, coll. J. N. A. Hooper; NTM 2782: Angler's Reef, Lee Pt, 12°18.7'S.,130052.0'E., 5-6 m depth, 19.vii. 1982, coll. P. N. Alderslade, SCUBA; NTM 22101, 2103: Fish Reef, Bynoe Harbour, 12"26~2'S.,130°26.2'E., 9-10 m depth, 5.ix.1984, coll. J. N. A. Hooper, SCUBA. Cobourg Peninsula region N.T.: NTM 252, 54, 65: Coral Bay, 6 m depth, Port Essington, Cobourg Pen. Marine Natl Pk, 11'09.4-11~5'S.,132"02~0-04~0'E., 17.x.1981, coll. J. N. A. Hooper, SCUBA; NTM 288: same locality, 4 m depth, 18.x.1981; NTM 294: same locality, 5 m depth, 19.x.1981; NTM 2442: same locality, 4-5 m depth, 18.x.1981; NTM 21354, 1375: same locality, 4-5 m depth, 16.v.1983; NTM 22515: same locality, 2-3 m depth, l 5 . i ~1985; . NTM 22520: Orontes Reef, mouth of Port Essington, 11°03 .6'S.,l32"05.4'E., 18-20 m depth, 16.ix.1985, coll. J. N. A. Hooper, SCUBA; NTM 2626: Unnamed Shoal, N. of Melville I., Arafura Sea, 11°36.2'S.,129052.0'E., 47 m depth, 22.v.1982, coll. R. Lockyer, SCUBA; NTM 2923: Cootamundra Shoals, N. of Melville I., Arafura Sea, 10°50.2'S.,129"13 .2'E., 36 m depth, 7.v.1982, coll. R. Lockyer, SCUBA. Northwest shelf, W.A.: NTM 21222: W. of Port Hedland, 19"28.5'S., 118"55.3'E., 40 m depth, 26.iv.1983, coll. J. N. A. Hooper, CSIRO RV 'Soela', beam trawl; NTMl810: same locality, 19°26~9'S.,l18054.2'E., 50 m depth, 30.viii.1983, coll. T. Ward, CSIRO RV 'Soela', beam trawl; NTM 21421: NE. of Port Hedland, 19°1.0'S.,119025.0'E., 80 m depth, 19.iv.1983, coll. R. S. Williams, pair trawl; NCI Q66C-1557-5 (fragment NTM 23401): 0.5 km E. of Hermite I., Monte Bello Is, 20°27.6'S.,115"32.98'E.,3 m depth, 29.viii.1988, coll. D. Low Choy and NCI, 50 m depth, SCUBA; NTM 23043: N. of Amphinome Shoals, 19°19~7-23.3'S.,119008.8-12.2'E., 19.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', beam trawl. Mid-west coast, W.A.: NTM 22968: W. of Carnarvon, 24°55.6-56.5'S.,112050.8-53.5'E.,80-85 m depth, 14.vii. 1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', beam trawl; NTM 23373: 4 km N. of Barrow I., 20°38.8'S.,115028.8'E., 22 m depth, 26.viii.1988, coll. D. Low Choy and NCI, SCUBA; NTM 23397: 3 km W. of Hermite I., Small Is group, Monte Bello Is, 20'27. lfS.,115"34.2'E., 6 m depth, 29.viii.1988; NCI Q66C-1557-5 (fragment NTM 23401): 1 km E. of Hermite I., Small Is group, Monte Bello Is, 20°27.6'S.,115"32.8'E., 10 m depth, 29.viii.1988; NTM 23446: 2 km N. of Bessieres I., Anchor Is, Exmouth Gulf, 21°30~6'S.,114045.4'E., 17 m depth, 23.viii.1988. Queensland: BMNH 1881.10.21.288: Thursday I., Torres Strait, 10°35'S.,142013'E., 8-10 m depth, coll. HMS 'Alert'; Bird I., Torres Strait, Qld, 11°42'S.,143"05'E., coll. HMS 'Alert'; QM GL708: Moody Reef, Home Is, Cape York, 12°05.6'S.,143014.5'E., 29.7 m depth, 25.ix.1979, coll. QFS, trawl; QM GL863, 1986, 2755: Green I., GBR, 16°46'S.,146058'E,, Feb. 1979-Oct. 1980, coll. L. Cannon and B. Goeden, trawl; QM GL705: Osterland Reef, SE. of Cooktown, 15°36.2'S.,145026.8'E., 24 m depth, 10.ix.1979, coll. QFS, trawl; QM GL710: Noggin Passage, Frankland Is, SE. of Cairns, 17"13~5'S.,146"17~0'E., 37.8 m depth, lO.x.1979, coll. QFS, trawl; NTM 22859: Low Isles, N. of Cairns, 16°23'S.,145034'E., 0.3 m depth, 18.x.1986, coll. A. Ayling. Description Colour. Vivid red-orange alive (Munsell 5R 4/10), beige in ethanol. Shape. Numerous folded and rejoined lamellae together producing elongate-tubular, tubulo-digitate, flabellate or sometimes simply subspherical growth forms, always with cavernous reticulate construction; size varies from small (70 mm diameter) to massive lobes (up to 300 mm diameter). Surface. Many rounded conules, small digits, or spiky projections on ridges and margins of surface, with area between conules highly excavated to produce cavernous interior; large oscules (up to 6 mm diameter) on apex of surface projections or on margins of branches, 1248 J. N. A. Hooper and C. LCvi usually raised above surface and with a membraneous lip. Surface fleshy alive, harder when preserved, not noticeably porous. Ectosome. Variably developed ectosomal skeleton, ranging from typical Thalysias condition consisting of erect brushes of both ectosomal and subectosomal auxiliary styles (mainly on surface conules), to Clathria condition where paratangential or tangential bundles lie above plumose subectosomal skeleton (mainly between surface conules). Subectosomal region usually cavernous, with tracts of choanosomal styles becoming plumose towards the peripheral skeleton, but usually not protruding through surface. Choanosome. Skeletal architecture regularly reticulate, with well-developed spongin fibres forming more-or-less square meshes; fibres differentiated into primary, multispicular, predominantly ascending fibres, where spicules occupy about half of fibre diameter, and secondary, uni- or paucispicular, mostly transverse connecting fibres; fibres cored by both choanosomal principal styles and long, slender subectosomal styles, although the former are most common; fibres relatively heavily echinated by acanthostyles. Mesohyl with light collagen, relatively few free megascleres, and only rare microscleres. Fig. 11. Clathria (Thalysias)vulpina (Lamarck):A, specimen OM G300692, choanosomal principal styles; B, subectosomal auxiliary subtylostyles; C, ectosomal auxiliary styles; D, echinating acanthostyles; E, larger toxas; F, smaller toxas; G , section through peripheral skeleton. New Caledonian Poecilosclerids 1249 Megascleres (refer to Table 6 for dimensions). Choanosomal principal styles, entirely smooth, straight or slightly curved near base, with rounded or only faintly subtylote bases, tapering to sharp points; similar in morphology to subectosomal auxiliary styles, although thicker and shorter; moderately common. Subectosomal auxiliary styles long, thin, straight, rounded smooth bases, rarely subtylote, rarely microspined, tapering to sharp points; abundant. Ectosomal auxiliary styles short, thin, straight, rounded smooth bases or slightly subtylote, rarely microspined, tapering to sharp points; less common. Echinating acanthostyles short, thin, slightly subtylote bases, sharp points, spined mostly on basal and distal ends, with a more-or-less spine-free area below basal swelling; abundant. Fig. 12. Clathria (Thalysias) vulpina (Lamarck): A, specimen QM G300692 in situ (photo P . Laboute); B, SEM of peripheral skeleton (magnified 80 x); C, SEM of fibre characteristics (magnified 600x). Table 6. Comparison in spicule dimensions between Clathria vulpina from New Caledonia and populations from northern Australia Measurements are given in micrometres. Refer to text for sources of information Spicule type Dimension New Caledonia Northern Australia Choanosomal principal styles L W L 140-(156.8)-195 5-(6.2)-8 142-(205.8)-244 2-(3.1)-5 79-(97.7)-128 2-(2.6)-4 37-(51.1)-59 2 5-(3.5)-6 18-(79.3)-194 0.5-(0.6)-0.8 Absent Absent 113-(184.4-253 4-(11.2)-19 124-(221.4)-3 11 1-(4.7)-10 64-(99.1)-161 1-(3.3)-6.9 52-(73.1)-95 1-(6.6)-12 7-(97.9)-200 0.5-(1.2)-3.2 7-(12.4)-18 1-(3.8)-7 Subectosomal auxiliary styles W Ectosomal auxiliary styles Echinating acanthostyles Toxas Isochelae L W L W L W LI LII 1250 J. N. A. Hooper and C. L h i Microscleres. Isochelae absent. Toxas always thin, hair-like, divided into 2 forms, the smaller with central curvature and slightly reflexed arms, the larger with small central bend and straight arms; both forms uncommon. Distribution Widely distributed throughout tropical and subtropical waters of the Indo-Pacific, from Mozambique, Madagascar, Amirante Is, Seychelle Is, Aldabra Is, Red Sea, west coast of India, Gulf of Manaar, Sri Lanka, Mergui Archipelago, Andaman Sea, Straits of Malacca, throughout the Indo-Malay archipelago, Philippines, Vietnam, Japan, Guam, Papua New Guinea, west, north, and eastern Australia and New Caledonia; intertidal to 80 m depth, coral reef, coral rubble, Halimeda beds, beach rock (Hooper, unpublished data). Remarks This species is now known from collections made throughout the Indo-Pacific region: Madagascar, Sri Lanka, Vietnam, Philippines, Guam, Papua New Guinea, Indonesia, Western Australia (Shark Bay to Exmouth Gulf), northern Australia (Darwin region to the Wessel Islands), and eastern Australia (Cape York to Cairns). The New Caledonia population described here is a new locality record and also the most easterly record for the species so far recorded. Not surprisingly, therefore, specimens were found to vary in a number of features from the Indian Ocean-Australasian populations. However, these New Caledonian specimens are still easily recognisable as belonging to C. vulpina, and this recorded variability is not considered to warrant the creation of a new taxon. The New Caledonia population is consistently brighter red in life than other populations (which are typically mauve-red in shallow water, or drab brown-beige in deeper water material). The bases of subectosomal and ectosomal auxiliary styles are predominantly smooth (v. predominantly spined), isochelae appear to be entirely absent (v. isochelae of two size categories present, and in only 3% of material were they rare or absent), toxas are invariably thin, hair-like, never more than 0.8 pm diameter [v. toxa thickness varying from very thin to relatively thick (0.5-3.2 pm diameter)]; ectosomal skeleton is thin, paratangential, relatively poorly developed (v. more substantial development of ectosome, although specimens range from the bushy, erect condition typical of Thalysias, to paratangential as described above), and all categories of spicules are marginally smaller in the New Caledonian specimens (Table 6 ) . Detailed analysis of most morphological characters from large samples of the species (Hooper, unpublished data), in an attempt to document its intra-specific variability, failed to find any substantial, cryptic differences between widely dispersed populations, suggesting that C. vulpina is truly a circum-Indo-Pacific species. Clathria (Thalysias) flabellifera, sp. nov. (Figs 13-14, Table 7) Material Examined Holotype. QM G300693 (ORSTOM R1416; fragment NTM 23884): stn 106, N. entrance, RCcif des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 3O.i~.1976, coll. P . Laboute, SCUBA. Other New Caledonian material. QM G301271 (ORSTOM 'cfR1416'): stn 456, Lagoon pool, E. coast Iles des Pins, 22°35.1'S.,167032.0'E., 38 m depth, 28.xi.1987, coll. J.-L. Menou, SCUBA; ORSTOM 'cfRl416': stn 457, E. coast Ile des Pins, 22"36.0'S.,167"33.O'E., 15 m depth, 28.xi.1987, coll. G. Bargibant, SCUBA. Description Colour. Orange-red alive (Munsell 5R 5/10), pale cream in ethanol. Shape. Flabelliform, flattened in 1 plane, produced by fusion of adjacent, flattened branches, but branch fusion incomplete producing large gaps on surface of fan; fan 170 mm high, 230 mm wide, up to 8 mm maximum breadth, with uneven, digitate margins. New Caledonian Poecilosclerids 1251 Surface. Mostly even, slightly folded, especially towards digitate margins; without conules or other processes; prominent oscules visible in live specimen, approximately 2 mm diameter, each surrounded by white subectosomal canals (astrorhizae) converging on oscules in all directions, but oscules not visible optically in preserved material; surface microscopically porous, with 'dusty' coloured surface in preserved state. Ectosome. Variable development of ectosomal skeleton, ranging from specialised Thalysias condition with erect brushes of intermingled ectosomal and subectosomal auxiliary spicules forming continuous palisade, to Clathria condition with sparse tangential or paratangential tracts of auxiliary spicules; subectosomal region slightly cavernous, with relatively large mesh sizes (200-350 pm maximum diameter); tips of choanosomal principal megascleres may protrude through ectosome, but usually masked by 'true' ectosomal skeleton. Choanosome. Skeletal architecture regularly reticulate, composed of long, primary, ascending spongin fibres (70-150 pm diameter), and short, thinner, secondary, transverse fibres (40-100 pm), together producing elongate-rectangular or square meshes near periphery, more compact oval meshes towards axis (150-200 pm diameter); axial fibres cored by multispicular tracts of choanosomal principal spicules occupying most of fibre diameter; secondary interconnecting fibres paucispicular; all fibres in peripheral skeleton cored by uni- or paucispicular tracts of spicules, forming plumose brushes at surface; echinating acanthostyles abundant throughout skeleton; collagen light, choanocyte chambers oval (40-90 pm diameter), usually lined by abundant isochelae and toxas. Megascleres (refer to Table 7 for dimensions). Choanosomal principal styles relatively short, ranging from narrow to robust, usually straight, slightly subtylote or with rounded, smooth bases, tapering to fusiform points. Subectosomal auxiliary styles long, slender, straight, with rounded or slightly subtylote smooth bases, tapering to sharp points. Fig. 13. Clathria (Thalysias)jlabeNifera,sp. nov.: A, holotype QM G300693, choanosomal principal styles; B, subectosomal auxiliary styles; C, ectosomal auxiliary styles; D, echinating acanthostyles; E, smaller toxas; F, larger toxas; G, palmate isochelae; H, section through peripheral skeleton. J. N. A. Hooper and C. L6vi 1252 Ectosomal auxiliary styles short, slender, straight, with rounded or slightly subtylote bases, usually smooth, sometimes microspined, tapering to sharp points. Echinating acanthostyles relatively robust, subtylote, sharply pointed, with small spines located mainly on bases and points of spicules. Fig. 14. Clathria (Thalysias) Jabellfera, sp. nov.: A , holotype QM G300693 (ORSTOM R1416) in situ (photo J.-L. Menou); B, SEM of axial skeleton (magnified 1 0 0 ~ ) C, ; SEM of fibre characteristics (magnified 500 x ). Table 7. Comparison in spicule dimensions between the New Caledonian Clathriaflabellifera, sp. nov., and the related Western Australian species Clathria cancellaria Measurements are given in micrometres Spicule type - - Dimension - Choanosomal principal styles Subectosomal auxiliary styles Ectosomal auxiliary styles Echinating acanthostyles Toxas I (short, thick, recurved) Toxas I1 (long, thin, straight) Isochelae I Isochelae I1 A C. Jabellifera (New Caledonia) - C. cancellariaA (Western Australia) - 139-(151.4-163 6-(7.6)-9 209-(253.8)-293 3-(4.4-6 98-(131.7)-179 2-(2.3-4 49-(66.7)-84 3-(5.1)-7 13-(33.4-48 1.O-(I.7)-2.5 49-(124.8)-204 O.5-(0.8)-1 .O 11-(13.1)-15 Absent 161-(185.5)-218 6-(12.6)-16 135-(218.8)-286 3-(6.5)-8.5 86-(97. 6)-142 2-(3.3)-5 62-(79.8)-92.5 3-(8.6)-11 .5 Absent Holotype MNHN DT528 and specimen NTM 21249 from the Northwest Shelf, W.A. New Caledonian Poecilosclerids 1253 Microscleres. Palmate isochelae simple shape. Two categories of toxas present: (I) most common short, relatively thick, large central curve, reflexed arms; (11) long, hair-like, with small central curve, straight arms. Distribution Known so far only from the south-west coast of the Noumea lagoon, 8-38 m depth, growing on coral and coral rubble substrates. Remarks Clathria (Thalysias)j7abellifera is very closely related to (and possibly may even eventually be merged with) C. (T.) cancellaria (Lamarck, 1814). Both species possess several similar features, indicating that they are at least sibling species (i.e. similar skeletal architecture, comparable spicule geometry and spicule size in most cases, and nearly identical growth form, surface features and live coloration). Clathria cancellaria was originally collected from an unknown locality thought to be on the south-west coast of Australia-the type material of Lamarck's (1814) (MNHN DT528) has recently been re-examined-and its Western Australian distribution was also recently confirmed by the discovery of additional fresh material from the Houtman-Abrolhos Islands and Northwest Shelf of Western Australia (Hooper, unpublished data). There are no known populations of either species living in between these widely separated New Caledonian and Western Australian populations, a fact that has been established with a reasonable degree of confidence from recent widespread collecting efforts in tropical Australasia. Consideration that these two disjunct populations might be recent isolates of a single species are rejected, given that there are several major, albeit relatively cryptic differences between the two groups. These differences are considered to justify the erection of a new taxon for the New Caledonian population, being relatively more substantial than allowing us to recognise it simply as a geographical 'morph' of C. cancellaria (as done above for C. vulpina, for example). Clathria flabellifera has two different sorts of toxa microscleres (neither of which resemble the single toxon category found in C . cancellaria), one size of isochelae microsclere (the latter species has two), acanthostyle spines are small and found mainly towards the extremities of the spicule (whereas those on the latter species are much larger, the spicules more robust, and spines more evenly dispersed over the spicule), with the exception of a few smaller ectosomal megascleres most auxiliary spicules are completely smooth (whereas C. cancellaria has basal spines on most of the smaller and larger auxiliary spicules), and most spicules are substantially thinner in the New Caledonian population (Table 7). Of course, it is possible that these differences may be trivialised in future re-evaluations of the Microcionidae, but on the basis of morphological criteria currently accepted for this particular group (e.g. Hooper 1991a), and for the Poecilosclerida in general (e.g. van Soest 1984a; Bergquist and Fromont 1988), and corroborated by chemotaxonomic evidence (e.g. Hooper et al. 1990; Hooper et al. 1992), there is certainly sufficient morphological divergence to recognise these two sibling species as distinct. Etymology Named for its fan-shaped growth form. Clathria (Thalysias) corneolia, sp. nov. (Figs 15-16, Table 8) Material Examined Holotype. QM G300691 (ORSTOM R527; fragment NTM 23877): stn 106, N. entrance, RCcif des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA. Other New Caledonian material. ORSTOM R340: stn 114, NE. IlBt TarCti, Banc Gail, 22022.11S., 166"39.0rE., 35 m depth, 29.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R526: stn 147, mid24 m depth, 8.ix.1976, coll. channel between Ton Du (TiarC) and Ilat Ie, 22°12~1'S.,166016~5'E., 1254 J . N. A. Hooper and C. LCvi P. Laboute, SCUBA; ORSTOM R475: stn 110, SE. Ilbt RCdika, 22°31~1'S.,166036~0'E., 19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA. Description Colour. Pale red-pink alive (Munsell 2.5R 8/4), pale pink-brown in ethanol. Shape. Clumped, branching growth form, with bulbous subcylindrical branches or thin arborescent branches, attached to a common base, entirely free or partially fused with adjacent branches, and with one or more points of attachment to substrate. Growth forms range from 70 to 190 mm high; branches bulbous-cylindrical alive up to 45 mm diameter, contracted and slightly flattened when preserved, 5-12 mm diameter, usually bifurcated and reticulated, tapering to rounded tips. Fig. 15. Clathria (Thalysias) corneolia, sp. nov.: A , holotype QM G300691, principal styles (echinating fibres); B, subectosomal auxiliary styles; C,ectosomal auxiliary styles; D, echinating acanthostyles; E, toxas; F, two sizes palmate isochelae; G , section through peripheral skeleton. New Caledonian Poecilosclerids 1255 Surface. Optically even, without any noticeable protrusions; surface with membraneous ectosome stretched between branches, most clearly seen in live specimens, partially collapsing upon preservation, but still retaining prominent subectosomal canals (astrorhizae) running along lateral margins of branches. Large oscules, up to 25 mm diameter, on apex of branches, surrounded by prominent membraneous lip. Ectosome. Specialised ectosomal skeleton present, consisting of well-developed, discrete plumose bundles of ectosomal styles, perched on peripheral spongin fibres; ectosomal spicule bundles not always associated with ascending fibres or spicule tracts that arise from sub- Fig. 16. Clathria (Thalysias) corneolia, sp. nov.: A , specimen ORSTOM R340 in situ (photo P. Laboute); B, holotype QM (3300691 (ORSTOM R527); C , SEM of fibre characteristics (magnified 300 x); D, SEM of axial skeleton (magnified 8 0 ~ ) . Table 8. Comparison in spicule dimensions between the New Caledonian Clathria corneolia, sp. nov., and the related species Clathria cervicornis (Thiele) from Oceania and the Indo-Malay archipelago Measurements are given in micrometres Spicule type Dimension Choanosomal principal styles L Subectosomal auxiliary styles W L W Ectosomal auxiliary styles Echinating acanthostyles Toxas Isochelae I Isochelae I1 L W L W L W L L C . corneolia (New Caledonia) C. cervicornis (holotype SMF 679) Undifferentiated from subectosomal styles 180-(258.9)-304 4-(5.8)-9 112-(128.3)-155 2-(3.3)-5 48-(62.0)-74 3-(5.9-7 12-(35.3)-55 O.5-(0.9)-1.5 10-(12.4)-15 3-(6.9)-9 1256 J. N. A. Hooper and C. Ltvi ectosomal skeleton. Peripheral spongin fibre skeleton well developed, lying directly below ectosomal membrane. Fibres usually terminate as swollen nodes on surface. Choanosome. Subectosomal organic skeleton relatively cavernous, with large, elongate fibre meshes, 160-350 pm diameter; subectosomal mineral skeleton more-or-less plumose, with tracts of subectosomal auxiliary styles, in paucispicular bundles, coring fibres and many also protruding from fibres in plumose array. Choanosomal skeleton irregularly reticulate, with well-developed spongin fibre system divided into primary and secondary components; fibre reticulation in choanosomal core produces elongate meshes, 230-400 pm maximum diameter; primary fibres paucispicular, 100-180 pm diameter, cored by subectosomal auxiliary styles; secondary fibres 50-90 pm diameter, aspicular or cored by uni- or paucispicular tracts of subectosomal styles. Both acanthostyles and principal styles echinate fibres, but these are sparse. Choanocyte chambers oval, 50-85 pm diameter, often paired. Mesohyl reinforced by light collagen, with abundant microscleres and many auxiliary spicules also dispersed between fibres. Megascleres (refer to Table 8 for dimensions). Principal styles (echinating fibres) scarce; straight, sharply pointed or with stepped points (telescoped ends); bases evenly rounded or tapering, hastate, invariably smooth. Intermediate forms between principal styles and acanthostyles also occur. Subectosomal auxiliary styles long, usually straight, sharply pointed, with slightly subtylote, smooth bases. Ectosomal auxiliary styles short, thin, straight, sharply pointed, with slightly subtylote bases. Acanthostyles (echinating fibres) scarce; short, stout, sharply pointed or stepped tips, with evenly rounded or slightly subtylote bases; spines concentrated on bases and mid-way along shaft, sometimes in more-or-less verticillate rows. Microscleres (refer to Table 8 for dimensions). Palmate isochelae scarce, two size categories, both small, the larger with some twisted forms. Toxas common, moderately thin, short or long, usually with large central curvature, with or without straight arms. Distribution Known so far only from the south-west coast of the New Caledonian lagoon, sand, coral rubble, Halimeda beds. Remarks This species is very similar to C. (T.) cervicornis (Thiele, 1903), which is widespread throughout the western Pacific and Indo-Malay archipelago-Marshall Is, Palau Is, Caroline Is, Moluccas, Arafura Sea (Thiele 1903; Brondsted 1934; de Laubenfels 1954; Bergquist 1965). However, there are several cryptic but important morphological differences indicating that two species should be distinguished. In C. cervicornis, principal styles are virtually undifferentiated from subectosomal spicules; most spongin fibres are cored by multi- or paucispicular columns of subectosomal styles, and there is no differentiation of primary or secondary fibre elements; most categories of spicules are substantially longer and more robust (Table 8); and spination of acanthostyles is also quite different. Etymology Named for the bulbous, horny spongin fibres predominant in the surface and choanosoma1 skeletons (L. cornu, horn). Clathria (Thalysias) araiosa, sp. nov. (Figs 17-18, Table 9) Material Examined Holotype. QM G300694 (ORSTOM R1370; fragment NTM 23886): stn 106, N. entrance, Rtcif des Cinq Milles, SW. New Caledonia lagoon, 22"29.3'S.,166"44.4'E., 8 m depth, 30.iv.1976, coll. G. Bargibant, SCUBA. New Caledonian Poecilosclerids Description Colour. Pale red alive (Munsell 5R 6/10), beige in ethanol. Shape. Bulbous, lobate, thickly encrusting, up to 15 mm thick. Surface. Uneven surface composed of numerous fused bulbs, each with apical osculum, up to 4 mm diameter, raised on membraneous lip and with white or pale red subectosomal canals (astrorhizae). Surface also excavated by parasitic barnacles. Ectosome. Dense ectosomal skeleton, composed of a single layer of ectosomal auxiliary styles forming compact brushes, perched on the ends of subectosomal spicule brushes. Choanosome. Hymedesmoid skeletal architecture, with thin layer of spongin lying on coralline substrate; bases of choanosomal principal styles and echinating acanthostyles embedded in basal spongin layer, standing perpendicular to substrate; multispicular brushes of subectosomal auxiliary styles arise from ends of erect choanosomal styles, in 2-3 discrete layers, producing straight or sinuous tracts within mesohyl, and with peripheral layer of subectosomal spicules intermingled with ectosomal spicules to produce the ectosomal skeleton. Choanosome slightly cavernous near substrate, with oval choanocyte chambers, 40-130 pm diameter, and granular collagen permeating mesohyl. Megascleres (refer to Table 9 for dimensions). Choanosomal principal styles short, slender, straight or slightly curved towards base, sharply pointed, with smooth evenly rounded or very slightly subtylote bases. Subectosomal auxiliary styles long, slender or relatively thick, straight, sharply pointed, with smooth, rounded or slightly subtylote bases. Fig. 17. Clathria (Thalysias) araiosa, sp. nov.: A, holotype QM G300694, choanosomal principal styles; B, subectosomal auxiliary styles; C, ectosomal auxiliary styles; D, echinating acanthostyles; E, toxas, F, palmate isochela; G, section through peripheral skeleton. 1258 J. N. A. Hooper and C. Ldvi Ectosomal auxiliary styles only slightly differentiated from choanosomal principal styles, marginally thinner, straighter, with smooth, evenly rounded, non-tylote bases. Echinating acanthostyles relatively long, slender, slightly subtylote bases, sharply pointed, with small spines; most spines on basal end and midway along spicule, with points and areas below basal constriction free of spines. Microscleres (refer to Table 9 for dimensions). Palmate isochelae rare, relatively large, single size class. Toxas small, thin, slightly curved at centre, with straight arms, rarely with reflexed tips. Distribution Known only from the New Caledonia lagoon, dead coral substrate, 8 m depth. Remarks Clathria araiosa is unremarkable in most of its characters, resembling many other encrusting microcionid species [e.g. C. (C.) aceratoobtusa (Carter, 1887) (with representative specimen seen by the authors NTM 23676 from the Gulf of Thailand, the holotype having been destroyed), C. (C.) grisea (Hentschel, 1911) (fragment of the holotype ZMB 4435), C. (T.) aruensis (Hentschel, 1912) (holotype SMF 955), C. (T.) toxifera (Hentschel, 1912) (with representative specimen NTM 22136 from Darwin, N.T., the holotype not yet found in the ZMH)]. Like these species, C.araiosa has a rnicrocionid skeletal structure (Fig. 17G), red pigmentation, and a surface sculptured by subectosomal canals (astrorhizae) (Fig. 18A). It is distinguished from these species only by a combination of characters such as the size, Fig. 18. Clathria (Thalysias) araiosa, sp. nov.: A , holotype QM G300694 (ORSTOM R1370) in situ (photo G. Bargibant); B, SEM of peripheral skeleton (magnified 80 x); C, SEM fibre characteristics (magnified 500 x). New Caledonian Poecilosclerids 1259 Table 9. Comparison in spicule dimensions between Clathria araiosa, sp. nov., and the related species Clathria coralliophila (Thiele) from Indonesia, Papua New Guinea and the Great Barrier Reef Measurements are given in micrometres C. araoisa (New Caledonia) C. coralliophila (lndonesiaA) C. coralliophila (Great Barrier ~ e eB,f C. coralliophila (Papua New ~uinea~) L W 126-(186.6)-267 6-(7.3-10 Choanosomal principal styles 285-(347.5)-408 145-(252.8)-406 12-(13.8)-15 7-(10.5)-13 224-(345.2)-452 10-(12.4)-15 L W 278-(339.2)-365 2.5-(4.6)-6 Subectosomal auxiliary styles 312-(390.4)-488 321-(355.4)-390 5-(7.9)-10 5-(7.3)-11 223-(347.3)-430 5-(7.2)-9 L W 121-(142.3)-183 2.5-(3.2)-4 Ectosomal auxiliary styles 118-(140.9)-174 104-(150.1)-208 2-(3.1)-4.5 2-(3.1)-4 109-(121.5)-138 2-(3.5)-5 L W 50-(61.8)-72 2.5-(4.1)-5 48-(56.3)-7 1 4-(5.2)-6 84-(95.0)- 105 4-(6.3)-8 76-(85.8)-95 5-(6.6)-8 L W 12-(25.3-42 0.5-(1.1)-2 Toxas 18-(76.6)-118 0.5-(0.9)-1.2 68-(140.3)-244 0.5-(1.4)-2 32-(125.5)-211 0.5-(1.2)-2 11-(13.5)-17 10-(12.1)-15 2-(4.4)-8 3-(5.1)-8 Echinating acanthostyles Isochelae I L 15-(16.6)-18 11-(13.1)-16 L Absent 2-(4.6)-8 Isochelae I1 A Lectotype SMF 1784 from Ternate, Moluccas. Redescription of Burton's (1934) specimen from Low Is, BMNH 1930.8.13.107. Specimen NCI Q66C-4518-A (fragment QM G300377) from Motupore I. geometry and ornamentation of spicules. It is closest to C.(T.) coralliophila (Thiele, 1903) from the Moluccas, Papua New Guinea and the northern Great Barrier Reef (lectotype SMF 1784), although C. araiosa differs from that species in the dimensions of many categories of spicules (Table 9), in lacking basal spines on either principal or auxiliary spicules, the pattern of spination on acanthostyles, and in lacking a smaller category of palmate isochelae. Etymology From the Greek araios, thin, porous. Clathria (Thalysias) hirsuta, sp. nov. (Figs 19-20, Table 10) Material Examined Holotype. QM GL2746 (fragment NTM 21551): Cairns region, Qld, 16"56'S.,146"00'E., depth unknown, 1982, coll. QFS (stn 7, Cairns inter-reef survey, trawl). Paratypes. QM GL2750 (fragment NTM 21555), QM GL2754 (fragment NTM 21560): Cairns region, Qld, 16°56'S.,146000'E., depth and date of collection unknown, coll. QFS (stn 7, Cairns inter-reef survey, trawl). 1260 J. N. A. Hooper and C. L h i Other Queensland material. NCI Q66C-0813-X (fragments QM G300603, NTM 23494): N. end of Deloraine I., Whitsunday Is, GBR, 20°09'S.,149004'E., 20 m depth, 15.x.1987, coll. NCI, SCUBA; QM G300081: Shag Rock, N. Stradbroke I., vicinity of Moreton Bay, 27°25.0'S.,153031.4'E., 15 m depth, 5.ii.1992, coll. J. N. A. Hooper and J. Wilkinson, SCUBA. New Caledonian material. QM G301274 (ORSTOM R1548): 'Patate Proner', Ile des Pins, S. lagoon, New Caledonia, 22°31.2'S.,167024.5'E., 30 m depth, 10.x.1991, coll. G. Bargibant, SCUBA. Description of typical material (Queensland waters) Colour. Bright red alive conules (Munsell 2.5R 5/10), paler between conules, dark brown in ethanol. Shape. Small, erect, tubular or flattened-lobate digits, reticulate-honeycombed in construction, 98-165 mm long, 57-68 mm wide, 25-46 mm in diameter, with large digitate, prominently conulose margins; with or without cylindrical basal stalk, up to 9 mm long, 6.5 mm diameter (Fig. 20E). Surface. Highly clathrous, reticulate, Echinoclathria-like surface, consisting of numerous small or large interconnected flattened or lobate branches, up to 15 mm long, with rounded or pointed conulose edges (Fig. 20F). Surface membraneous between conules, and shaggy on conules. Branches forming 'body' of sponge are completely fused although perforated by large cavities, and adjacent branches are joined by a membraneous covering; porous meshes between adjacent branches up to 4.5 mm diameter, excavating sponge entirely. Texture firm, compressible, difficult to tear, slightly arenaceous. Large oscules, 3-5.3 mm diameter, scattered between digitate surface projections; small pores, 0.5-0.8 mm diameter dispersed over surface between conules. Ectosome. Ectosomal skeleton lightly arenaceous, with a sparsely dispersed, thin, tangential or paratangential, uni- to paucispicular skeleton composed of subectosomal auxiliary subtylostyles lying on or just below surface; at tips of conules the surface skeleton is raised to form irregular paratangential tracts which project through the surface. Choanosomal principal styles also pierce surface in places on the ends of conules, occurring individually or in light bundles, but these are relatively infrequent. Choanosomal fibres are immediately subectosomal, with only a small subectosomal region, and the extra-fibre spicule tracts composed of subectosomal auxiliary styles are confined to peripheral skeleton. Choanosome. Choanosomal skeletal architecture irregularly reticulate, with thick, flattened, relatively heavy spongin fibres divided into primary and secondary components. Primary fibres, 58-95 pm diameter, contain multispicular tracts of intermixed choanosomal principal styles and subectosomal auxiliary styles. Primary tracts relatively straight, ascending to the surface (running longitudinally through sections), although near periphery they curve towards surface and terminate in light plumose brushes; secondary fibres relatively short, thick, 15-85 pm diameter, interconnecting primary fibres, with a uni- or paucispicular core of intermingled principal and subectosomal spicules. Secondary fibres run mostly perpendicular to surface, and predominant in deeper regions of choanosome. Echinating acanthostyles moderately heavy on all fibres, particularly on those near periphery. Circular, thick fibre nodes on all major fibre junctions, 60-98 pm diameter, contain a heavier core of choanosomal mpgascleres than in other parts of fibres. Branching of spongin fibre produces variable ovoid or elongate meshes, usually small, sometimes cavernous, 49-210 pm diameter. Few spicules or detritus scattered between fibres. Abundant, unpigmented, granular collagenous spongin in mesohyl, with oval choanocyte chambers, 42-57 pm diameter. Megascleres (refer to Table 10 for dimensions). Choanosomal principal styles thin, fusiform, straight, rounded or slightly subtylote, smooth bases; not well differentiated from auxiliary subtylostyles, although shorter and thicker, with less well-developed basal swelling. Subectosomal auxiliary subtylostyles long, slender, fusiform, straight, evenly cylindrical, rounded or subtylote bases, usually microspined. Acanthostyles straight, prominently subtylote, with light or vestigial spination, mostly aspinose necks. New Caledonian Poecilosclerids 1261 Microscleres (refer to Table 10 for dimensions). Palmate isochelae not common (holotype and paratype GL2750), or common (other material), small, imperfectly divided into two size categories. Toxas very thin, raphidiform, abundant, very slight and gradually rounded central curvature, with straight arms or very slightly reflexed points, rarely sinuous. Description of Atypical Material (New Caledonian waters) Colour. Tips of protruding surface conules red alive (Munsell 5R 5/10), honeycombed portion of sponge pale mauve-red alive (2.5R 8/2), with slightly striated, paler surface sculpturing (subectosomal canals (astrorhizae)) also visible in between surface conules; sponge evenly beige in ethanol. Fig. 19. Clathria (Thalysias) hirsuta, sp. nov.: A , New Caledonian specimen QM G301274, choanosomal principal styles; B, subectosomal auxiliary styles; C, ectosomal auxiliary styles; D, echinating acanthostyles; E, toxas; F, Queensland paratype QM GL2754, choanosomal principal styles; G , subectosomal auxiliary subtylostyles; H, echinating acanthostyle, I, toxas, J, palmate isochelae; K, specimen QM G301274, section through peripheral skeleton. 1262 J. N. A. Hooper and C. Livi Shape. Erect flattened digit, 65 mm long, 40 mm wide, 30 rnm thick, with honeycombed construction, excavated by pores and canals passing completely through sponge, with highly conulose surface (Fig. 200, G). Surface. Numerous close-set, pointed surface conules, 2-4 mm apart, 3-8 mm long, 2-3 mm diameter, usually bifurcate or trifurcate at apex, situated on ends of erect flat, lamellate surface processes, 5-7 mm long, 5-9 mm wide, up to 2 mm thick, which encircle edges of the excavated canals and rejoin to produce the honeycombed surface structure. Texture firm, compressible, difficult to tear. Oscules, up to 5 mm diameter, scattered over surface between microconules. Ectosome. Ectosomal skeleton consists of sparse tangential or paratangential tracts of intermixed ectosomal and subectosomal auxiliary styles, not producing a specialised Thafysias-like skeleton. Collagen in ectosomal region only lightly developed. Choanosome. Skeleton dominated by well-developed, horny spongin fibres, with mineral skeleton greatly reduced in comparison with Queensland material. Spongin fibre skeleton Fig. 20. Clathria (Thalysias) hirsuta, sp. nov.: A , holotype QM GL2746, SEM acanthostyles ; SEM skeletal structure (magnified 1 0 0 0 ~ )B, ; SEM ectosomal spicule brushes (magnified 1 5 0 ~ ) C, (magnified 1 0 0 ~ )D, ; specimen QM (3301274 (ORSTOM R1548) from New Caledonia; E, holotype (QM GL2746) from Cairns region; F, surface of specimen QM G300603 from Great Barrier Reef (photo NCI); G, specimen QM G301274 (ORSTOM R1548) from New Caledonia in situ (photo P. Laboute). New Caledonian Poecilosclerids 1263 differentiated into primary, ascending fibres, 65-110 pm diameter, cored by multispicular tracts of spicules; secondary, transverse fibres shorter and marginally thinner, 80-170 pm long, 20-50 pm diameter, interconnecting ascending fibres, cored by pauci- or unispicular tracts of megascleres. All fibres cored by abundant subectosomal auxiliary styles intermixed with fewer principal choanosomal styles, and echinated by sparse echinating acanthostyles. Coring spicules occupy only small proportion of fibre diameter, usually confined to centre of fibre. Fibre reticulation forms triangular, oval or elongate meshes, more compact in axial region (45-110 pm diameter) than in peripheral skeleton (90-180 pm diameter). Mesohyl with abundant collagen, slightly granular, lightly pigmented, but with few interstitial spicules. Choanocyte chambers large, oval, 25-50 pm diameter. Megascleres (refer to Table 10 for dimensions). Choanosomal principal styles (exclusively in fibres) scarce, very slender, straight, sharply pointed, with smooth evenly rounded bases. Subectosomal auxiliary styles (both coring fibres and in the subectosomal skeleton) long, very slender, straight or rarely slightly curved, with tapering sharp points, and bases nearly anisoxeote, tapering to pointed or small rounded ends. Ectosomal auxiliary styles short, straight, very slender, with evenly rounded bases. Echinating acanthostyles scarce, short, slender, sharply pointed, with vestigial spines and very slight subtylote bases. Microscleres (refer to Table 10 for dimensions). Isochelae absent. Toxas uncommon, raphidiform, with slightly angular or rounded central curvature and straight arms. Table 10. Comparison in spicule dimensions between Queensland and New Caledonian populations of Clathria hirsuta, sp. nov. Measurements are given in micrometres Holotype (QM GL2746) Paratypes (QM GL2750, 2754) Other material Queensland New Caledonia (n = 1 ) (n = 2) Choanosomal principal styles 132-(158.9)-195 140-(158.8)-183 3.5-(6.2)-10 4-(6.0-9 Subectosomal auxiliary styles 183-(237.9-293 141-(202.2)-254 2-(3.6)-5 2-(3.5)-4.5 Ectosomal auxiliary styles Absent Absent Absent Echinating acanthostyles 43-(63.2)-79 3-(5.2)-8 52-(63.8)-75 3-(5.1)-8 Toxas 35-(100.7)-163 0.5-(0.7)-1.0 22-(104.6)-165 0.5-(0.8)-1 .O Isochelae I 3-(4.3-6 3-(3.8)-5 Absent 9-(11.2)-14 Absent Isochelae I1 9-(10.8)-14 1264 J. N. A. Hooper and C . L6vi Distribution Northern, central and southern inshore coastal waters of Queensland, and the New Caledonia lagoon; coral reef, rock and coral rubble substrates, 20-30 m depth. Remarks Two independent descriptions of this species are given in order to highlight the differences between Queensland and New Caledonian populations. Although these differences are not considered here to be sufficient to recognise two distinct, sibling species, many characters seen in New Caledonian material have been modified to a greater or lesser degree, or have been lost completely, from those present in Australian populations. We consider that the Queensland morph is typical of the species, whereas the loss or reduction of characters seen in New Caledonian material, which is obviously secondary, is an apomorphy for the species. Hence, the New Caledonian populations are interpreted as being derived sister-species of the Queensland populations. Nevertheless, both populations are clearly conspecific, as seen by their growth forms (spiky surface, vaguely digitate, reticulate-honeycombed construction), live coloration (darker red conules, paler red or whitish between conules, with subectosomal striations) and skeletal architecture (particularly the predominance of spongin fibres in the skeleton, with well-differentiated primary multispicular and secondary paucispicular fibres), whereas they are contrasted substantially in their spicule geometries (compare Fig. 19A-E, F-J) and spicule sizes (Table 10) to the extent that comparison of these features alone may not be enough to recognise conspecificity. Additional, non-morphological evidence is required to determine the degree to which these two populations have diverged, but for the present they are treated as conspecific. In comparison with New Caledonian material, Queensland specimens have much more robust spicules in all categories, and their relative abundance in the skeleton also differs considerably: choanosomal principal styles are abundant inside fibres; there is only one size class of auxiliary spicule (lacking specialised ectosomal megascleres) which is a large, subtylote auxiliary style; acanthostyles are subtylote and relatively heavily spined; two size classes of isochelae are present; and toxas are similar in thickness and central curvature, but have slightly reflexed points. Clathria hirsuta belongs to a species-group having vaguely similar skeletal characteristics, but are generally morphologically depauperate in comparison with many other Clathria species. To this group belong Clathria (ThaZysias) cactiformis (Lamarck) (lectotype MNHN DT580), C. (T.) clathrata, sensu Topsent (not Schmidt nor Whitelegge) (no material seen), C. (T.) cervicornis (Thiele) (holotype SMF 679), C. (T.) juniperina (Lamarck) (lectotype MNHN DT570), C. (T.) paucispina (Lendenfeld) (lectotype AM G9121a), and the present species. All these species, termed here the 'juniperina' species complex, and distributed amongst the nominal genera Clathria and Thalysias, have choanosomal (coring) megascleres that are barely differentiated from subectosomal auxiliary spicules (and in the present species both are intermingled within the fibres), an irregular, heavy fibre skeleton, and raphidiform toxas, but each species differs in one or more other significant features. In the case of the present species, characteristics that distinguish it from other members of the 'juniperina' species-group are its growth form, the possession of vestigially spined acanthostyles, very thick, relatively flattened fibres, relatively thin megascleres of all categories, two sizes of isochelae, and slightly curved, reflexed raphidiform toxas. Hallmann (1912) contends that some of these species (i.e. those he suggested were related to C. paucispina) are barely different in their gross morphology (i.e. more-or-less tubular, digitate, with a spiky surface), and therefore should be combined, but resolution of this species complex is not possible without much more comparative (preserved) material and corroboration using non-skeletal characters. Whether or not the 'juniperina' species group is a true monophyletic group has yet to be decided, requiring rigorous investigation of the relative importance, and phylogenetic value, of the morphological characters concerned. Etymology The species name refers to the highly conulose, shaggy surface. New Caledonian Poecilosclerids Clathria (Clathria) kylista, sp. nov. (Figs 21-22, Table 11) Material Examined Holotype. QM G300035: Inner Gneering Shoals, off Mooloolaba, SE. Qld, 26°38.5'S.,153009.5'E., 10 m depth, 10.xii.1991, coll. J. N. A. Hooper and S. D. Cook, rock reef. Paratype. QM G300690 (ORSTOM R1338; fragment NTM 23876): stn 106, N. entrance, Rkcif des Cinq Milles, SW. New Caledonia lagoon, 22"29.3'S., 166"44.4'E., 8 m depth, 3O.i~.1976, coll. G. Bargibant, SCUBA. Description Colour. Dark orange to pale orange alive (Munsell 10R 6/10-2.5YR 7/8), dark brown to beige in ethanol. Shape. Growth form ranges from erect, branching, bulbous, digitate tubes, with several tubes attached to a common base, some bifurcating some single and each terminating with a single osculum (paratype), to irregular more-or-less tubular lobes and low digits arising from a bulbous, thickly encrusting base, with oscules scattered over the apical regions of lobate digits (holotype). Specimens range from about 130 mm basal diameter up to encrusting individuals exceeding 320 mm basal diameter, and lobate digits range from 20 mm long, 25 mm diameter, to over 300 mm long, 70 mm diameter. Surface. Prominently conulose, large, bulbous surface tubercles, producing regular (paratype) or irregular (holotype) surface sculpturing. Oscules large, up to 35 mm diameter, surrounded by a prominent membraneous lip. Ectosome. Membraneous, with a sparse, tangential layer of irregularly dispersed subectosomal auxiliary styles and raphidiform spicules, presumably younger forms of these spicules; ectosomal skeleton also contains erect brushes of auxiliary spicules scattered Fig. 21. Clathria (Clathria) kylista, sp. nov.: A , paratype QM G300690, choanosomal principal styles; B, subectosomal auxiliary subtylostyles; C, smooth echinating subtylostyles; D, toxas; E, section through peripheral skeleton. 1266 J. N. A. Hooper and C. L6vi throughout both peripheral and subectosomal regions, generally protruding towards exterior. In peripheral region mesohyl contains moderately light collagen and few small sand particles. Choanosome. Skeletal structure plumoreticulate, with differentiated primary and secondary skeletal lines. Primary fibres long, ascending, 50-70 pm diameter, paucispicular, with never more than 4-5 spicules abreast; secondary fibres shorter, 15-30 pm diameter, transverse, connecting primary fibres, uni- or aspicular. Primary fibres cored predominantly by principal spicules but also with moderate quantities of longer, more slender auxiliary megascleres interdispersed; spicules in primary fibres occupy only small proportion of fibre diameter. Echinating styles moderately common on primary ascending fibres, sparse on secondary connecting fibres, inserted at oblique angles to fibres; this uneven distribution of both coring and echinating spicules gives skeleton an almost plumose appearance. Choanosomal mesohyl contains sparse collagen, fibre meshes widely spaced, 160-350 pm diameter, choanocyte chambers oval, 35-60 pm diameter, and small quantities of sand grains scattered between fibre meshes. Megascleres (refer to Table 1 1 for dimensions). Choanosomal principal styles slender, straight, relatively short, with hastate points and smooth, slightly swollen, subtylote bases. Subectosomal auxiliary styles long, slender, straight, with hastate points and smooth, elongated, swollen subtylote bases. Echinating styles entirely smooth, short, slender, sharply pointed, with prominent basal constriction ('neck'), subtylote base and widest just below basal constriction. Microscleres (refer to Table 11 for dimensions). Isochelae absent. Toxas abundant, moderately long, ranging from slender to raphidiform, with straight arms and prominent central curve. Clathria (Clathria) kylista, sp. nov.: A , paratype QM G300690 (ORSTOM R1338) in situ (photo G. Bargibant); B, SEM of skeletal structure (magnified 80 x); C , SEM fibre characteristics (magnified 500 x). Fig. 22. New Caledonian Poecilosclerids 1267 Distribution Although this species is known only from the Noumea lagoon and Sunshine coast region o f Queensland, it is hypothesised that it also lives on the southern Great Barrier Reef region and some other subtropical western Pacific islands; depth range 8-10 m . Remarks Under previous schemes for the Microcionidae (e.g. de Laubenfels 1936), this species would probably have been referred to Echinoclathria because its echinating spicules are entirely smooth. However, as noted elsewhere (Hooper 1991a), Echinoclathria differs from Clathria not in the presence or absence o f spines on these spicules, but in the structure (or derivation) o f spicules localised to particular parts o f the skeleton: in Echinoclathria, spongin fibres are both cored and echinated by principal spicules (i.e. spicules with identical geometry), whereas in Clathria there are distinct categories o f coring (principal) and echinating megascleres. Clathria kylista is remarkable in its growth form (Fig. 22A), skeletal architecture (Fig. 22B-C) and spiculation (Fig. 21A-D). It superficially resembles a Callyspongia (Haplosclerida) in its tubular digitate morphology, although it lacks the specialised surface sculpturing characteristic o f that genus. Its skeletal structure o f sparsely cored ascending primary fibres and uncored secondary connecting fibres is similar to that o f C. (C.) angulifera Dendy (holotype NMV G2286), which has acanthose echinating spicules and isochelae, and an undescribed species from South Australia (Clathria provisional species number 503) which has acanthose echinators, isochelae and slightly curved toxas. The possession o f entirely smooth echinating styles with basal constrictions, as well as long slender toxas with prominent central curvature is reminiscent o f similar spicules found in C. dubia (Kirkpatrick) (holotype BMNH 1898.12.20.37 from Christmas Island, re-examined). This latter species has a specialised ectosomal skeleton (placing it with Thalysias), a hymedesmoid skeletal structure, and anchorate isochelae. The unusual geometry o f the echinating styles in C. kylista is the strongest apomorphy amongst other species o f Clathria. Only two specimens are so far known for C. kylista, one from Queensland and one from New Caledonia, and both o f these are very similar in their morphology and skeletal characters. Several characters have been contrasted between this material in the description above, and comparisons in spicule dimensions are given in Table 11. Etymology From the Greek kylistos, rolled, alluding to the tubular bulbous growth form peculiar to this species. Table 11. Comparison in spicule dimensions between New Caledonian and Australian specimens of Clathria kylista, sp. nov. Measurements are given in micrometres Spicule type Choanosomal principal styles Subectosomal auxiliary styles Ectosomal auxiliary styles Echinating acanthostyles Toxas Isochelae Dimension New Caledonia (paratype QMG300690) South-east Queensland (holotype QMG300035) L W L W L W L W L W L 61-(81.6)-97 1.6-(2.3-3.6 138-(159.4)-183 0.4-(2.1)-3.5 Absent 64-(84.7)-112 2.5-(2.9)-4.0 143-(168.9)-181 1.0-(2.4)-4.1 Absent 32-(35.5)-39 1.1-(3.1)-5.0 65-(135.6)-266 0.2-(1 .I)-1 -6 Absent 26-(32.8)-40 1 .O-(2.6)-4.0 65-(129.6)-231.4 0.5-(I .3)-2.5 Absent J. N. A. Hooper and C. Ltvi 1268 Clathria (Clathria) bulbosa, sp. nov. (Figs 23-24, Table 4) Material Examined Holotype. QM G300021 (ORSTOM R1350): stn 423, E. rtcif Boody, Chesterfield Is (territory of New Caledonia), 20°58.1'S.,158034.6'E., 42 m depth, 23.vii.1984, coll. G. Bargibant, SCUBA. Description Colour. Bright red alive (Munsell 5R 4/10), pale beige in ethanol. Shape. Thickly encrusting on plate coral (Astreopora sp., Family Acroporidae), bulbous, mushroom-shaped sponge (40 mm high, 60 mm diameter), with short, slightly constricted basal attachment (10 mm long, 25 mm diameter), and two large oscular atria on apex of sponge (10-18 mm diameter). Surface. Few rounded irregular lobate bumps on lateral sides, whereas surface more even on apex; entire surface microscopically porous. Oscular atria surrounded by symmetrical membraneous lip. Large openings of excurrent canals (true oscules, 2-5 mm diameter) visible inside central oscular atrium. Texture soft, compressible, difficult to tear. Ectosome. Ectosomal skeleton with a special category of quasidiactinal auxiliary styles erect or lying tangential to surface, usually not projecting from it, forming sparse bundles or scattered individually on surface; these spicules confined to peripheral skeleton and do not occur deeper within choanosomal region. Light collagenous spongin in mesohyl of peripheral skeleton, with sparsely dispersed microscleres. Fig. 23. Clathria (Clathria) bulbosa, sp. nov.: A, holotype QM G300021, choanosomal auxiliary subtylostyles; B, subectosomal auxiliary quasidiactinal styles; C, echinating acanthostyles; D, toxas; E, palmate isochelae; F, section through peripheral skeleton; G, fibre characteristics. New Caledonian Poecilosclerids 1269 Choanosome. Skeletal architecture reticulate, with a well developed organic spongin fibre system predominant over inorganic spicule components. Fibres differentiated into primary and secondary elements. Primary fibres (45-70 pm diameter) generally ascending, usually cored by plumose paucispicular brushes of auxiliary styles (different from those on ectosome). These spicule brushes not completely confined to axis of fibres, occasionally bundles lie outside, adjacent to primary fibres. Secondary fibres (12-35 pm diameter) usually transverse connecting, mostly uncored or rarely with single auxiliary styles lying in axis of fibre. Both primary and secondary fibres heavily echinated by acanthostyles, heaviest at fibre nodes. Fibre meshes oval to elongate, 70-350 pm diameter, more cavernous in choanosome than in peripheral region. Mesohyl heavily permeated by collagenous spongin containing abundant rnicroscleres. Choanocyte chambers oval (48-60 pm diameter), often lined by microscleres. Megascleres. 'True' principal megascleres absent; fibres cored by choanosomal auxiliary styles, long, slender, straight, with smooth subtylote bases and hastate points: length 124-(167.6)-180 pm, width 2-(2.8)-4 pm. Subectosomal auxiliary spicules quasidiactinal modifications of styles, long, slender, straight, with smooth subtylote bases and smooth rounded or slightly swollen tips, often with a terminal mucronate point: length 184-(196.8)-207 pm, width 1.5-(2.3)-3.0 pm. Second category of specialised ectosomal spicules absent. Echinating acanthostyles slender, straight, cylindrical, with slightly subtylote bases, rounded or slightly swollen tips, often with terminal spines obvious; granular spines evenly dispersed over entire spicule: length 55-(66.9)-74 pm, width 2-(2.9)-4 pm. Microscleres. Palmate isochelae abundant, relatively large: length 14-(15.9)-19 pm. Toxas abundant, slender, relatively short, usually with widely curved centres and slightly reflexed arms, sometimes with straight arms: length 32-(56.3)-92 pm, width 0-8-(1.4)2 - 5 pm. Fig. 24. Clathria (Clathria) bulbosa, sp. nov.: A , holotype QMG300021 (ORSTOM R1350) in situ (photo G. Bargibant); B, SEM of skeletal structure (magnified 150 x ); C, SEM of fibre characteristics (magnified 800 x). 1270 J. N. A . Hooper and C. L h i Distribution Known so far only from the Chesterfield Islands Group, coral reef habitat, 42 m depth. Remarks This species is obviously closely related to C. rugosa, although a number of characters indicate that they are not conspecific but probably sibling species. Clathria rugosa is characteristically flabellate-digitate (Fig. 8C-F), whereas C. bulbosa is bulbous encrusting (Fig. 24A). Both species have quasidiactinal auxiliary spicules forming the inorganic part of the ectosomal skeleton, but in C. bulbosa these auxiliary megascleres are completely smooth. Echinating acanthostyles are also similar between the two species, with a tapering cylindrical shape, rounded points and evenly distributed granular spines, and spicule sizes are similar for several categories of spicules but different for others (Table 4). Nevertheless, unlike C. rugosa, this species lacks principal spicules. Fibres are cored by auxiliary styles: i.e. there are two categories of auxiliary styles, those coring the spongin fibres differing only slightly in shape and dimension from the quasidiactinal forms located in the ectosomal skeleton, but both forms appear to be derived from the auxiliary category of spicules. Both species have reticulate skeletons and differentiated primary and secondary spongin fibre systems, but the skeleton of C. rugosa contains abundant detritus, placing it in the subgenus Clathriopsamma, whereas C. bulbosa lacks any foreign spicules or detritus and is typical of most other Clathria (and included in the subgenus Clathria). These minor differences between obvious sibling species, each of which on paper is referrable to a different nominal genus, illustrates the possible artificiality of the genus Clathriopsamma, as already suggested by Hooper (1991~). However, all these subgeneric groups (e.g. Clathriopsamma, Clathria, Thalysias) do appear to contain homogeneous assemblages of sister-species, with possible, but as yet undetermined, monophyletic origins. Interpretations of these levels of divergence, and indeed the whole phylogenetic basis for the existing systematics of Microcionidae, require investigation. Etymology From the Latin, bulbosus, swollen, referring to the bulbous encrusting growth form of this species. Clathria (Clathria) menoui, sp. nov. (Figs 25-26) Material Examined Holotype. Q M G301267 (ORSTOM R1535): stn 490, Ilot N'Da, S. lagoon, New Caledonia, 22°41.5'S.,166058.VE.,47 m depth, 23.v.1991, coll. J.-L. Menou, SCUBA. Description Colour. Orange-brown alive (Munsell5YR 7/10), with bright yellow larvae (2.5Y 8/10); light beige in ethanol. Shape. Massive, encrusting on coral, up to 40 mm high, 60 mm wide, and extending for 180 mm along substrate, composed of fused, bulbous (in life) or flattened flabellate (when preserved), lobate-digitate projections, 15-35 mm high, 21-53 mm wide, 3-6 mm thick (preserved material), attached to a common encrusting base. Smaller bulbous (live) or flattened (preserved) conulose digits cover apex of larger digits, 4-5 mm diameter, up to 5 mm high. Surface. Fleshy, porous, membraneous ectosome when alive, becoming detached in most places when preserved, leaving highly porous surface. Margins of digits longitudinally striated, uneven, and apex of most digits have single large oscule, 2-6 mm diameter, with prominent membraneous lip. Each oscule surrounded by converging subectosomal canal system (astrorhizae), with large canals covered by smooth membraneous surface (in contrast to adjacent surface which is highly porous, excavated by large ostia, up to 0.8 mm New Caledonian Poecilosclerids 1271 diameter); subectosomal canals collapse when preserved, leaving an excavated, highly porous surface. Texture spongy, elastic, difficult to tear. Ectosome. Membraneous, with a thin layer of subectosomal auxiliary subtylostyles lying tangential to surface, through which protrude tips of spongin fibres from primary, ascending spongin fibre system; terminal fibres tapering to sharp points, single or bifurcate at apex, 50-100 pm apart, protruding approximately 50 pm through surface, and each with a sparse paucispicular core of vestigial spicules which protrude through end of fibres. Collagen in peripheral skeleton abundant but very lightly pigmented, with small quantities of detritus scattered throughout (although not embedded in fibres). Choanosome. Skeletal structure irregularly reticulate, cavernous, with spongin fibre system dominant over spicule skeleton. Spongin fibres poorly differentiated into primary and secondary categories; primary fibres more-or-less ascending, 40-60 pm diameter, cored by uni- or paucispicular tracts of principal subtylostyles, sometimes uncored or cored by vestigial spicules of indeterminant origin (merely resembling fibre pith), and coring spicules occupy very small proportion of fibre diameter; secondary fibres 15-35 pm diameter, interconnecting primary fibres at irregular intervals, more-or-less transverse, usually uncored or rarely unispicular; fibre meshes irregular, generally large, oval or elongate, 80-250 pm diameter; echinating spicules abundant, regularly dispersed over fibres. Collagen in axial Fig. 25. Clathria (Clathria) rnenoui, sp. nov.: A, holotype QM G301267, smaller choanosomal principal subtylostyles; B, larger subectosomal auxiliary subtylostyles; C , smooth echinating subtylostyles;D, toxas; E, section through peripheral skeleton. 1272 J. N. A. Hooper and C. Ltvi region of skeleton abundant but very lightly pigmented, with numerous subectosomal auxiliary subtylostyles scattered in fibre meshes; choanocyte chambers oval, 25-45 pm diameter. Megascleres. Choanosomal principal spicules, coring spongin fibres, nearly vestigial, always with blackened axial canals, and barely different from auxiliary spicules, consisting of short, straight, slender subtylostyles, with prominent tylote bases and sharply pointed tips. Length 82-(91.2)-96 pm, width 2-(2.6)-3.5 pm. Subectosomal auxiliary subtylostyles, in ectosomal skeleton and scattered between fibre meshes, long, slender subtylostyles, straight or very slightly curved near tip, with prominent subtylote bases, often tear-drop shaped, and long, tapering, sharply pointed tips. Length 154-(172.6)-185 pm, width 1 8-(2.5)-3 a0 pm. Echinating spicules entirely smooth, straight subtylostyles, with well-marked constricted 'necks', evenly rounded bases and sharply pointed tips. Length 27-(30.1)-34 pm, width 1.5-(2.1)-3.0 pm. Microscleres. Isochelae absent. Toxas uncommon, exceedingly slender (raphide-like), with only slight angular curvature at centre and straight arms (not reflexed). Length 62-(76.3)-85 pm, width 0.5-(0.7)0.8 pm. Clathria (Clathria) menoui, sp. nov.: A , holotype QM G301267 (ORSTOM R1535), peripheral spongin fibres (scale=200 pm); B, holotype; C, SEM of skeleton (magnified 80x); D, holotype in situ (photo J.-L. Menou); E, SEM fibre characteristics (magnified 1OOOX). Fig. 26. New Caledonian Poecilosclerids Distribution Known so far only from the southern New Caledonian lagoon, coral reef habitat, 47 m depth. Remarks This species is greatly reduced in its inorganic skeletal characters. The species is distinctive in its porous, ornamented surface features, live coloration, skeletal characteristics where the organic fibre skeleton dominates over the coring spicule skeleton, the smooth echinating spicules and the raphide-like toxas with only a slight angular centre and completely straight arms. In growth form and surface characteristics, C. menoui resembles to some extent the common tropical Australian microcionid species C. (Isociella) eccentrica (Burton), but this resemblance is purely superficial. In its fibre characteristics (well-developedhorny fibres, few coring spicules; Fig. 26A, C, E), C. menoui also resembles the southern Australian species Echinoclathria leporina (Lamarck), and in the geometry and ornamentation of its echinating spicules (smooth, prominent bases, constricted 'necks', thickest at the centre of the shaft; Fig. 25C) it shows some similarities to C.(T.) transiens Hallmann. However, none of these species are closely related and the specific affinities of C. menoui are uncertain. Etymology Named for M. Jean-Louis Menou, ORSTOM, the original collector of the holotype, in recognition of his substantial contributions to the documentation of the marine biology of New Caledonia. Genus Echinochalina Thiele Echinochalina Thiele, 1903: 961 [type species Ophlitaspongia australiensis Ridley, 1884: 4421. Protophlitaspongia Burton, 1934: 562 [Siphonochalina bispiculata Dendy, 1895: 2461. Diagnosis (emended) Microcionidae with monactinal, quasi-monactinal or thin diactinal auxiliary megascleres coring irregularly reticulate or plumo-reticulate spiculo-spongin skeleton, with smooth or acanthose principal spicules echinating fibres; ectosomal skeleton also with auxiliary spicules, usually tangential or erect on surface; microscleres present or absent. Remarks Including the three New Caledonian species described here, there are now 20 nominal species of Echinochalina worldwide, 18 of which appear to be valid, and 16 of which are endemic to the Australasian region. The genus is divided into two subgenera: Echinochalina, with diactinal, quasimonactinal or monactinal auxiliary spicules coring fibres, and monactinal principal spicules echinating fibres; and Protophlitaspongia with diactinal or quasimonactinal auxiliary spicules coring fibres and diactinal or quasimonactinal principal spicules echinating fibres. This subdivision is based on an earlier phylogenetic analysis of the family (Hooper 1991~). Echinochalina (Echinochalina) intermedia (Whitelegge) (Figs 27-28, Table 12) ?Echinoclathria viminalis (in part). -Whitelegge, 1901: 87-8. Not Thalassodendron viminalis Lendenfeld, 1888: 225. Echinoclathria intermedia (in part). -Whitelegge, 1902: 214-15; Dendy, 1922: 7 1, pl. 2, fig. 8. Echinochalina intermedia.-Hallmann, 1912: 294-95, text-fig. 69; Shaw, 1927: 427; Burton, 1934: 563. Material Examined Holotype. AM, Lendenfeld no. 365, presently missing. J. N. A. Hooper and C. LBvi 1274 New Caledonian material. QM G300684 (ORSTOM R1299, fragment NTM 23868): stn 106, N. entrance, RCcif des Cinq Milles, SW. New Caledonia lagoon, 22'29.3'S., 166"44.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA; ORSTOM R433: stn 104, NE. RCcif des Cinq Milles, 22"27.5'S., 166"43.5'E., 12 m depth, 29.iv.1976, coll. P. Bourret, SCUBA; ORSTOM R267, R268, R472: stn 110, SE. IlBt RBdika, 22"31~1'S.,166°36~0'E.,19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R1278: stn 184, SE. of IlBt Ua, 22"42. 1'S.,166°49.0'E., 16 m depth, 8.vi.1977, coll. G. Bargibant, SCUBA; ORSTOM R1070: stn 249, NW. I1Bt Vua, Lagon Sud, 21°33.4'S.,166043.5'E., 35 m depth, 10.xii.1978, coll. P. Laboute, SCUBA. Comparative material. Northwest shelf, W.A.: NTM 21021: NW. of Darnpier Archipelago, 20°14'S.,117037'E., 29 m depth, 20.x.1982, coll. J. Blake, CSIRO RV 'Soela', trawl. Great Barrier Reef, Qld: QM GL707 (fragment NTM 21529): Noggin Passage, E. of Frankland Is, 17'13.5'S., 146"17'E., 42 m depth, date of collection unknown, coll. QFS, dredge; QM GL2748 (fragment NTM 21553): W. of Sudbury Reef, E. of Cairns, 17°03'S.,1460071'E., 34.5 m depth, 28.i.1981, coll. QFS, dredge; QM GL2752 (fragment NTM 21558): Stanley Reef, outer slope and lagoon, 19O16'S., 14S006'E., 13 m depth, 18.xi.1983, coll. A. Kay, SCUBA. South-east Queensland: QM GL2747 (fragment NTM 21552): off Mooloolabah, 26"43'S.,153"08'E., 86 m depth, 12.xii.1980, coll. N. Hall, trawl; QM G300395: Inner Gneerings, off Mooloolabah, 26"38.SS., 153°09.5'E., 14 m depth, 1.v. 1991, coll. J . Thorogood, SCUBA; QM G300025: Outer Gneerings, off Mooloolabah, 26°39'S.,153010'E., 25 m depth, 10.xii.1991, coll. J. N. A. Hooper and S. D. Cook, SCUBA. Description Colour. Orange-brown alive (Munsell 10R 6/10), beige in ethanol. Shape. Massive, irregular, lobate, bulbous-digitate sponge, up to 400 mm high, 300 mm maximum breadth, with small bulbous lobes forming digits (but collapsing and becoming flattened in preserved material) covering entire mass, and whole sponge excavated by large oscules; together the excavations and fused bulbous digits produce a honeycombed clathrous growth form. Surface. Paler, 'dusty', translucent, skin-like membrane covers entire surface, except around large oscules; oscules usually distributed on ends of bulbous digits, 10-30 mm diameter. Fig. 27. Echinochalina (Echinochalina) intermedia (Whitelegge): A , specimen QM G300684, choanosomal (auxiliary) quasidiactinal spicules (coring fibres); B, principal styles (echinating fibres); C, section through peripheral skeleton. New Caledonian Poecilosclerids 1275 Fig. 28. Echinochalina (Echinochalina) intermedia (Whitelegge): A , specimen ORSTOM R1278 in situ (photo P . Laboute); B, specimen QM G300684 (ORSTOM R1299) in situ (photo P. Laboute); C, SEM skeletal structure (magnified 1 0 0 ~ )D, ; SEM fibre characteristics (magnified 8 0 0 ~ ) E, ; preserved specimen QM G300684 (ORSTOM R1299). Table 12. Comparison in spicule dimensions between New Caledonian and Australian material of Echinochalina (Echinochalina) intermedia (Whitelegge) Measurements are given in micrometres New Caledonia Great Barrier Reef South-east Qld 22%. 26"s. NW. Shelf, W.A. ? Illawarra Cargados Carajos, N.s.w.~ Indian oceanA 20% 16% 34%. 200 3 140-185 4 Latitude 17-19"s. Coring auxiliary styles L W 169-(I 75.3)-189 184-(193.8)-205 184-(196.2)-208 89-(148.2)-180 1.2-(1.9)-2.5 2.0-(3.1)-5.0 1.5-(2.2)-3 1.5-(2.5)-4.0 Echinating oxeas A Dendy (1922). Hallmann (1912). 1276 J. N. A. Hooper and C. L h i Ectosome. Membraneous, arenaceous surface, without specialised spiculation, although auxiliary spicules may poke through surface forming sparse, erect bundles, or lying individually tangential to surface. Primary (ascending) choanosomal fibres also terminate close to surface, sometimes protruding slightly through ectosomal membrane and forming low conules. Collagenous spongin in peripheral skeleton heavy, granular, infiltrated with detritus, including both sand grains and foreign spicules. Choanosome. Skeletal architecture irregularly plumo-reticulate, with well-developed spongin fibre skeleton, although fibres only poorly differentiated into primary and secondary components. Primary fibres (60-100 pm diameter), more-or-less ascending to the surface, meandering through choanosome, cored by multispicular tracts composed of auxiliary spicules. Spicule tracts occupy only about 40-60% of fibre diameter, relatively well compacted although becoming more plumose towards periphery. Secondary fibres (30-60 pm diameter), usually interconnecting primary tracts, uni-, pauci- or rarely aspicular. Fibre meshes cavernous, irregular, eliptical or oval, 180-440 pm diameter, smaller in choanosomal region than in peripheral skeleton. Choanosomal mesohyl contains abundant granular collagenous spongin, heavy deposits of detritus, although detritus occurs only within fibre meshes and not inside spongin fibres, and few auxiliary spicules dispersed between fibres. Fibres sparsely echinated by thin principal styles. Choanocyte chambers difficult to observe due to dense detritus in skeleton, oval, 30-110 pm diameter. Megascleres (refer to Table 12 for dimensions). Choanosomal (auxiliary) spicules, coring fibres, long, very thin, straight, with hastate points, rounded smooth bases, varying from styles, tornostyles, or strongyles in about equal proportions; usually with blackened axial canals. Echinating (principal) styles, short, thin, straight, with hastate points and rounded, rarely subtylote smooth bases. Microscleres. Absent. Distribution Probably widely distributed throughout the Indo-west Pacific, although so far known only from Great Barrier Reef, Queensland [Direction Is (Burton 1934); Frankland Is, Sudbury Reef, Stanley Reef (present study)], southern Queensland coast [Mooloolabah (present study)], New Caledonia (present study), N.S.W., possibly the Illawarra region (Lendenfeld 1888; Whitelegge 1902; Hallmann 1912), Tasmania (Shaw 1927), Northwest Shelf of Western Australia [Dampier Archipelago (present study)] and Cargados Carajos, Indian Ocean (Dendy 1922). Depth range in the New Caledonian region is 8-35 m. Remarks The New Caledonian population of E. intermedia, a new locality record and most easterly record for the species, differs from other known populations in having abundant detritus within both the choanosomal and ectosomal mesohyl, lying between fibre meshes (but not inside fibres). In all the other specimens examined, and from the published literature, detritus is never more than minimal in this species, and usually absent completely. New Caledonian populations of E. intermedia also differ from most other specimens in having much more slender echinating styles, and in this regard these populations are most similar to those from Mooloolabah, southern Queensland (Table 12). In all other respects, all specimens are clearly conspecific in growth form, skeletal architecture and spicule geometry, although the geometry of auxiliary spicules may range from exclusively styles (seen in the specimen from the Northwest Shelf, W.A.) to virtually all strongyles [in material from Mooloolabah, southern Queensland, and also described by Dendy (1922) for material from Carnados Carajos]. In using the name 'intermedia' for this species, we follow Dendy (1922: 71, pl. 2, fig. 8), who, according to Whitelegge (1902), was familiar with the type specimen, which is presently missing, and who describes and illustrates a specimen from the Indian Ocean that agrees closely with present material. Nevertheless, the synonymy of this species is still New Caledonian Poecilosclerids 1277 slightly confused, and the validity of the 'type' material of Thalassodendron viminalis and Echinoclathria intermedia, and their relationships to the concept of Echinochalina intermedia requires further clarification. For the present, Echinochalina intermedia is used in the sense of Hallmann (1912) and Dendy (1922). Echinochalina (Protophlitaspongia) laboutei, sp. nov. (Figs 29-30, Table 13) Material Examined Holotype. QM G 300685 (ORSTOM R1301 +E084; fragment NTM 23870): stn 106, N. entrance, Rkcif des Cinq Milles, SW. New Caledonia lagoon, 22"293'S.,l66"44.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA. Description Colour. Bright red alive (Munsell 5R 4/10), beige in ethanol. Shape. Branching, digitate, arborescent sponge (220 mm high), stipitate, with holdfast and short cylindrical stalk (28 mm long, 9 mm diameter), long cylindrical branches (3-6 mm diameter), frequently bifurcate midway along their length, tapering to rounded tips. Surface. Macroscopically even; surface slightly bulbous, with concentric swellings and constrictions at more-or-less regular intervals along branches (2-4 mm apart), superficially reminiscent of an Zsis gorgonian; surface microscopically porous, finely hispid, with fibre endings from choanosome protruding slightly through surface. Oscules not visible. Texture firm, slightly compressible, branches flexible, whip-like. Ectosome. Auxiliary styles stand perpendicular or lie paratangential to surface, usually forming sparse bundles, and always lying outside spongin fibres. Ectosomal skeleton dominated by well-developed peripheral fibre skeleton, with tapering erect fibre endings protruding about 100-250 pm from the surface of the tangential spongin fibre skeleton. Peripheral spongin fibres lying perpendicular to surface always have a unispicular core of larger oxeas, protruding a short distance through fibres, whereas echinating oxeas sparsely dispersed on peripheral fibres. Mesohyl with very light collagenous spongin seen within fibre meshes. Fig. 29. Echinochalina (Protophlitaspongia) laboutei, sp. nov.: A, holotype QM G300685, choanosomal principal oxeas (coring fibres); B, echinating oxeas; C, ectosomal auxiliary styles; D, section through peripheral skeleton; E, surface fibres. 1278 J. N. A. Hooper and C. Lkvi Choanosome. Skeletal architecture regularly reticulate, renieroid, with very tightly condensed axial skeleton, occupying about 30% of branch diameter, and regularly reticulate extra-axial skeleton. Organic spongin fibre skeleton emphasised over inorganic spicular skeleton. Fibres heavy, stratified in cross-section, with differentiated primary and secondary elements and differences between axial and extra-axial fibre skeletons. Primary fibres in axis, condensed, 120-180 pm diameter, run longitudinally through branches and are cored by paucispicular plumose tracts of choanosomal oxeas; these fibres are interconnected by short, thinner, uncored secondary axial fibres, 50-90 pm diameter, together forming compressed oval meshes in axial region, 90-180 pm diameter. Primary fibres in extra-axial region are radial, 50-90 pm diameter, and extend from edge of axis, standing perpendicular to it to surface, cored by uni-, bi- or paucispicular tracts of choanosomal oxeas, usually end-on-end, rarely overlapping; primary fibres interconnected by aspicular, rarely unispicular, transverse secondary fibres, 18-40 pm diameter; fibre meshes in extra-axial and peripheral regions of skeleton usually elongate, generally larger in periphery (160-210 pm maximum diameter) than in axial region (100-170 pm maximum diameter). Sparse tertiary fibre network also present, 10-20 pm diameter, aspicular, subdividing extra-fibre skeletal meshes. Echinating oxeas heavier in axial than peripheral region, scattered mainly (but not exclusively) on primary fibres, standing perpendicular to fibres. Mesohyl with very light collagenous spongin; choanocyte chambers small elongate-oval, 40-60 pm diameter. Megascleres (refer to Table 13 for dimensions). Choanosomal principal oxeas (coring fibres) relatively short, straight, stout, with hastate, sharply pointed, telescoped ends. Echinating oxeas similar in morphology to principal megascleres, but substantially shorter, marginally thinner, also with hastate points and telescoped tips. Fig. 30. Echinochalina (Protophlitaspongia)laboutei, sp. nov.: A-B, specimen QM G300685 (ORSTOM R1301+ E084) in situ, with commensal ophiuroid (photos P. Laboute); C, SEM fibre characteristics (magnified 8 0 0 ~ )D, ; SEM skeletal structure (magnified 8 0 ~ ) . ' New Caledonian Poecilosclerids 1279 Ectosomal auxiliary styles long, slender, straight or sinuous, with rounded bases and tapering raphidiform points. Microscleres. Absent. Distribution Known only from the south-west lagoon of New Caledonia, on coral rubble and coral reef substrates, 8 m depth. Remarks Echinochalina laboutei, sp. nov., is closely related to three other species of Protophlitaspongia from the Indo-west Pacific region: E. bispiculata (Dendy, 1895: 246) from Port Phillip Heads, Vic., E. oxeata (Burton, 1934: 562) from the Great Barrier Reef and Moreton Bay, Qld, and another new species from the Noumea lagoon described below, E. bargibanti, sp. nov. All four species are very similar in several features (more-or-less digitate growth forms, reticulate skeletal structure with very well developed spongin fibres, possession of principal and echinating oxeas and subtylote auxiliary styles, all very thin and poorly silicified), but differ subtly in many other characters (size of fibres, fibre mesh diameter, presence or absence of an axial skeleton, density of spicules in the skeleton, degree of silicification of spicules, presence or absence of particular megascleres and microscleres, specific spicule size and geometry). Nevertheless, these subtle differences are considered here to justify the distinction of these taxa at the species level, and they provide a good example of character transformations between allopatric populations of sibling species. These four species (together with another, undescribed, deeper-water species from southern Queensland), represent the only known members of the subgenus Protophlitaspongia, which is probably indigenous to this region. Echinochalina bispiculata is similar to E. laboutei in its spiculation (smaller oxeas echinating fibres and larger oxeas of the same geometry coring fibres) and skeletal structure (subisodictyal-renieroid skeletal structure dominated by spongin fibres), but E. bispiculata Table 13. Comparison in spicule dimensions between the New Caledonian and Australian species of Echinochalina (Protophlitaspongia) Measurements in micrometres E. laboutei, sp, nov. (QM G300685) E. bargibanti, sp. nov. (QM (3301270) E. bispiculata (Dend~) (NMV G23 19) E. oxeata (Burton) (QM G300030) 105-(186.8)-266 2.0-(3.8)-5.0 94-(101.7)-163 1.5-(3.8)-7.5 Coring oxeas L 52-(75.1)-108 1.8-(2.9)-4.0 55-(73.3)-98 0.8-(1.9)-3.0 W 115-(156.2)-194 1.O-(1.3-2.5 Ectosomal auxiliary styles 144-(216.1)-278 Absent 1.0-(2.4-3.5 142-(181.9)-228 1.5-(2.2)-3 L W 28-(33 5)-42 2.0-(2.5)-4.0 Echinating oxeas 32-(46.3-58 31-(39.0)-45 2.0-(3.4-5.5 1.O-(2.0)-2.5 67-(76.2)-88 1.5-(2.4)-3 $ 5 L Absent Isochelae 14-(15.5)-21 Absent Absent L W Absent Absent Absent 31-(44.9)-62 0.5-(1.1)-1.5 W L Toxas 1280 J. N. A. Hooper and C. Ltvi has a very different growth form (massive, lobate-digitate), a well-developed ectosomal skeleton composed of choanosomal oxeas (same as those coring fibres) forming relatively dense, erect, multispicular brushes on the surface, lacks auxiliary styles, has well-developed extra-fibre collagenous spongin permeating the mesohyl, and the dimensions of coring spicules are nearly twice as large as those in E. laboutei (Table 13). Echinochalina oxeata has a very similar arborescent growth form as E. laboutei, but differs in its skeletal structure, spicule geometry and spicule dimensions (Table 13). Echinochalina bargibanti has a clumped-digitate growth form, lacks an axial skeleton, differs in spicule geometry and spicule dimensions (Table 13). Further comparisons between the two New Caledonian species are provided below. Etymology Named in honour of M. Pierre Laboute, ORSTOM, who has been studying and photographing the marine fauna of New Caledonia since 1971, for his significant contributions to the marine biology of New Caledonia. Echinochalina (Protophlitaspongia) bargibanti, sp. nov. (Figs 31-32, Table 13) Material Examined Holotype. QM G301270 (ORSTOM R1251): stn 271, between Ilot Tere and Ilot N'Da, S. New Caledonia, 22"48.5'S.,166"51.4'E., 30 m depth, 1.iv.l98O, coll. P. Laboute, SCUBA. Description Colour. Dark red alive (Munsell 5R 5/10), dark grey-brown in ethanol. Shape. Lobate-digitate mass (65 rnm high, 57 mm wide), composed of clumps of partially fused, small, thick digits, joined to a common base encrusting on a dead bivalve. Digits are bulbous, cylindrical (18-35 mm long, 8-15 mm diameter), thicker at apex than at point of attachment to base, usually single or bifurcate, sometimes trifurcate at apex, often partially fused with adjacent digits midway along stem, together producing a vaguely reticulate, excavated, honeycombed mass. Surface. Digits with even, rounded tips, either unornamented or with few, large conules (or secondary digits) near apical margins; conules 6-12 mm high, 4-9 mm diameter, approximately 3-7 mm apart. Oscules small, 1-2 mm diameter, located mainly on ends of, or between, secondary digits (surface conules). Oscules have slight subectosomal sculpturing (astrorhizae), converging towards each pore, but this is not extensively developed. Texture rubbery, compressible, difficult to tear. Ectosome. Sparse, tangential or paratangential tracts of auxiliary subtylostyles dispersed over ectosomal region. Where thickest auxiliary subtylostyles may form sparse plumose brushes, but usually these spicules are scattered. Terminal, ascending primary fibres protrude through ectosomal spicule skeleton for up to 150 pm, containing a core of principal oxeas, the ends of which barely protrude through ends of fibres. Relatively sparse clumps of lightly pigmented collagen usually associated with ectosomal spicule skeleton. Choanosome. Skeletal architecture regularly reticulate throughout, without any differentiation of axial or extra-axial/peripheral regions. Spongin fibres dominate skeleton, and spicules both greatly reduced in abundance and poorly silicified (all with blackened axial canals). Spongin fibres divided into primary ascending and secondary transverse (connecting) components, with a very thin tertiary network also apparent. Primary fibres, 40-60 pm diameter, meandering throughout skeleton but more-or-less ascending to surface, terminating in small digitate microconules, cored by uni- or paucispicular tracts of choanosomal principal oxeas, occupying only a small proportion of fibre diameter. Secondary connecting fibres usually transverse, 15-25 pm diameter, aspicular or unispicular, and tertiary fibres, 8-20 pm diameter, invariably aspicular and interconnecting both secondary and primary fibre system. Fibre meshes cavernous, square or rounded, 80-180 pm diameter, containing very little New Caledonian Poecilosclerids 1281 visible collagen, although distinct clumps of granular collagen plus principal oxeas and isochelae scattered within choanosome, particularly around major fibre nodes. Echinating acanthostyles sparse, usually associated with small fibre nodes protruding from a primary or secondary fibre. Choanocyte chambers small, elongate-oval, 35-60 pm diameter. Megascleres (refer to Table 13 for dimensions). Choanosomal principal oxeas (coring fibres) relatively short, slender, with hastate, sharply pointed ends, occasionally telescoped points. Ectosomal auxiliary subtylostyles long, slender, invariably straight, with enlarged subtylote bases, rounded or slightly pointed, and tapering to sharply pointed or slightly rounded tips. Echinating oxeas rare, similar in morphology to principal oxeas, but marginally shorter and of similar thickness. Microscleres. Palmate isochelae moderately large, common, usually found aggregated in clumps of collagen in choanosome. Toxas are absent. Fig. 31. Echinoclathria (Protophlitaspongia) bargibanti, sp. nov.: A, holotype QM G301270, ectosomal auxiliary subtylostyles; B, choanosomal principal oxeas (coring fibres); C, echinating oxeas; D, palmate isochelae; E, section through peripheral skeleton. 1282 J. N. A. Hooper and C. L6vi Distribution Known only from the south-west lagoon of New Caledonia, on coral rubble substrate, 30 m depth. Remarks If a phylogenetic study of Protophlitaspongia eventually shows E. laboutei (New Caledonia) and E. bispiculata (south-east Australia) to be sister-species, then it is quite probable that E. bargibanti (New Caledonia) is also a sister-species of E. oxeata (north-east Australia). Both pairs of species have comparable similarities in their skeletal structure and fibre characteristics, although they differ subtly in many other ways. Echinochalina bargibanti differs from the other three described sister-species in having isochelae microscleres (whereas E. laboutei has toxas, and the others have no microscleres Fig. 32. Echinoclathria (Protophlitaspongia) bargibanti, sp. nov.: A, holotype QM G301270 (ORSTOM R1251), SEM fibre characteristics (magnified 4 0 0 ~ ) ; B, SEM skeletal structure (magnified 100 x ) ; C, three specimens in situ (holotype in right foreground) (photo P. Laboute); D, holotype; E, fibre structure, showing coring spicules and sparse aggregates of collagen plus spicules located near fibre nodes (indicated by arrows) (scale =200 pm). New Caledonian Poecilosclerids 1283 at all). This species also differs from E. laboutei in growth form: lacking a condensed axial skeleton, having instead undifferentiated axial and extra-axial regions; having a more cavernous skeleton, with rounded or square fibre meshes, whereas E. laboutei has elongate meshes; echinating oxeas are very scarce in the present species and the geometry of auxiliary spicules differs between the two species. Spicule measurements of all these species are compared in Table 13. Etymology Named for M. Georges Bargibant, ORSTOM, in recognition of his substantial contributions in documenting the marine biology of the New Caledonian region. Family RASPAILIIDAE Hentschel Definition Poecilosclerida usually with very hispid surface, specialised ectosomal skeleton of brushes of small thin styles or oxeas, surrounding individual long thick styles or oxeas; choanosomal skeleton varies from compressed axial skeleton, to plumo-reticulate or exclusively reticulate structures; spongin fibres enclose choanosome styles, oxeas or both; spined styles (acanthostyles) or highly modified styles echinate fibres; microscleres usually absent, although single raphides or bundles (trichodragmata) occur in some genera (from Hooper 1991b). Remarks The present work brings the total number of raspailiid species in the New Caledonian region to seven, three apparently endemic and four also recorded from other Indo-west Pacific localities. These species are: Raspailia wilkinsoni Hooper (Great Barrier Reef), Ceratopsion clavata Thiele (Moreton Bay, Qld, and Japan), C. palmata Hooper (tropical western and northern Australia), Aulospongus clathrioides LCvi (endemic), A. gardineri (Dendy) (Amirante, W. Indian Ocean), Plocamione pachysclera ( L h i & LCvi) (endemic, deep water), and Lithoplocamia dolichosclera LCvi & LCvi (endemic). Three of these species (R. wilkinsoni, C. clavata and C. palmata) are new records for the region. The taxonomic system used here follows Hooper (1991b). Genus Raspailia Nardo Raspailia Nardo, 1833: 522 [type species Raspailia typica Nardo, 1833: 5221 [fullsynonymy and discussion given by Hooper (1991b: 1195)l. Diagnosis Raspailiidae with axial compression and differentiated axial and extra-axial skeletons, or sometimes reduced to reticulate structures (subgenus Clathriodendron); axial skeleton typically condensed, reticulate, with fibres and spicules forming distinct tracts; extra-axial skeleton plumo-reticulate, perpendicular to axis, cored by large styles or oxeas in tracts, or plumose, with single extra-axial spicules; specialised ectosomal skeleton present, with bundles of erect, small styles surrounding bases of protruding extra-axial megascleres; choanosomal spongin fibres echinated by club-shaped acanthostyles (subgenus Raspailia), acanthose rhabdostyles (subgenus Raspaxilla), modified cylindrical styles with greatly enlarged bases (subgenus Hymeraphiopsis), or secondarily lost (subgenus Syringella); microscleres absent (from Hooper 1991b). Remarks This genus has been recently revised, mainly on the basis of Australian material, and five subgenera were recognised (as defined above). It is possible, however, that several more subdivisions within the genus may be appropriate eventually, although more extensive studies of other faunas are first required. Prior to the present work, no species of Raspailia had been recorded from the New Caledonian region, although Plocamione pachysclera was initially referred to the genus by LCvi and LCvi (1983b). 1284 J. N. A. Hooper and C. LCvi Raspailia (Raspailia) wilkinsoni Hooper (Figs 33-34, Table 14) Raspailia wilkinsoni Hooper, 1991b: 1220-22, figs 17-1 8. Material Examined Holotype. NTM 22734: Back Reef, Davies Reef, off Townsville, Great Barrier Reef, Qld, 18"5O'S.,147"39'E.,15 m depth, ll.viii.1989, coll. C. R. Wilkinson (AIMS ref. R22). New Caledonian material. QM G300744 (ORSTOM R766): stn 206, Banc de la Torche, Ile des Pins, 22°51.1'S.,167040.0'E., 36 m depth, 8.iii.1978, coll. P. Laboute, SCUBA; QM G300747 (ORSTOM R322): stn 196, barrier reef M'Bkre, false passe de Uitoe, 22°18~1'S.,166011.1'E.,45 m depth, 9.xi.1977, coll. J.-L. Menou, SCUBA; ORSTOM unregistered: RCcif exterior de Passe Boulari, 60 m depth, l.vi.1977. coll. P. Laboute, SCUBA. Comparativematerial. ORSTOM TI: precise locality unknown, Tanna, S. Vanuatu (New Hebrides). Description Colour. Dark orange-brown alive (Munsell 5YR 5/8), dark brown in ethanol. Shape. Arborescent, digitate, bushy, 80-200 mm high, 60-110 mm maximum breadth of branches, with short or long cylindrical stalk and basal holdfast, 12-85 mm long, 4-8 mm diameter, and thick, bifurcate, cylindrical branches, 25-95 mm long, 5-1 1 mm diameter, tapering to sharp points at apex. Surface. Shaggy, prominently microconulose surface on branches, more compact on stalk. Surface conules up to 3 mm long, narrow, close-set. Oscules not observed; inhalent pores small, up to 0 . 5 mm diameter, scattered between surface conules. Texture firm, compressible, with easily flexible branches and less flexible stalk. Ectosome. Membraneous, without specialised raspailiid skeleton, although smaller, thinner, wispy auxiliary spicules scattered in peripheral skeleton mostly tangential to surface. Subectosomal region dominated by micro-digitate projections, 1 5-3 mm long, predominantly cored by ascending, plumose, multispicular tracts of long, thick, subectosomal auxiliary spicules, protruding through surface for about 100 pm, usually in bundles. Ascending subectosomal spicule tracts interconnected by transverse spicule tracts composed of choanosomal principal oxeas. Whole peripheral skeleton cavernous, with widely spaced extra-axial spicule tracts producing more-or-less rectangular meshes, up to 500 pm diameter. Peripheral spicule tracts multi- or paucispicular, heavily echinated by acanthostyles, more so than in axial region. Collagenous spongin abundant in mesohyl of peripheral skeleton. Choanosome. Skeleton reticulate, with well differentiated compressed axial and reticulate extra-axial regions; axial skeleton tightly compressed, composed of close-set, reticulate, heavy spongin fibres, 100-150 pm diameter, with stratified spongin fibres producing tight meshes, up to 150 pm diameter. Fibres cored by choanosomal oxeas in pauci- or multispicular tracts, and lightly echinated by acanthostyles. Mesohyl in axial region only lightly invested with collagenous spongin. Megascleres (refer to Table 14 for dimensions). Choanosomal principal oxeas long, thin or thick, usually symmetrically curved, tapering to sharp points. Subectosomal auxiliary spicules predominantly oxeas, less commonly styles (cf. holotype), long, thick, straight or slightly curved, often anisoxeote, with rounded bases (for styles) and tapering to sharp points. Ectosomal auxiliary spicules short, thin, whispy, varying from symmetrically curved oxeas to sinuous anisoxeas, tapering to sharp points. Echinating acanthostyles short, thixf, straight, with subtylote bases and tapering rounded blunt tips; spines long, slender, conical, sharply pointed, concentrated mainly on bases and points. Microscleres. Absent. New Caledonian Poecilosclerids 1285 Distribution Central region of the Great Barrier Reef, south-west lagoon of New Caledonia and Tanna, southern Vanuatu, depth range 15-60 m, on live and dead coral substrates. Remarks The material described above from New Caledonia and Vanuatu is only the second record of this species, which was previously known only from the holotype. There are a number of differences between these three populations, but these are considered here to be relatively minor and in our opinion cannot justify their differentiation at the species level. Both the holotype and the Vanuatu specimen lack any sort of ectosomal spiculation, whereas New Caledonian specimens have sparse, wispy tracts of auxiliary oxeas (or anisoxeas) mostly lying tangential to the surface. These ectosomal auxiliary spicules do not form a specialised ectosomal skeleton, synapomorphic for the Raspailiidae, and hence in this regard the original concept of the species is retained. Similarly, the larger subectosomal auxiliary spicules in both the holotype and the Vanuatu specimen are exclusively styles, Fig. 33. Raspailia wilkinsoni Hooper: A, specimen QM G300744, choanosomal principal oxeas; B, subectosomal auxiliary oxeas and styles; C, ectosomal auxiliary oxeas/ anisoxeas; D, echinating acanthostyles; E, section through peripheral skeleton. J. N. A. Hooper and C. L6vi 1286 Fig. 34. Raspailia wikinsoni Hooper: A , specimen QM G300744 (ORSTOM R766), SEM skeletal structure (magnified 200x); B, preserved specimen; C, unregistered ORSTOM specimen in situ (photo P . Laboute); D, specimen QM G300744 (ORSTOM R766), SEM fibre characteristics (mangified 600 x ); E, preserved specimen QM G300747 (ORSTOM R322). Table 14. Comparison in spicule dimensions between New Caledonian, Vanuatuan and Great Barrier Reef populations of Raspailia (Raspailia) wilkinsoni - Measurements are given in micrometres; comparative data from Hooper (1991b) - New Caledonia Vanuatu Great Barrier Reef - - - - Choanosomal oxeas L W 188-(238.1)-285 4-(6.2)-10 650-800 6-8 151-(214.0)-346 4-(6.6)-10 Subectosomal auxiliary oxeas and styles L W 255-(382.9-524 8-(9.3-12 L W Ectosomal auxiliary oxeas and anisoxeas 106-(159.6)-208 Absent I-(]. 3-2.5 250-350 10 Echinating acanthostyles 267-(353 ~3)-542 3-(5.6)-10 Absent New Caledonian Poecilosclerids 1287 whereas in New Caledonian material these spicules are mostly oxeas, less commonly styles, occasionally anisoxeas. Nevertheless, the dimensions of these spicules between Australian and New Caledonian material are closely comparable (Table 14). Similarly, transverse sections of the subectosomal spicule and fibre skeletons in New Caledonian specimens are thicker, and contain a larger core of multispicular tracts, whereas those in the holotype are much less robust. Genus Ceratopsion Strand Ceratopsion Strand, 1924: 33 [type species Ceratopsis expansa Thiele, 1898: 571 [full synonymy and discussion given by Hooper (1991b: 1327)l. Diagnosis Raspailiidae with well-differentiated axial and extra-axial skeletons, and condensed reticulate axial skeleton cored by styles, anisoxeas or oxeas, often sinuous; extra-axial skeleton radially arranged long styles, anisoxeas or strongyles, sinuous or straight, projecting through surface, surrounded by specialised raspailiid ectosomal skeleton of oxeas or styles; echinating megascleres absent; microscleres absent (from Hooper 1991b). Remarks Eleven species of Ceratopsion have been described (Hooper 1991b), extending in distribution from Japan, New Zealand, southern Indonesia, northern and eastern Australia, South Africa and the Mediterranean. The present records of C. clavata and C. palmata from New Caledonia are the easternmost records of the genus in the Pacific. Ceratopsion clavata Thiele (Figs 35-37, Table 15) Ceratopsis clavata Thiele, 1898: 57, pl. 5, fig. 23, pl. 8, fig. 42a-c. Ceratopsion clavata. -Hooper, 1991b: 128, table 18. Material Examined Holotype. Unknown, possibly ZMB: Enoshima region, Sagami Bay, Japan, 139°30'N.,35010'E., 130 m depth. New Caledonian material. QM G300743 (ORSTOM R685): stn 202, NW. Rtcif Ana, 21°22. l'S., 166"01.11E., 35 m depth, 8.ii.1978, coll. G. Bargibant, SCUBA; QM G300745 (ORSTOM R755) (4 specimens): stn 209, NNE. I16t Ugo, Ile des Pins, 22°28.1'S.,166055.2'E., 25-35 m depth, l0.iii. 1978, coll. P. Laboute, SCUBA; QM G300018 (ORSTOM R1260): stn 282, 'travers de la cascade', Tao, 20°32~1'S.,166"49~0'E.,37 m depth, 26.v.1980, coll. P. Laboute, SCUBA; QM G300720 (ORSTOM R328): stn 114, NE. Ili3t Tareti, Banc Gail, 22°221'S.,166039.0'E., 35 m depth, 29.vi. 1976, coll. P. Laboute, SCUBA. Comparative material. QM G301289 (2 specimens): Hanlon Light, Moreton Bay, Qld, 27"28+5'S., 153"20.9'E., 10 m depth, 3.xi.1992, coll. J. N. A. Hooper and S. D. Cook, SCUBA. Description Colour. Yellow (Munsell 2.5Y/8-10) or pale orange alive (5YR 7-10), pale beige to light brown in ethanol. Shape. Arborescent digitate, variable morphology ranging from bushy to elongate digitate, whip-like, 60-360 mm maximum height, 25-115 mm maximum breadth (width of branching), with short, thick, cylindrical basal stalk, 5-28 mm long, 2-11 mm diameter, enlarged basal holdfast, and long, bifurcate, bulbous-cylindrical branches, 25-125 mm long, 5-12 mm diameter (inclusive of surface conules), tapering towards ends. Surface. Prominently conulose surface, although varying between specimens (poorly developed in G300720); conules close-set, irregular, sharply pointed, shaggy, up to 5 mm long, 3 mm basal diameter. 1288 J. N. A. Hooper and C . L h i Ectosome. Predominantly membraneous ectosomal skeleton, with subectosomal auxiliary spicules protruding for up to 200 pm from surface, and with sparse tangential tracts of ectosomal auxiliary oxeas dispersed around the vicinity of larger subectosomal spicules, although not producing a specialised raspailiid skeleton. Heavy collagen in peripheral region, with large areas completely clear of any spicules. Choanosome. Skeleton divided into three distinct components: compressed axial core, consisting of short, tightly meshed, heavy spongin fibres, 60-100 pm diameter, producing small oval meshes, 30-70 pm diameter, sparsely to moderately cored by sinuous choanosomal principal spicules; radial extra-axial skeleton, consisting of subectosomal auxiliary spicules, occurring individually or in bundles, embedded in and standing perpendicular to axial fibres, Fig. 35. Ceratopsion clavata Thiele: A , specimen QM G300743, choanosomal principal strongyles/ strongyloxeas/ anisoxeas; B, subectosomal auxiliary oxeas/ anisoxeas/ styles; C , ectosomal auxiliary oxeas; D, section through peripheral skeleton. New Caledonian Poecilosclerids 1289 in parallel rows, and extending outwards for only one spicule length; and plumose extra-axial skeleton, consisting of large (up to 5 mrn) plumose (or less obviously plumo-reticulate), multispicular tracts of subectosomal auxiliary spicules, and including intermingled sinuous coring spicules from choanosome; latter part of skeleton most prominent, corresponding to large surface conules. Mesohyl in axial region only lightly invested with collagenous spongin, whereas closer to periphery there are heavier deposits of more granular collagen. Choanocyte chambers oval, up to 120 pm diameter. Megascleres (refer to Table 15). Choanosomal principal megascleres, predominantly in axial skeleton, usually sinuous, sometimes merely curved, rarely straight, ranging from strongyles with evenly rounded ends, strongyloxeas with telescoped, mucronate ends, to anisoxeas with a combination of rounded and telescoped ends. Queensland specimens predominantly with sinuous principal spicules; New Caledonian specimens predominantly with curved principal spicules. Subectosomal auxiliary megascleres, forming extra-axial skeletal tracts, range in size and geometry from small wispy oxeas, anisoxeas or styles, to large thick styles and anisoxeas; most spicules have telescoped ends, but larger ones may have evenly rounded ends; spicules straight, symmetrically or asymmetrically curved or less often sinuous. Queensland specimens predominantly with thinner, sharply pointed telescoped oxeas; New Caledonian specimens predominantly with thicker, rounded telescoped oxeas. Ectosomal auxiliary spicules thin whispy oxeas, slightly curved, usually asymmetrical curvature, with tapering fusiform points. Echinating megascleres absent. Microscleres. Absent. Fig. 36. Ceratopsion clavata Thiele: A-E, New Caledonian material; A , specimen QM G300720 (ORSTOM R328) in situ (photo P. Laboute); B, 3 specimens QM G300745 (ORSTOM R755); C , specimen QM G300743 (ORSTOM R685); D, specimen QM G300018 (ORSTOM R1260), SEM axial and extra-axial skeletons (magnified 100x); E, SEM extra-axial skeleton (magnified 3 0 0 ~ ) . 1290 J. N. A. Hooper and C. LBvi Distribution Sagami Bay, Japan; Moreton Bay, Queensland; and south-west lagoon of New Caledonia. Remarks This is the first record of the species since it was initially described from Japan by Thiele (1898). The species shows remarkable variability in the extent to which surface conules are developed (Figs 36A-C, 37A), in spicule size (Table 15), and in the terminations on sub- Fig. 37. Ceratopsion clavata Thiele: A-C, Moreton Bay material; A , specimen QM G301289 in situ (photo J . N . A. Hooper); B, SEM peripheral skeleton (magnified 200 x ) ; C , SEM transverse section through branch (magnified 80 X ) (scale = 500 km). New Caledonian Poecilosclerids 1291 Table 15. Comparison in spicule dimensions between specimens of Ceratopsion clavata Thiele, 1898 The great range of spicule dimensions within this species is illustrated; measurements are given in micrometres; comparative data from Hooper (1991b) Japan ~ o l o t y ~ eQM ~ G300720 New Caledonia QM G300743 QM G300745 QM G300018 Moreton Bay QM G301289 Choanosomal strongyles/anisoxeas L W 690-1500 10-30 294-(405 '3)-680 252-(439.1)-790 251-(447.8)-868 284-(577.8)-912 2.0-(2.8)-4.0 1.5-(2.9)-4.0 2.0-(3.6-4.5 4-(5.6)-9 L W 550-1500 10-50 234-(310.8)-418 258-(381.2)-590 172-(317.4)-453 232-(319.8)-422 3-(4.7)-6 4-(6.3)-8 6-(7.8)-10 7-(10.3-13 L W 60-1 10 3 192-(251.4)-302 126-(193.3)-263 165-(189.6)-232 163-(213.0)-246 1.0-(1.7)-3.0 0.8-(1.4)-2.0 0.8-(1.3)-2.0 1-(2.1)-4 422-(573 '3)-805 2.0-(3.9)-6.0 Subectosomal auxiliary styles/anisoxeas 216-(258.0)-288 4.0-(5.4)-8.0 Ectosomal auxiliary oxeas A 142-(202.4)-268 0.8-(2.1)-3.0 From Thiele (1898: 57). ectosomal auxiliary spicules (oxeas with rounded ends, telescoped ends, sharply pointed, etc.; Fig. 35A-C). In growth form, one specimen (QM G300720) has greatly elongated and spindly branching, with nearly vestigial surface sculpturing, whereas others (e.g. QM G300743 from New Caledonia and QM G301289 from Moreton Bay) are very bushy, and also have prominent close-set conules, as in the holotype. In spiculation, one specimen from New Caledonia in particular (QM G30018) has generally larger and thicker spicules than any of the other New Caledonian specimens, although these appear to vary greatly between all known specimens, and spicule thickness seems to be highly variable in the species. Moreover, all specimens described above from New Caledonia and southern Queensland have substantially smaller spicule dimensions than those published for the Japanese holotype (Thiele 1898). Nevertheless, resemblance is very close in all other features between the present material and Thiele's (1898) original description, and this material is considered here to be conspecific. Ceratopsion clavata differs from most other species of the genus in lacking a clearly defined, specialised raspailiid ectosomal skeleton: specialised ectosomal auxiliary spicules are present in this species, but these are obviously vestigial and do not form plumose surface structures. By comparison, in all other species examined of Ceratopsion, this feature is consistently present (cf. Hooper 1991b: 1327-39). Ceratopsion clavata has sinuous axial strongyles (some modified to anisoxeas or strongyloxeas, cf. the holotype description) and extra-axial styles (many modified to anisoxeas or oxeas, usually with telescoped ends). In this regard, the species shows some similarities to species from South Africa [C. microxephora (Kirkpatrick, 1903)l and the Mediterranean (C. minor Pulitzer-Finali, 1983). In growth form it resembles C. dichotoma (Whitelegge, 1907) most closely, although surface sculpturing is usually more prominent in the present species. The species is contrasted further with other members of the genus in Hooper (1991b). Ceratopsion palmata Hooper (Figs 38-39, Table 16) Ceratopsion palmata Hooper, 1991b: 1337-9, figs 78-9, table 18. Material Examined Holotype. N T M 23051: N, of Amphinome Shoals, Northwest Shelf, W.A., 19"19.7-23.3'S., 119"08.8-12~2'E.,50 m depth, 19.vii.1987, coll. J. N . A. Hooper, USSR RV 'Akademik Oparin', beam trawl. 1292 J. N. A. Hooper and C. L6vi New Caledonian material. NTM 23899 (fragment QM G300698): exterior of Grand Rkif Mbere, off Noumea, 22°20'S.,166013'E., 28 m depth, 27.ix.1990, coll. J. N. A. Hooper, SCUBA, stn JH-90-021. Comparative material. Refer to Hooper (1991b: 1337). Description Colour. Yellow-beige alive (Munsell 7.5YR 7-10), beige in ethanol. Shape. Small fan, 240 mm high, 150 mm wide, 8 mm maximum thickness, with folded margins and short stalk, 12 mm long, 5 mm diameter. Fig. 38. Ceratopsionpalmata Hooper: A, specimen QM G300698, choanosoma1 principal oxeas; B, subectosomal auxiliary styles/ oxeas; C,ectosomal auxiliary oxeas; D, section through peripheral skeleton. New Caledonian Poecilosclerids 1293 Surface. Optically even, microscopically villose and hispid,surface; oscules not seen; texture firm, rubbery. Ectosome. Darkly pigmented collagenous surface, with long extra-axial styles protruding, more-or-less surrounded by sparse bundles of thin ectosomal oxeas but not necessarily confined to the region at the point of insertion of the extra-axial spicules. Fig. 39. Ceratopsion palmata Hooper: A , specimen NTM 23899 in situ; B, specimen QM G300698 in situ (photo J. Hooper); C, SEM skeletal structure (magnified 8 0 ~ ) . Table 16. Comparison in spicule dimensions between specimens of Ceratopsion palmata Hooper, 1991 Measurements, given in micrometres, are presented as range (mean) of lengths and widths; comparative data from Hooper (1991b) Holotype Australia (n = 5) New Caledonia (n=1) L W 229-(348.4)-391 4-(12.9)-18 Choanosomal oxeas 231-612 3-20 269-(293.0)-3 12 6-(9.9)-13 L W 521-(722.3-808 5-(10.3)-18 Subectosomal styles 583-2232 3-49 549-(645.4)-751 5-(6.6)-8 L W 202-(340.1)-442 1-5-(3.5)-6 Ectosomal oxeas 191-741 0.5-15 194-(247.9)-357 2-(3.3)-4 1294 J. N. A. Hooper and C. Ltvi Choanosome. Axial and extra-axial skeletons clearly differentiated and extra-axial skeleton composed of two components. Axis compressed, consisting of bundles of oxeas mostly running longitudinally through the sponge, bound together by primary spongin fibres, 60-1 10 pm diameter, in vaguely cylindrical bundles, interconnected with thinner secondary fibres. Extra-axial skeleton perpendicular to axis, running to surface, composed of plumose, ascending multispicular tracts interconnected by fewer transverse, uni- or paucispicular tracts, both sorts of fibres cored by choanosomal oxeas, producing a plumo-reticulate extra-axial skeleton. Embedded in the ends of extra-axial plumose tracts are individual subectosomal megascleres, standing perpendicular to surface. Mesohyl contains abundant, darkly pigmented, granular collagen. Choanocyte chambers small, oval, up to 70 pm diameter. Megascleres (refer to Table 16 for dimensions). Choanosomal principal oxeas, in both axial and plumo-reticulate extra-axial skeletons, usually robust, slightly curved, rarely straight, with sharply pointed or faintly telescoped ends. Subectosomal auxiliary styles, in peripheral extra-axial skeleton, long, slender, slightly curved near base, tapering to fusiform points, with evenly rounded bases or occasionally modified to oxeas. Ectosomal auxiliary spicules are symmetrical oxeas, relatively long, slender, slightly curved or straight. Echinating megascleres absent. Microscleres. Absent. Distribution Tropical Western Australia, northern Australia and New Caledonia; coral reef and interreef regions, 5.5-76 m depth. Remarks The present specimen is the most easterly record for the species, and there are some slight differences between it and the Australian population. The New Caledonian specimen is yellower in coloration; it is a flat fan with folded margins rather than with digitate projections on the margins; the ectosomal skeleton is more poorly developed, with only few ectosomal bundles seen; extra-axial styles are sometimes modified to oxeas; and spicule sizes differ slightly from other material (Table 16). However, these differences are minor and the species appears to be relatively homogeneous across its broad distribution. Relationships between C. palmata and other species are discussed in detail by Hooper (1991b). Genus Aulospongus Norman Aulospongus Norman, 1878: 267 [type species Haliphysema tubulatus Bowerbank, 1873: 291 [full synonymy and discussion given by Hooper (1991b: 1307)l. Diagnosis Raspailiidae with specialised growth form of fused plumose skeletal columns, without or with greatly reduced reticulate skeletal structure; axial and extra-axial skeletons poorly differentiated, without axial compression; choanosomal skeletal tracts plumose or bifurcate microcionid-like; extra-axial skeletal tracts radial, plumose, diverging at periphery; choanosomal megascleres rhabdostyles; extra-axial skeletal columns echinated by acanthorhabdostyles or acanthostyles; specialised raspailiid skeleton present or absent; microscleres may include raphides (from Hooper 1991b). Remarks Two species of Aulospongus have been previously recorded from the New Caledonian region, one deeper-water species [A. gardineri (Dendy) from 300-430 m depth] and one shallow-water species (A. clathrioides LCvi). The former species is also known from Amirante, whereas the latter is endemic and relatively abundant in the lagoon. 1295 New Caledonian Poecilosclerids Aulospongus clathrioides LCvi (Figs 40-41) Aulospongus clathrioides LBvi, 1967: 21, text-fig. 5, pl. 2, fig. c. -Hooper, 1991b: 1307, fig. 67a-d. Material Examined Holotype. MNHN DCL 823: Canala, Melasceu, New Caledonia, 04.i.1962, coll. Mission SingerPolignac. Other New Caledonian material. QM G300721 (ORSTOM R714): stn 198, pinnacle S. of Canyon Central, Chenal des Cinq Milles, 22"30.4'S.,166"45.1'E., 35 m depth, 153.1978, coll. G. Bargibant, SCUBA; QM G300742 (ORSTOM 'cfR714'): stn 106, N. entrance, RBcif des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 16.ii.1978, coll. P. Laboute, SCUBA; QM G300746 (ORSTOM R515): same locality, 30.iv.1976, 8 m depth; ORSTOM R854: stn 198, pinnacle S. of Canyon Central, Chenal des Cinq Milles, 22°30.4'S.,166045.1'E., 40 m depth, ll.iv.1978, coll. P. Laboute, SCUBA; ORSTOM R696: Plateau des Cinq Milles, 22°29.3'S.,166044.4'E., 35 m depth, 16.ii.1978, coll. P. Laboute, SCUBA. Fig. 40. Aulospongus clathrioides L h i : A , specimen QM G300746, choanosomal principal styles/ anisoxeas; B, echinating acanthostyles; C , section through peripheral skeleton. 1296 J. N. A. Hooper and C. LCvi Description Colour. Yellow orange alive (Munsell 7.5YR 7/10), khaki brown in ethanol. Shape. Arborescent, bushy, 55-180 mm high, 40-110 mm maximum branching width, with short woody stalk, 22-40 mm long, 2-8 mm diameter, and enlarged basal attachment; branches prolific, usually repeatedly bifurcate and anastomosing, producing a reticulate bushy mass. Branches thinly cylindrical or slightly flattened, up to 6 mm diameter, tapering to rounded bifurcated tips. Surface. Thin translucent skin-like ectosomal membrane stretched between surface conules, intact in only few places in preserved material. Branches with uneven, shaggy, very prominent microconulose surface, and macroscopically hispid; basal stalk woody, more darkly pigmented; branch tips even, hispid only. Oscules occasionally seen on lateral sides of branches, between surface conules, up to 2 mm diameter. Texture of branches soft, flexible, spiky; stalk firm, flexible. Ectosome. Without specialised spiculation, with plumose brushes of choanosomal principal styles protruding up to 150 pm from peripheral fibres. Peripheral skeleton dominated by swollen spongin fibres, with fibre nodes protruding through surface and forming surface conules; fibre nodes 350-600 pm long, up to 250 pm maximum diameter. Mesohyl in peripheral skeleton with heavy collagenous spongin. Choanosome. Skeleton predominantly plumose, although a slightly compressed axial skeleton also present; both axial and extra-axial skeletons dominated by well-developed spongin fibre system, with spicule skeleton proportionally reduced. Axial fibres small, close-set, with vestigial core of choanosomal principal styles; axial fibre meshes 60-90 pm diameter, with only moderate quantities of collagen. Extra-axial fibres occupy most of the branch diameter, with primary ascending spongin fibres, 60-200 pm diameter, cored by multispicular plumose columns of choanosomal principal styles (although spicule columns Fig. 41. Aulospongus clathrioides Lhi: A , specimen QM G300721 (ORSTOM R714) in situ (photo G . Bargibant); B, specimen (ORSTOM R854) in situ (photo P . Laboute); C, specimen QM G300742 (ORSTOM 'cfR714'), SEM of peripheral skeleton (magnified 8 0 ~ ) D, ; SEM of fibre characteristics (magnified 1 0 0 0 ~ ) . New Caledonian Poecilosclerids 1297 occupy less than 50% of fibre diameter, and plumose brushes composing columns are relatively sparse); secondary fibres short, thin, 100-150 pm long, 30-60 pm diameter, cored by 0-2 choanosomal styles, and anastomoses between primary and secondary fibres produce elongate cavernous meshes, 200-550 pm diameter. Echinating acanthostyles moderately sparse, dispersed relatively evenly over primary and secondary fibres, but probably more abundant in extra-axial rather than axial skeleton. Collagenous spongin heavy in extra-axial region; choanocyte chambers small, oval, 15-40 pm diameter. Megascleres. Choanosomal principal styles of a single category only, although length and thickness variable; usually long, slender, slightly curved, occasionally sinuous, with rounded smooth bases, sometimes anisoxeote, tapering fusiform points. Length 145-(249.3)454 pm, width 3-(5.1)-7 pm. Subectosomal and ectosomal megascleres absent. Echinating acanthostyles long, slender, slightly curved towards basal end, with rounded or slightly swollen, mostly smooth bases, pointed or slightly swollen tips, and evenly covered with small granular spines (except for basal region). Length 58-(70.5)-82 pm, width 1.5(2.4)-4.0 pm. Microscleres. Absent. Distribution Known only from the New Caledonia lagoon, inter-reef region, coral and sand substrate, 8-40 m depth. Remarks Aulospongus clathrioides is slightly unusual to the genus in retaining the rudiments of an axial skeleton, although this structure is greatly reduced in relation to the extra-axial region. By comparison, most species of Aulospongus lack axial compression or any marked differentiation between axial and extra-axial regions. Similarly, although typical members of the genus have a peculiar growth form consisting of fused bundles of plumose fibres, together forming rounded, massive and clathrous structures, A. clathrioides is digitate (Fig. 41A), and as such, more similar to many species of Raspailia. However, the prominent surface fibre bundles in this species, consisting of large, swollen spongin fibres protruding through the peripheral skeleton, are distinctly homologous to the typical Aulospongus condition, and in this regard there are also clear affinities with other members of the genus. Further discussion of A. clathrioides, and its relationships to other Aulospongus, has been presented by Hooper (1991b: 1307-8). Discussion From our relatively rudimentary knowledge of the sponge fauna of the New Caledonian region, based exclusively on the recent works of LCvi (1967 et seq.), Ltvi and LCvi (1978 et seq.) and Desqueyroux-Faundez (1984, 1987), it has been suggested that levels of endemism for the whole sponge fauna in this region are relatively high (about 60% for all Demospongiae). However, most of these 'apparent endemics' were restricted to the deeperwater shelf and slope fauna, with ancestral links to the Cretaceous fauna of northern New Zealand (LCvi and LCvi 1978): of the 76 species described from deeper-waters, 55 have not been recorded from any other province (72% endemism). In contrast, it was previously thought that the shallower-water fauna contained far fewer indigenous species, having a 'widespread Indo-Pacific' fauna (Ltvi 1979), in common with many other groups of marine invertebrates (e.g. Crosnier and Bouchet 1991). Of the 68 shallow-water species described prior to the present study, only 25 had not been recorded from any other province (3770 endemism). The present results, concentrating on the shallow-water Poecilosclerida, generally support these trends, although levels of endemism were substantially higher for this fauna than had been predicted on the basis of Ltvi's (1979) preliminary analysis of the species composition. These data probably reflect the high diversity of Poecilosclerida in shallow waters. Thirty 1298 J. N. A. Hooper and C . Ltvi species o f Myxillidae, Crellidae, Desmacellidae, Microcionidae and Raspailiidae are now known from the New Caledonian region, including 13 new species and six new locality records for the fauna provided here. Eighteen o f these species (60%)are apparently endemic to this fauna: seven species are restricted to the deeper-water fauna ( o f which 5 are endemic), whereas 21 are predominantly or exclusively shallow-water species ( o f which 13 are endemic). The non-endemic, deeper-water sponge fauna shows similarities to the northern New Zealand sponge fauna, whereas the non-endemic species found in the shallowwater fauna o f New Caledonia also occur in both the temperate (Peronian and Maugean provinces) and tropical Australian faunas (Solanderian province). In most cases, shallowwater species living in both New Caledonian and Australian waters represent the easternmost extent o f species' distributions in the Indo-west Pacific, and, accordingly, many o f these species were substantially different in some o f their characters. Furthermore, many o f the endemic New Caledonian species were also usually immediately recognisable as (transformed) sister-species o f Australasian species. More detailed biogeographic affinitieso f this fauna will be discussed in a separate contribution. A summary o f the New Caledonian poecilosclerid fauna (for the five families indicated) is presented below. S , predominantly or exclusively shallow-water species (0-100 m depth); D, predominantly or exclusively deeper-water shelf species (100-500 m depth); species marked with an asterisk are presently thought to be endemic to the region. Myxillidae (only 3 species have been described, all 'apparently endemic'): Lissodendoryx bifacialis LCvi & LCvi, 1983b [Dl* L. stylophora LCvi & LCvi, 1983b [Dl* Acarnus caledoniensis, sp. nov. [S]* Desmacellidae (only 1 'endemic' species has been described; the hypercalcified 'sclerosponge' Merlia is also included here, after van Soest (1984b) and others]: Neofibularia hartmani, sp. nov. [S]* Merlia normani Kirkpatrick, 1908 [S] Crellidae (only 2 species described, 1 'apparently endemic'): Yvesia acanthosclera LCvi & LCvi, 1983b [Dl* Crella spinulata (Hentschel, 191 1 ) [ S ] Microcionidae (14 species are known, 10 o f which are 'apparently endemic'): Antho novizelanica (Ridley, 1881) [Dl Clathria (Clathriopsamma) rugosa, sp. nov. [S]* C. (CI.) litos, sp. nov. [SI* C. (Thalysias) vulpina (Lamarck, 18 14) [S] C. (T.) flabellifera, sp. nov. [SI* C. (T.) corneolia, sp. nov. [S]* C. (T.) araiosa, sp. nov. [S]* C. (T.) hirsuta, sp. nov. [S] C. (Clathria) kylista, sp. nov. [S]* C. (C.) bulbosa, sp. nov. [S]* C. (C.) menoui, sp. nov. [S]* Echinochalina (Echinochalina) intermedia (Whitelegge, 1902) [S] E. (Protophlitaspongia) laboutei, sp. nov. [SI* E. (P.) bargibanti, sp. nov. [S]* Raspailiidae (7 species have been described, 3 o f which are 'apparently endemic'): Raspailia wilkinsoni Hooper, 1991b [S] Ceratopsion clavata Thiele, 1898 [S] C. palmata Hooper, 1991b [ S ] Aulospongus clathrioides LCvi, 1967 [S]* A . gardineri (Dendy) [Dl Plocamione pachysclera (LCvi & Lkvi) [Dl* Lithoplocamia dolichosclera LCvi & LCvi [Dl* New Caledonian Poecilosclerids 1299 Acknowledgements This paper benefited greatly from the comments of an anonymous reviewer. 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 Barbigant, Jean-Louis Menou, and other staff of ORSTOM, Noumea, for the hospitality, helpful assistance and cooperation during this project. References Bakus, G . J. (1966). Marine poeciloscleridan sponges of the San Juan Archipelago, Washington. Journal of Zoology, London 149, 415-531. Bergquist, P. R. (1965). The sponges of Micronesia, Part 1. The Palau Archipelago. Pacific Science 19(2), 123-204. Bergquist, P. R., and Fromont, J. (1988). The marine fauna of New Zealand: Porifera, Demospongiae, Part 4 (Poecilosclerida). 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