Cheilostomatous Bryozoa from Vanuatu

Transcription

1 Zoological Journal of the Linnean Society (2001), 131: With 22 figures doi: 10.l006/zjls , available online at httpj/ on 10 E Cheilostomatous Bryozoa from Vanuatu KEVIN J. TILBROOK FLS~*, P. J. HAYWARD FLS ~ AND D. P. GORDON FLS3 Marine and Environmental Reseamh Group, School of Biological Sciences, University of Wales, Swansea, Singleton Park, Swansea SA2 8PP Reseamh Associate, Department of Zoology, The Natural History Museum, London SW7 5BD JNational Institute of Water and Atmospheric Reseamh, PO. Box , Kilbirnie, Wellington, New Zealand Received October 1999; accepted for publication February 2000 The present account of some Bryozoa from Vanuatu is the first study of reef-flat species from the eastern Coral Sea. Based on samples collected from several shallow water localities on the island of Efate, a total of 92 species is described, including a new family, three new genera and 20 new species. Some of the newly recognized species are the result of taxonomic revision stimulated by these new samples, while others are entirely new to science. Of the new taxa described here, 16 are presently known only from Vanuatu. The total number of species recorded does not accurately represent the true taxonomic diversity of the Cheilostomatida of Vanuatu, and further sampling, at other localities and in other habitats, is needed before it can be reasonably estimated. The fauna is predominantly Indo-West Pacific in character with a high proportion of species apparently known only in the Southwest Pacific The Linnean Society of London ADDITIONAL KEYWORDS: Bryozoa - taxonomy - systematics - Vanuatu - Southwest Pacific - tropical - new species - faunal review. INTRODUCTION The bryozoan faunas of the tropical southwestern Pacific region are probably the most taxonomically diverse in the world, but are still relatively poorly known for much of the region and for many depths, including the intertidal. In particular, their most accessible component, the shallow-water, coral reef-associated communities, have been especially neglected. For a long time, the most comprehensive accounts of the Bryozoa of the Indo-West Pacific region was contained in the four volumes of the Siboga Expedition Reports (Harmer, 1915,1926,1934,1957) and a volume on the bryozoans of the Philippines (Canu & Bassler, 1929), but these described mostly benthic faunas and scarcely referred to reef-associated species. Winston s (1986) global checklist of coral reef-associated bryozoans tallied just 284 species, only a moderate proportion of which had been reported from tropical Australia and its adjoining areas, while Ryland & Iiayward (1992) * Corresponding author. Present address: Dept. Zoology, The National History Museum, Cromwell Rd., South Kensington, London SW7 5BD. k.tilbrook@nhm.ac.uk ~0llol $35.00/0 35 provided a bibliography of just 13 published sources referring in whole or in part to the Bryozoa of the Great Barrier Reef from 1852 onwards. The potential systematic diversity of the Great Barrier Reef province was demonstrated in two studies of the Bryozoa from Heron Island reef-flat habitats (Ryland & Hayward, 1992; Hayward & Ryland, 1995) which together reported 124 species from 54 samples, including 34 species new to science and 32 species that had not previously been reported from Australia. While the bryozoan faunas of the Great Barrier Reef have at last begun to be explored, those of the islands fringing the eastern Coral Sea, from Papua New Guinea to New Caledonia, and those of eastern Polynesia are very little known (d Hondt, 1985; Gordon, 1989a). There is just a single faunal study, viz. Gordon s (1989a) account of 10 species from Western Samoa, and isolated references to Southwest Pacific localities in systematic reviews of certain tropical genera (e.g. Soule & Soule, 1973; Tilbrook, 1998, 1999, companion paper). The recent MUSORSTOM campaigns around New Caledonia have resulted in an extraordinary yield of new taxa (d Hondt, 1986; Gordon & d Hondt, 1991, 1997; d Hondt & Gordon, 1996, 1999; Gordon, 1993a; The Linnean Society of London

2 36 K. J. TILBROOK ET AL. EFATE (--yguna * Espiritu Santo Pentecost &fate Erromango Q Q Tannab Anatorn 0 Figure 1. Map of Vanuatu. Efate is approximately 45 km across at its widest point. Gordon & Braga, 1994), but again, with the exception of dhondt (1986) these were mostly from deep benthic communities. The present account of some Bryozoa from Vanuatu is the first study of reef-flat species from the eastern Coral Sea. It is based on samples collected in 1990 and 1992 from several localities on the island of Efate (16.44"S, "E). A total of 92 species is described, including a new family, three new genera and 20 new species. Some of the newly recognized species are the result of taxonomic revision stimulated by these new samples, while others are entirely new to science. MATERIALS The collections forming the basis of this study were made during two periods, October 1990 (D. P. Gordon) and November 1992 (l? J. Hayward), at four shallow water locations on the island of Efate (Fig. 1); Iririki Island, off Port Vila, Port Vila itself, Erakor Island (so called as it is opposite Erakor village at the seaward edge of Ekasuvat (Erakor) Lagoon, though not named on local maps) and Poanangisu, in the north of the island. All material was returned dried and examined using a WILD light microscope after initial preparation with 5-10% bleach solution. Scanning electron micrographs were taken at the Natural History Museum using an IS1 ABT-55 scanning electron microscope with an environmental chamber. The images presented here are back scattered electron images of uncoated specimens (for index of species described in paper, see Appendix). SYSTEMATICS The taxonomic order is based on that adopted by Ryland & Hayward (1992), Hayward & Ryland (1995) as advocated by Gordon (1984, 1986, 1989a,b,c). All material is deposited in the Department of Zoology, The Natural History Museum, London. SUPERFAMILY AETEOIDEA SMITT, 1867 FAMILY AETEIDAE SMI'IT, 1867 GENUS AETEA LAMOUROUX, 1812 Type species: Sertularia anguina Linnaeus, There are relatively few known species of Aetea, but identifying them with certainty is not straightforward as the commonest, presently recognized species are either highly variable or complexes of more than one species (Gordon, 1984). Aetea species often produce

3 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 37 extended colonies which are very small and fragile, usually occur in cryptic habitats and are thus extremely difficult to identify. Aetea anguina (Linnaeus, 1758) has been accorded ii worldwide distribution in shallow shelf seas, with the exception of polar waters, but it is probable that this encompasses several distinct species. Specimens attributable to A. anguina have been reported from Heron Island (Ryland & Hayward, 1992) but as with the present material they were mostly too incomplete for adequate comparison with European specimens. Many fragmentary colonies of Aetea species were found in samples of coral rubble from Erakor Island and Port Vila Harbour, some of which seem attributable to the following taxa. AETEA?A USTRALIS FIDE GORDON (Fig. 2A,B) Aetea?australis Gordon, 1984: 39, pl. 8, figs A-D. Material NHM , Erakor Island, Efate, Vanuatu. Actea?australis is characterized by its very fine, closely packed, annulated erect mid-section, which merges into a punctate distal portion. The material figured here agrees well with that illustrated by Gordon (1984) as Aetea?australis Jullien, As Gordon (1984, 1986) discussed, his New Zcaland material did not correspond to European specimens of A. anguina (Linnaeus, 1758) or A. sica (Couch, 1H44), hence its nominal assignment to Jullien s (1888) species, which was described from magellanic South America and western Australia. Numerous small colonies were found in interstices of coral rubble at Erakor Island. AETEA LIGULATA BUSK (Fig. 2A) Avtea ligulata Busk, 1852b: 31, pl. 42. At>tea ligulata: Prenant & Bobin, 1966: 89, fig. 21, IV, VI (cum syn.); Gordon, 1984: 39, pl. 8, figs E,F. Mu terial NHM , Erakor Island, Efate, Vanuatu. Rcmarks Actea ligulata is characterized by its broad, rounded, horizontal corrugations, rather than the smaller annulations seen in A.?australis (above), on the erect midsection. It is the larger of the two species found at Erakor Island during this study. Described by Busk (1852b) from the Straits of Magellan and Patagonia, this species has been accorded a circumglobal distribution (Gordon, 1984). SUPERFAMILY MEMBRANIPOROIDEA BUSK, 1852b FAMILY MEMBRANIP~RIDAE BUSK, 1852b GENUS JELLYELLA TAYLOR & MONKS, 1997 Diagnosis Colony an encrusting unilaminar sheet. Autozooids lightly calcified, frontal surface largely membranous, gymnocystal calcification moderate to well developed, producing tubercles or longer processes proximally and around opesia; cryptocystal calcification wanting or reduced; opesia bordered by small branched marginal spinules. No articulated spines, avicularia, or ovicells. Planktotrophic cyphonautes larva, which gives rise to a twinned ancestrula. m e species: Membranipora eburnea Hincks, Taylor & Monks (1997) erected the genus Jellyella for two species, Membranipora eburnea Hincks, 1891 and Flustra tuberculata Bosc, 1802, epiphytic on Sargassum. They characterized the genus by skeletal characters (the presence of intricately branched spinules projecting into the zooidal chamber, and a calcitic ultrastructure of transversely arranged elongate needles, more or less parallel to the wall surfaces), lack of lateral wall cracks and tower cells found in Membranipora sensu stricto, and the presence of a twinned ancestrula. JELLYELLA TUBERCVLATA (BOSC) (Fig. 20) Flustra tuberculata Bosc, 1802: 143. Nichtina tuberculata: Harmer, 1926: 208, pl. 13, fig. 10. Membranipora tuberculata: Osburn, 1950: 23, pl. 2, figs 4-6 (cum syn.); Cook, 1985: 82, pl. 3A (cum syn.); Hayward & Ryland, 1995a: 537, fig. 3C. Jellyella tuberculata: Taylor & Monks, 1997: 41, figs 3,14,15. The species found in Vanuatu seems to agree well with the figures of Winston (1982) and Hayward & Ryland (1995a). Autozooids are x mm, and gymnocystal tubercles are clearly evident in frontal proximal angles. The cryptocyst is minutely tuberculate, deeper proximally and many autozooids have a single marginal spinule on the inner edge of the

4 38 K. J. TILBROOK ET AL. opesia. Generally, there are two distolateral and two proximolateral septula on each lateral wall. The distal wall, which appears as a slight shelf, bears up to ten uniporous septula close to the basal wall. The type material of Membranipora tenella Hincks, 1880b, on Sargassum from Florida, is very similar to Jellyella tuberculata. However Hincks specimen has a deeper proximal gymnocyst and the tubercles are far closer together as a consequence, i.e. not in the proximal angles. The species illustrated by Winston (1982) as Membranipora tenella Hincks, is not Hincks species, rather it is more similar to Crassimarginatella maxillaria sp. nov. (see below). This species has been quoted as being found wherever Sargassum drifts over the warmer seas...= (Osburn, 1950). Originally described from the Atlantic, this species has been recorded from warm temperate and tropical seas worldwide. However, many variations have been noted in morphology both within and between geographic areas (Harmer, 1926; Osburn, 1950) which perhaps indicates a complex of species. It was found encrusting algae (of indeterminable assignment) from Port Vila Harbour, Efate. Diagnosis GENUS BIFLUSTRA D ORBIGNY, 1852 Colony encrusting, or erect, unilaminar. Autozooids variably calcified, the frontal surface membranous, with no gymnocystal calcification; cryptocyst well developed proximally, forming a shelf, generally granular. No spines, avicularia or ovicells. Type species: Membranipora ramosa d orbigny, he form of the ancestrula is not known for Membranipora ramosa dorbigny, 1852, a feature that has been used to differentiate genera, for example Jellyella Taylor & Monks, Flustra savartii Audouin, 1826 (variously assigned to Biflustra, Mem branipora de Blainville, 1830 and Acanthodesia Canu & Bassler, 1920) which has a twinned ancestrula (P.D. Taylor pers. comm.) is not conspecific with d orbigny s species and might not even be congeneric with it. If after further investigation the latter is found to be true then there may be some justification for resurrecting Acanthodesia for l? savartii and allied species with demonstrably twinned ancestrulae. BZFLUSTRA RETICULATA SP. NOV. (Fig. 2E) Material Holotype: NHM , Port Vila Harbour, Efate, Vanuatu, October Paratypes: NHM , same locality as holotype, October 1990; NHM , Erakor Lagoon, by the Radisson Royal Palm Resort, Efate, October Colony encrusting, unilaminar. Autozooids irregularly oval to rectangular, distinctly separated by shallow grooves, quincuncially arranged in ordered columns; lateral walls shallow. Gymnocyst reduced, most prominent proximally as two triangular-shaped areas, either side of the distal end of the preceding zooid; a thin wall of gymnocyst surrounding the entire cryptocyst, with a slightly raised tuberculate rim. Cryptocyst occupying one third to one half frontal area, extending laterally either side of the opesia, negligible distally. most extensive proximally, flattened or very slightly convex, smooth other than for a raised reticulate pattern of tubercles; rim of cryptocyst smooth proximally, slightly tuberculate laterally. Opesia oval or subtriangular. Generally four large multiporous septula in lateral walls, two distolateral and two proximolateral. A line of up to ten very small uniporousl septula in the distal wall. Basal wall generally entire, but appears to be lacking in small areas in certain autozooids. Measurements Holotype: means and standard deviations, mm (n= 30). Autozooid length 0.56k0.05; width 0.27 ko.02. Opesia length 0.28 k0.03; width 0.23 k0.02. Etymology From reticulatus, L.-netlike. Named for the reticulate patterning of the frontal cryptocyst. The most characteristic feature of BifEustra reticulata is the reticulate patterning on the proximal area of cryptocyst. The suite of septula observed in this species is similar to those of Jellyella tuberculata described above. The form of the ancestrula has not been observed. Two small colonies of Biflustra reticulata were found in Port Vila Harbour encrusting discarded wall tiles,

5 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 39 Figure 2. A, Aetea ligulata (centre) and Aetea?australis (right). B, Aetea?australis. C, Nellia tenuis. D, Jellyella tiiberculata. E, Biflmtra reticulata sp. nov. F. Conopeurn papillorurn sp. nov. G, Antrwpora granulifera. Note colony origin, bottom right. Scale bars: A = 0.2 mm; B = 0.10 mm; C = 0.25 mm; D = 0.25 mm; E = 0.25 mm; F = 0.50 mm; G = 0.25 mm.

6 40 K. J. TILBROOK ET AL. and another on a specimen of Halimeda sp. from Erakor Lagoon. FAMILY ELECTRIDAE STACH, 1937 GENUS CONOPEUM GMY, 1848 Type species: Flustra lacmixii Audouin, CONOPEUM PAPZLLORUM SP. NOV. (Fig. 2F) Material Holotype: NHM , Vila Waterfront, Efate, Vanuatu. Colony encrusting, unilaminar. Autozooids elongateoval, rectangular or polygonal, distinct, separated by deep grooves, frontal surface largely membranous; opesia oval, occupying majority of frontal area. Cryptocyst tuberculate, narrow, surrounding whole opesia, widening very little proximally, many small spinules around the inner edge, orientated towards the interior of the opesia; gymnocyst smooth, well developed, surrounding entire cryptocyst, deepest proximally where generally two conical bud-like processes are produced. Three or four pairs of lateral marginal spines, short, stout, pointed, incurved over the opesia. Space-filling kenozooids present, small, with a circular opesia surrounded by cryptocyst similar to autozooids but with cryptocystal spinules on the inner rim; no spines. Measu pem ents Holotype: means and standard deviations, mm (n= 30). Autozooid length 0.41 ko.08; width 0.29 f0.03. Etymology From papilla, L.-buds. Named for the two bud-like processes on the proximal gymnocyst of each autozooid. Conopeurn papillorum is characterized by the proximal gymnocystal processes, marginal spines and cryptocystal spinules. This species is very similar to Electra angulata Levinsen, 1909, originally described from Koh Samet, Thailand, and subsequently recorded by Harmer (1926) from North Sumbawa, Malaysia. Although E. angulata has a well-developed proximal gymnocyst and two budlike processes, these are very small and thin; there are also six to eight pairs of marginal spines, as well as a single proximomedial spine near the edge of the gymnocyst/cryptocyst boundary, unlike the rest of the spines which are at the junction, though originating on the gymnocyst. Mawatari (1974) synonymized Electra angulata Levinsen with Membranipora tenella Hincks (as Electra tenella), but with some reservations. Mawatari s (1974) text-figures and plates are far more reminiscent of E. angulata as described and illustrated both by Levinsen (1909) and Harmer (1926) than of the type specimen of Hincks M. tenella (see discussion of Jellyella tuberculata above) for several reasons. Firstly, M. tenella sensu stricto bears no marginal spines and, in particular, no single proximomedial spine (as described above), but also the gymnocystal processes are far more robust and knob-like, occupying a greater area of gymnocyst, thanineithere. angulata or C. papillorum. Membranipora tenella and E. angulata are clearly separate species; the latter is the species recorded by Mawatari (1974) and whilst the two species each bear a superficial resemblance to C. papillorum it is distinct from both. Conopeum papillorum differs greatly from the two species of Conopeum described by Harmer (1926) from Malaysia-C. eriophorum (Lamouroux, 1816) and C. reticulum (Linnaeus, 1767). Having examined Harmer s material of each (C. eriophorum NHM , New Guinea; C. reticulum NHM , Kei Islands), it is obvious that they are conspecific, although the actual species identity remains uncertain: there is no visible gymnocyst, therefore no gymnocystal processes or marginal spines, and they may represent an undescribed species. A single colony of Conopeum papillorum was found encrusting a shell from Vila Waterfront, Efate, itself having been grown over by a colony of Watersipora subovoidea sensu Harmer, SUPERFAMILY CALLOPOROIDEA NORMAN, 1903a FAMILY ANTROWRIDAE VIGNEAUX, 1949 GENUS ANTROPORA NORMAN, 1903b Diagnosis Colony encrusting, unilaminar or multilaminar. Autozooidal cryptocyst moderately developed, gymnocyst negligible or absent. Spines absent. Small interzooidal avicularia and autozooid-sized vicarious avicularia present. Ovicells endozooidal, their presence generally indicated by a slight cap-like thickening at the distal end of the autozooid. Basal pore-chambers (dietellae) present. Type species: Membranipora granulifera Hincks, 1880a.

7 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 41 I ilbrook (1998) provided a comprehensive re-evaluation of the species assigned to this genus. ANTROPORA GRANULIFERA (HINCKS) (Fig. 2G) Membranipora granulifera Hincks, 1880a: 72, pl. 9, fig. 4. Antmpora granulifera: Norman, 1903b: 87, pl. 8, fig. 4; Tilbrook, 1998: 27, fig. 1A. Colony forming flat unilaminar sheets. Autozooids irregularly polygonal to hexagonal, separated by discernible grooves. Frontal surface bordered by a crenulated mural rim, raised particularly towards the distal end of the autozooid. Gymnocyst proximal, very reduced or negligible; cryptocyst occupying generally less than one half of total autozooid length, flat or very slightly convex, coarsely beaded. Opesia roughly triangular to slightly trifoliate, constricted at the proximal edge of the operculum opening. Distal to each autozooid a pair of medium-sized, triangular intcrzooidal avicularia. Rostra raised and acute to frontal plane, directed medially, often touching at the midline, or rarely very slightly distomedially, particularly in ovicellate zooids; mandibles long and acutely triangular. Vicarious avicularia absent. Ovicell endozooidal, small, indistinct. Mature colonies are chocolate brown in appearance, lighter and more translucent at the growing edge. The opercula are slightly darker than the rest of the frontal membrane. Cook (1968) described the opercula of fertile zooids as showing a slight dimorphism, i.e. wider and darker in colour than other autozooids. Antmpora granulifera was found at Port Vila Harbour and at Iririki Island. This species has a circumglobal distribution in shallow warm-temperate and tropical waters. ANTROPORA MlNOR (HINCKS) (Fig. 3A) Membranipora trifolium var. minor Hincks, 1880a: 87, pl. 11, fig. 6. Antmpora minus: Cook, 1968: 139, fig. 10; Mawatari & Mawatari, 1981: 34. Antmpora minor: Tilbrook, 1998: 34, fig. 2A-F (cum Syn.). Colony encrusting, multilaminar. Autozooids oval, bordered by a distinct thin mural rim, separated by deep grooves, often wide. Gymnocyst generally minimal; cryptocyst coarsely granular, concave, deeper proximally than laterally, sloping basally. Opesia large, broadly oval or subtriangular, narrowing distally, occupying over half of the frontal area. Small interzooidal avicularia, elongate, rounded, mostly distally directed; mandible rounded or more acute, triangular; no crossbar; a cryptocystal rim around avicularium opesia. Small kenozooidal papillae are also often found at the proximal ends of autozooids in the interzooidal angles, either in place of or in conjunction with the avicularia. Ovicells endozooidal, vestigial, indistinct, cap-like. Vicarious avicularia often present. Antmpora minor has a circumtropical distribution in shelf seas, which only became apparent following the recent revision of the genus by Tilbrook (1998), who provided a complete synonymy for this species. It was not common at Vanuatu, with only eight colonies being found in total, mainly from Erakor Island; others occurred in Port Vila Harbour and at Poanangisu on the north coast of Efate. Diagnosis GENUS PARANTROPORA TILBROOK, 1998 Colony encrusting. Autozooidal cryptocyst moderately developed around the opesia, gymnocyst negligible or absent. Spines absent. Small interzooidal avicularia present. Large spatulate vicarious avicularia present, much larger than autozooids. Ovicells endozooidal. Septula present. Type species: Parantmpora penelope Tilbrook, This genus is very similar to Antmpora, from which it differs in the lack of dietellae, presence of lateral-wall septula, and the occurrence of very large spatulate vicarious avicularia. The presence of special zooecia was first noted by Canu & Bassler (1929) in Membrendoecium lagunculum, though they failed to recognize them as avicularia. Species of Parantmpora generally have a more delicate structure to their side walls than do species of Antmpora. PARANTROPORA PENELOPE TILBROOK (Fig. 3F) Antmpora granulifera: Ryland & Hayward, 1992: 229, fig. 2c. Parantmpora penelope Tilbrook, 1998: 40, fig. 3D,E.

8 42 K. J. TILBROOK ET AL. Colony encrusting, unilaminar. Autozooids with minimal gymnocyst; cryptocyst occupying slightly less than half total length, flat or slightly concave, coarsely beaded; opesia subtriangular. At the distal end of each autozooid a pair (rarely one) of small, teardrop-shaped interzooidal avicularia; rostrum acute to frontal plane, directed distally or distomedially, mandible acutely triangular. Large vicarious avicularia also present, generally larger than autozooids; rostrum raised distally, cryptocyst narrow, coarsely granular, laterally constricted; opesia rounded; mandible spatulate, articulated on pointed, triangular condyles, angled slightly proximally, situated proximal to lateral rostral constriction. Ovicells endozooidal, small, granular, caplike, derived from distal zooid. Following examination of Ryland & Haywards (1992) material from Heron Island, attributed to Antmpora granuliferu (Hincks, 1880), it may confidently be assigned to I! penelope. Great Barrier Reef, Australia; Erakor Island, Vanuatu; Suva Barrier Reef, Fiji. This species appears to be limited to the tropical Southwest Pacific. FAMILY CALLOFORIDAE CANU & BASSLER, 1927 GENUS CORSULELLA GORDON, 1984 Qpe species: Membranipora corbula Hincks, 1880b. Corbulella was originally introduced as a subgenus of Crassimarginatella by Gordon (1984) together with Crassimalginatella and Valdemunitella. Owing to the long ranges of these subgenera in the Cenozoic, and the number of species in each, it is appropriate that they should be regarded as full genera. CORSULELLA CORBULA (HINCKS) (Fig. 3B) Membranipora corbula Hincks, 1880b: 378, pl. 17, fig. 6. Crassimarginatella corbula: Ristedt & Hillmer, 1985: 136, pl. 1, fig. 4. Crassimarginatella (Corbulella) corbula: Gordon, 1984: 29, pl. 3, figs D,E (cum syn.); Gordon, 1986: 32, pl. 5, fig. A; Ryland & Hayward, 1992: 230, fig. 3. Colony encrusting, forming small unilaminar sheets. Autozooids oval, distinct; x mm. Oval opesia occupying most of frontal area, bordered by a narrow granular cryptocyst; gymnocyst smooth, well developed, particularly proximally. Six to nine pairs of marginal spines, distal one or two pairs stout, longer and erect, the rest slender, incurved over opesia. Vicarious avicularia as long as autozooids but slightly wider, without pivot bar; oval opesia constricted midway by opposing pivots, bordered proximally by spines; narrow cryptocyst and smooth, variably developed, proximal gymnocyst; thin raised rostral rim, minutely serrated. Ovicell prominent, hyperstomial, with a broad frontal fenestra, notched apically. Kenozooids occasional. Corbulella corbula seems to be widely distributed in the western Pacific: Japan, Great Barrier Reef, and Kermadec Ridge and Tasman Bay, New Zealand. It was quite common in cryptic habitats on coral rubble from the reef flats of Erakor Island, Efate. GENUS CRASSMARGINATELLA CANU, 1900 %e species: Men branipora crassimarginata Hincks, 1880a. See discussion in Gordon (1984) of species assigned to this genus. CRASSKMARGKNATELLA CORNICULATA SP. NOV. (Fig. 3C) Material Holotype: NHM , Poanangisu, Efate, Vanuatu. Paratypes: NHM ,189, same locality as holotype. Colony encrusting, forming unilaminar sheets. Autozooids irregularly oval, separated by distinct deep grooves, in quincuncial arrangement. Gymnocyst distinct, smooth, particularly prominent proximally, narrowing laterally; cryptocyst narrow, granular, with beaded opesial border. Opesia oval, covering most of the frontal area. Single pair of distal gymnocystal protuberances (not articulated spines) present, short (protruding above the gymnocyst only slightly), laterally flattened, sometimes almost bifurcate. Avicularia not observed. Ovicells endozooidal, vestigial,

9

10 44 K. J. TILBROOK ET AL. distal edge of ovicellate zooids only slightly raised. A single distal and two lateral mural septula. Measurements Holotype: means and standard deviations, mm (n= 25). Autozooid length 0.58 k0.04; width 0.38 k0.03. Etymology From corniculatus, L.-horned. Named after the appearance of the distal gymnocystal protuberances. Crassimalginatella corniculata is very similar to Crassimarginatella similis and C. falcata described by Cook (1968) from West Africa. However, C. corniculata differs from these species in length of gymnocystal protuberances and extent of gymnocyst, and number of protuberances and extent of gymnocyst, respectively. Crassimarginatella corniculata is also similar to C. tubemsa Cook, 1968 from West Africa and C. spatulifera Harmer, 1926 from Indonesia: neither of the latter species bears marginal spines, which are also lacking in C. corniculata, and both bear gymnocystal protuberances similar to those of C. corniculata, but they are far more sporadic in these species than in C. corniculata in which generally all autozooids bear two. However, C. tubemsa and C. spatulifera possess autozooid-sized vicarious avicularia, spatulate in C. tubemsa and subquadrate with a serrated rostrum in C. spatulifera; none of the specimens of C. corniculata here described bears vicarious avicularia. Finally, the distal calcification, indicating the presence of endozooidal ovicells, is different in all three species. There is evidence of zooidal regeneration in the holotype and in a colony from Port Vila Harbour. Crassimalginatella corniculata was found in cryptic habitats on small pieces of coral rubble at Poanangisu, Erakor Island, and Port Vila Harbour, Efate. CRASSZMARGZNATELLA MAXZLLARZA SP. NOV. (Fig. 30) Material Holotype: NHM , Port Vila Harbour, Efate, Vanuatu. Paratype: NHM , same locality as holotype. Colony encrusting, forming large, flat unilaminar sheets. Autozooids irregularly oval to hexagonal, separated by distinct grooves, in quincuncial arrangement. Cryptocyst very narrow, granular; gymnocyst very narrow, smooth, rounded, deeper proximally. One, often two (sometimes lacking) tubercles (? spine bases) on proximal gymnocystal area. Avicularia vicarious, larger than autozooids ( x mm), distal portion comprising raised, smooth-edged rostrum supporting a spatulate mandible; proximal portion consisting of a shortened opesia. No crossbar. Ovicells not observed. Measurements Holotype: means and standard deviations, mm (n= 30). Autozooid length 0.66 k0.07; width 0.37 k0.04. Etymology From maxillaris, L.-of the jaw. Alluding to the appearance of the vicarious avicularia, the distal portions of which are surrounded by the raised spine bases (?) of the two distal zooids. The vicarious avicularia occur at the division of autozooid rows. The two distal autozooids produce several, generally three or four, spine bases along the edges of the gymnocyst next to the distal end of the avicularium, giving the appearance of a jaw. The spine bases appear similar to those illustrated by Gordon (1984) for his new species Crassimarginatella (Crassimarginatella) electra, though no evidence of smaller lateral spines has been seen in Crassimarginatella maxillaria. The species illustrated by Winston (1982) as Membranipora tenella Hincks, 1880b, is not Hincks species, being more similar to C. maxillaria; however. there is no evidence of vicarious avicularia in either the description or figure and the gymnocyst in Winston s figure is far more reduced than that seen in C. maxillaria. Only one large broken colony of Crassimalginatella maxillaria was found, on a discarded wall tile from Port Vila Harbour, Efate. GENUS CRANOSINA CANU & BASSLER, 1933 Type species: Membranipora comnata Hincks, 1881a.

11 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 45 CRANOSINA CORONATA (HINCKS) (Fig. 3E) Membranipora comnata Hincks, 1881a: 147, pl. 10, fig. 1. Setosellina comnata: Harmer, 1926: 265, pl. 16, figs 24. Cranosina comnata: Winston & Heimberg, 1986: 6, figs 3-6; Hayward, 1988: 281; Ryland & Hayward, 1992: 230, fig. 2e. This species is common throughout the Indo-West Pacific region, from the Philippines and Indonesia to Sri Lanka and Mauritius. Although not very frequent in the samples from Heron Island, Australia (Ryland & Hayward, 1992) it was very common at Poanangisu, Efate, as the most abundantly occurring species. FAMILY QUADRICELLARIIDAE GORDON, 1984 GENUS NELLIA BUSK, Type species: Nellia oculata Busk, 1852b. NELLIA TENUIS HARMER (Fig. 2C) Nellia tenuis Harmer, 1926: 245, pl. 14, figs 16, 17. Nellia tenuis: Hayward, 1988: 286, pl. 1, fig. a; Gordon, 1989: 450, fig. 3; Ryland & Hayward, 1992: 231, fig. 1oc. This species appears to be common in shallow reefal habitats where it forms part of the bryozoan sward on the undersides of coral rubble; however, its delicate spindly colonies may easily be overlooked. Originally described by Harmer (1926) from localities in Indo-Malaysia and the South China Sea, it has since been reported from the Great Barrier Reef, Fiji, Samoa and Mauritius. In Vanuatu it was a common member of the cryptic bryozoan sward found on the larger pieces of coral rubble, particularly from Erakor Island reef flat. SUPERFAMILY BUGULOIDEA GRAY, 1848 FAMILY BUGULIDAE GRAY, 1848 GENUS BUGULA OKEN, 1815 Type species: Sertularia neritina Linnaeus, BUGULA SCAPHULA SP. NOV. (Fig. 4A,B) Bugula cf. scaphoides Hayward, 1988: 289, fig. 4a. Material Holotype: NHM , Erakor Island reef flat, Efate, Vanuatu, Paratypes: NHM , NHM , 106, NHM ,191, NHM , same locality and collection data as holotype. Other material examined: NHM ,25 (as Bugula cf. scaphoides), Tamarin reef front, Mauritius, 5m, Colony comprising delicate sparse tufts; autozooids in biserial rows, bifurcating infrequently. Bifurcation of Type 3 (Harmer, 1923), zooids E and F forming the axis of the dichotomy and linked at their proximal ends by a slender connecting process. Autozooids (s0.45 x 0.15 mm) with broad distal portion constituting two-thirds of total length, frontal area entirely membranous, outer distal angle acute but not spinose, inner distal angle rounded; proximal portion narrow, cylindrical, fully calcified, with an apparent flexure just proximal to distal portion. Avicularia frequent, apparently attached at the proximal end of each autozooid, but actually borne on a short peduncle that originates from the basal wall of the preceding autozooid; its length almost equivalent to the width of the autozooid opesia; domed, the rostrum pointed and downcurved. Ovicell attached to inner distal angle, globular, pedunculate, the ectooecium appearing reticulate when dried. Etymology From scapha, L.-light boat, skiff. Alluding to the shape of the distal portion of the autozooid. Bugula scaphula from Erakor Island, Vanuatu, is undoubtedly conspecific with that described by Hayward (1988) from Mauritius as Bugula cf. scaphoides Kirkpatrick. Dried colonies appear very thin, translucent, and yellow-orange. The species occurs in cryptic habitats in coral rubble and may easily be overlooked. The present material was fragmentary, as was that from Mauritius (Hayward, 1988), but both have been compared with the type material of B. scaphoides (Kirkpatrick, 1890), which differs from B. scaphula particularly in its very small avicularia. Bugula scaphoides was redescribed by Harmer (1926) who noted its red coloration, small avicularia and white ovicells.

12 46 K. J. TILBROOK ET AL. Bugula scaphula has only been found at Tamarin reef, Mauritius and Erakor Island, Efate where it often cooccurred with Catenicella triangulifera Harmer, GENUS BRE'ITIELLA GORDON, 1984 Diagnosis Colony repent, supported above substratum by rhizoids; uniserial, branching. Autozooids claviform, with oval opesia and tubular proximal portion. No operculum. Spines and avicularia lacking. Ovicell globular, prominent. Pore-chambers present, with relatively large-pored septula. Type species: Brettiella ovicellata Gordon, BREZTZELLA CULMOSA SP. NOV. (Fig. 5B,C) Material Holotype: NHM , Erakor Island reef flat, Efate, Vanuatu, Paratypes: NHM , NHM , 274, NHM , same locality and collection data as holotype. Colony repent; each autozooid supported above the substratum by a rhizoid produced from its basal surface; translucent straw yellow when dried. Autozooids large (Holotype, x mm), claviform, widening distally, with extensive oval opesia and frontal membrane; proximally narrow and tubular. Autozooids arise distobasally from preceding zooid. Branches originate either left or right of the main axis, the most proximal zooid of a new branch arises laterobasally towards the distal end of the parent zooid. No ovicells or ancestrula observed. Etymology From culmosus, L.-strawlike. when dried. Alluding to the colour Brettiella culmosa differs from Brettiella ovicellata Gordon, 1984, the only other species in this genus, mainly in its size; autozooids of B. culmosa are almost twice as long as those of B. ovicellata whilst having a similar width. Additionally, the proximal end of the opesia in B. culmosa is bounded by a U-shaped border of cryptocyst, while the proximal edge of the gymnocyst is distinctly V-shaped. Brettiella culmosa was found only at Erakor Island but, owing to its inconspicuous appearance and cryptic habitat, it is probable that this species has been overlooked elsewhere and will be recorded from a broader geographical area. As well as loose fragments, a single colony was found in a cryptic area of a small coral boulder, associated with Beania hexamicomm. FAMILY BEANIIDAE CANU & BASSLER, 1927 GENUS BEANIA JOHNSTON, 1840 Type species: Beania mirabilis Johnston, BEANIA KLUGEI COOK Beania khgei Cook, 1968: 164, fig. 2A,B. Beania klugei: Cook, 1985: 119. Not Beania klugei: Hayward & Ryland, 1995a: 542, fig. 6A,B. Colony forming diffuse, branching, uniserial chains of slender autozooids. Autozooids almost parallel-sided, strongly tapered proximally; frontal surface entirely membranous. Spines lacking except for a pair of short, pointed distal processes. Each autozooid bears a small pair of frontally facing pedunculate avicularia, laterally adjacent to the operculum; rostrum far longer than deep; mandible acutely triangular, tip hooked. Autozooids budded from a single distobasal septulum and from paired proximolateral septula. No ovicells observed. Cook (1968) first distinguished this species from B. intermedia (Hincks, 1881b), from which it differed in having larger autozooids, comparatively smaller slender avicularia, and no marginal spines. Beania klugei was shown to be distributed from the Indian Ocean and the Red Sea to West Africa, the Caribbean, and Panama, while B. intermedia occurred in the Southwest Pacific. Bugula klugei of Hayward & Ryland (1995a) from Heron Island, Great Barrier Reef, had large autozooids and lacked marginal spines, but its avicularia are more similar to those of B. intermedin, with a highly domed profile. Their material is here referred to Beania cookae sp. nov. (see below) which occurred abundantly at two of the sites sampled in Vanuatu. Cook (1985) discussed the possibility that R. klugei may prove to be a junior synonym of Bugulella clavata Hincks, 1887, described from the Mergui Archipelago and subsequently reported and described by Menon & Nair (1969) from the Gulf of Manaar. Hincks' species lacks a type specimen and so this possibility

13

14 48 K. J. TILBROOK ET AL. cannot be further examined. The few autozooids found in Vanuatu are very similar in size (length approx. 1.00mm) to those described by Cook (1968), as are their avicularia ( x mm). Beania klugei was originally described from West Africa, but was shown to have a tropical/subtropical distribution from the Red Sea and Zanzibar to the Caribbean. Its putative first record from Australia (Hayward & Ryland, 1995) is here rejected, but its presence in the Southwest Pacific is established by the fragmentary specimens from Erakor Island. BEANIA PETIOLATA HARMER (Fig. 4E, F) Beania petiolata Harmer, 1926: 416, pl. 28, figs 18, 19. Material Holotype: NHM , South of Halmaheira (Djilolo), Indonesia, 45 m. Other material examined: NHM , west of north end of New Guinea, 32 m; NHM , NHM , Erakor Island reef flat, Efate, Vanuatu. Colony reticulate, forming thin sheets loosely attached to the substratum; autozooids equally spaced and connected by six relatively short tubes to its neighbours. Each autozooid has a single, tubular basal pore proximal to the most distal connecting tube; frontal wall entirely membranous. Autozooids have six short, straight, distal marginal spines, four on the distal margin and one bifurcating spine adjacent to each distally placed avicularium; there are two longer distally directed basal spines, and five to six pairs of generally equally spaced, longer, bifurcating lateral marginal spines, each with one ramus incurved over frontal membrane, the other ramus at approximately 45" to the first, directed frontolaterally. The paired frontally facing avicularia lateral to the operculum are attached just basal to the frontal rim; short with a squared head; rostrum slightly hooked; mandible triangular. No ovicells observed. Measurements Means and standard deviations, mm. Holotype (n = 18): Autozooid length 0.64 k0.05; width 0.29 f NHM (n = 30): Autozooid length 0.75f0.13; width NHM (n = 30): Autozooid length 0.59 k0.03; width 0.28 k0.02. Avicularium length (n = 15): 0.08 _ As noted by Harmer (1926), the presence of distal basal spines, branching lateral marginal spines, and the shape and size of the avicularia are the most distinctive characters of this species. He gave no measurements with his description of this species, but part of the type material, from south of Halmaheira (Djilolo), and another of Harmer's specimens, from New Guinea, were measured (above) and appear to have slightly larger autozooids than the specimens from Vanuatu (also above). This slight difference may be a result of preservation, however: Harmer's specimens were mounted in Canada balsam on slides, while the Vanuatu material was dried. The species is easily overlooked in its cryptic habitat on pieces of coral rubble, and is almost transparent when dry; it has very small colonies of fewer than 100 zooids, covering less than 20 mm'. Originally found in Malaysia and New Guinea, this is only the second record of Beania petiolata which was quite common at Erakor Island, Efate. BEANZA COOKAE SP. NOV. (Fig. 5E,G) Beania klugei Hayward & Ryland, 1995a: 542, fig. 6A, B. Material Holotype: NHM , Poanangisu, Efate. Paratypes: NHM ,50, NHM , Erakor Island reef flat, Efate, ; NHM , Port Vila Harbour, Efate; NHM , Paonangisu, Efate. Colony diffuse, uniserial, forming branching chains of slender, almost parallel-sided, autozooids. Frontal surface of autozooids entirely membranous. Spines may be lacking, except for a pair of short, acute distal processes; however, some autozooids may produce one or two further pairs of short lateral spines, one-third or two-thirds of their length proximally along the margin. Each autozooid bears a small pair of frontally facing pedunculate avicularia, attached laterally, adjacent to the operculum; rostrum highly domed, a5 deep as long; mandible acutely triangular. Autozooids are budded from a single distobasal septulum, and

15 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 49 Figure 5. A, Scrupocellaria spatulata. B,C, Brettiella culmosa sp. nov. B, autozooid showing budding pattern and basal postioning of rhizoid. C, close-up of frontal membrane. D, Beania cf mirabilis. E, Beania cookae sp. nov. Autozooid showing budding pattern. F, Beania cf mirabilis. G, Beania cookae sp. nov. Domed avicularia. Scale bars: A = 0.20 mm; B = 0.25 mm; C = 0.10 mm; D = 0.25 mm; E = 0.40 mm; F = 0.50 mm; G = 0.10 mm.

16 50 K. J. TILBROOK ET AL. paired proximolateral septula. No ovicells were observed. Measurements Holotype: means and standard deviations, mm. Autozooid length 1.23 ko.11 (n = 23); avicularium x Etymology Named for bryozoologist Patricia L. Cook. Beania cookae is very similar to Beania klugei Cook, 1968; however, despite being of a comparable size (if not a little larger), it produces occasional lateral marginal spines and has shorter, more-domed avicularia. The avicularia resemble those seen in B. intermedia (Hincks, 1881b), from Tasmania and New Zealand, but this species is only just over half the size of B. cookae. Younger zooids have no marginal spines, just two pairs of distal processes in the form of simple evaginations of the zooidal wall, one pair on the very distal edge and the other pair at the corner of the operculum. It may be that these latter processes are the precursors of the avicularia as they are not seen in zooids bearing avicularia. In later ontogeny two to four marginal spines may be present, proximal to the avicularia. Other specimens of Beania were found at Poanangisu and Erakor Island but they were unidentifiable as either B. klugei or B. cookae. The Heron Island material attributed to Beania klugei by Hayward & Ryland (1995a) bears the short, domed avicularia of B. cookae, and is accordingly assigned here to this species. In Vanuatu, B. cookae was fairly abundant in the material from Poanangisu, Erakor Island, and Vila Harbour, Efate, encrusting cryptic habitats in coral rubble. BEANlA HEXAMZCORUM SP. NOV. (Fig. 4C,D) Beania sp. Hayward, 1988: 288, fig. 4b. Material Holotype: NHM , Erakor Island reef flat, Paratypes: NHM , Erakor Island reef flat, Other material examined NHM (as Beania sp.), Tamarin reef front, Mauritius, 5 m; NHM (as Beania sp.), Flic en Flac, Mauritius, 25 m. Colony reticulate, forming thin sheets loosely attached to the substratum; each autozooid connected to its neighbours by six relatively short tubes, equally spaced around the basal periphery. Each autozooid has a single, tubular basal pore just proximal to the most distal connecting tube; frontal wall entirely membranous. Autozooids have five short, straight, distal spines, three on the distal edge and one adjacent to each of the distal avicularia, and six to eight pairs of longer, incurved, lateral spines equally spaced along its margins. The paired frontally facing, pedunculate avicularia lateral to the operculum are attached just basal to the frontal rim; short with a highly domed head and a strongly hooked rostrum; mandible acute triangular. A further six to eight avicularia, similar in shape and form to the others, appear later in astogeny, their peduncles forming as evaginations of the frontal rim, seemingly replacing the marginal spines. No ovicells observed. Measurements Holotype: means and standard deviations, mm. Autozooid length 0.57 & 0.03 (n = 16); width 0.31 k0.02 (n= 16). Avicularium length 0.14 & 0.01 (n = 15). Etymology From hex, Gr.-six; amicus, L.-friend, alluding to the six basal connections to neighbouring autozooids. Beania hexamicorum is similar to Beania discodermiae Harmer, 1926 which, however, only ever has two distal avicularia, with lengths equivalent to about one-third the total autozooidal length, substantially larger than those described above for B. hexamicorum. Beania petiolata Harmer, 1926 and B. hexamicorum are also very similar and occur in the same cryptic habitats. However, B. petiolata has more distal marginal spines than B. hexamicorum, six rather than five, and is further characterized by its basal spines and bifurcating marginal spines. Beania petiolata has more gracile, squared avicularia as opposed to the domed avicularia with strongly hooked mandible of B. hexamicorum, and does not produce the additional marginal avicularia found in the latter. Hayward s (1988) fragmentary specimens from Mauritius were insufficient to provide a type specimen,

17 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 51 or to make adequate comparisons with previously described species, but the more abundant material from Vanuatu, which has been found to be conspecific with the Mauritian specimens, allows this species to be properly characterized. Originally described from Mauritius by Hayward ( 1988) as Beania sp., Beania hexamicorum was a common species on the underside of coral rubble at Erakor Island, Efate. BEANIA CF. MIRABILIS HARMER (Fig. 5D,F) Reania mirabilis: Harmer, 1926 (?part): 419. Not Beania mirabilis Johnston, 1840: 272. Colony diffuse and ramifying, attached to the substratum by rhizoids which originate medially from the basal surface at the proximal end of the autozooid. Autozooids (zo.9 mm) boat-shaped, erect, the distal portion, i.e. that portion of the zooid free of the substratum, wider proximally than distally; frontal membrane occupying all of the erect part of the zooid. F roximal stolonic portions up to twice the length of the distal autozooids, originating latero- and meciioproximally from the wider proximal end. Two pairs of distal spines, one pair on the far distal edge, the other at the junction of the distal rim and the lateral margins, at the hinge of the operculum. Seven pairs of marginal spines overarch the frontal membrane. No ovicells or avicularia present. The single fragmentary colony of five autozooids from Iririki Island was found on a shell. The small number of autozooids is inadequate for description of a new species; however, they do show several differences from Johnston s type material from Scarborough, England, which throws doubt on their conspecificity with his species. The type specimen of Beania IH irabilis (NHM ) consists of only three autozooids but they still provide a guide to the identity of Johnston s species. The zooids are 0.70 mm long and spaced about 1.5 zooid lengths from each other. They have two pairs of very short, stout spines at their distal ends, the proximalmost pair being especially prominent, plus nine to ten pairs of marginal spines which arch over the frontal membrane. Another specimen (NHM ) in the Johnston Collection, from the type locality, is a better representation of the species; this consists of 22 autozooids (0.70 f 0.06 mm long (n = 22)), again on shell, which appear to be younger than those of the type specimen. The distal spines are very short, and the marginal spines range in number from two to six pairs, being absent from newly budded autozooids. A specimen from Guernsey (NHM , Norman Collection), is undoubtedly the same species as the type material. However, several other specimens attributed to B. mirabilis differ from the material just described. A specimen (NHM ) of seven zooids from Rapallo, North Italy, has slightly larger autozooids (0.80 mm) than the Johnston material of B. mirabilis, the same two pairs of distal spines, but 16 pairs of marginal spines; the zooids are spaced only one zooid length apart. Two specimens from Australia (NHM , Port Phillip Heads; NHM , Puebla Bay) are of another species altogether: this has two pairs of distal spines which are longer than in B. mirabilis and often the distal pair arches over to meet at the midline. The six to eight pairs of marginal spines are far more robust than in B. mirabilis and the zooids shorter (0.62 f 0.03 mm, n = 20) and deeper (frontobasally); finally, they are spaced only one zooid length apart. Harmer (1926), in describing B. mirabilis, had only a specimen of six zooids from the Philippines and one with just two zooids from the Torres Straits, yet he managed to count the number of tentacles, 12, far fewer, as he points out, than the 20 counted by Hincks (1880~) in, supposedly, the same species from Britain. Cook (1985) reported tentacles in a species with 2-22 marginal spines from West Africa. Harmer s (1926) material was not available for study, but he described two pairs of distal spines and 4-6 pairs of marginal spines, but provided no measurements. In the absence of Harmer s (1926) material it is difficult to say whether his specimens belong to B. mirabilis sensu stricto, to the same species as the specimens from Port Phillip Heads and Puebla Bay or to that described above from Vanuatu. Neither of these latter species is conspecific with B. mirabilis Johnston or with each other. FAMILY CANDIDAE D ORBIGNY, 1851 GENUS SCRUPOCELLARIA VAN BENEDEN, 1845 Type species: Sertularia scruposa Linnaeus, SCRUPOCELLARIA SPATULATA (D ORBIGNY) (Fig. 5A) Cellularia spatulata dorbigny, 1851: 50. Scrupocellaria spatulata: Harmer, 1926: 382, pl. 26, figs 1-10; Winston & Heimberg, 1986: 7, figs 11-12

18 52 K. J. TILBROOK ET AL. (cum syn.); Hayward, 1988: 283; Ryland & Hayward, 1992: 237, fig. 9. Scrupocellaria spatulata is widely distributed throughout the Indo-West Pacific region, having previously been reported from eastern Australia, East Africa, and the Philippines. Its occurrence at localities in Vanuatu provides the most easterly report so far. D Hondt & Gordon (1996) recorded and illustrated S. spatulata from New Caledonia, but in their description they mention not having observed ovicells, whilst figuring a specimen which clearly has ovicells, and although none of the characteristic avicularia are figured, they reported them as present in specimens from southern New Caledonia. This species was quite common at Erakor Island and Port Vila Harbour on small pieces of coral rubble and in loose debris. It appears to be a frequent component of cryptic coral reef communities. FAMILY EPISTOMIIDAE GREGORY, 1893 GENUS SYNNOTUM PIEPER, 1881 m e species: Loricaria aegyptiaca Audouin, SYNNOTUM AEGYPTIACUM (AUDOUIN) (Fig. 6A,C) Loricaria aegyptiaca Audouin, 1826: 243; Savigny, [1817]: pl. 13, figs Gemellaria (2) avicularis Pieper, 1881: 43, 47, pl. 2, figs 5-7. Synnotum aegyptiacum: Harmer, 1926: 398, pl. 27, figs 3, 4 (cum syn.); Winston, 1982: 127, fig. 53 (cum syn.); Gordon, 1984: 43, pl. 10, figs E,F. The type specimen of Pieper s (1881) Gemellaria avicularis, from the Adriatic Sea, is more heavily calcified than the specimens from Erakor Island, Vanuatu. Others (Harmer, 1926; Gordon, 1984) have also commented on less heavily calcified specimens of S. aegyptiacum. Older, basal ends of erect stems develop thicker calcification and opesial closure plates, apparently strengthening the colony. Thicker calcification may explain the survival of autozooids in the fossil record; Lagaaij (1968) reported its occurrence in the Tertiary (Miocene to Pliocene) of Indonesia and the Caribbean. It also has a circumglobal Recent distribution (Lagaaij, 1968). SUPERFAMILY MICROFQROIDEA GRAY, 1848 FAMILY ONYCHOCHELLIDAE JULLIEN, 1882 GENU SMITTIPORA JULLIEN, 1882 me species: Vincularia abyssicola Smitt, SMITTIPORA CORDIFORMIS HARMER (Fig, 6G) Smittipora cordiformis Harmer, 1926: 260, pl. 16, figs Smittipora cordiformis: Hayward & Ryland, 1995a: 543, fig. 6C. Several colonies of S. cordiformis occurred in the Vanuatu collections, encrusting small pieces of coral rubble from Erakor Island. Harmer (1926) described the species from material collected by the Siboga in Indonesia and reported additional material from the Indian Ocean (Burma and the Amirante Islands). Its presence in the SW Pacific was established by Hayward & Ryland (1995a), and it may prove to be more widely distributed in reef habitats in the Coral Sea. FAMILY MONOWRELLIDAE HINCKS, 1882a GENUS MONOPORELLA HINCKS, 1881b m e species: Monopodla nodulifera Hincks, 1881b. MONOPORELLA NODULIFERA (HINCKS) (Fig. 19E,F) Haploporella nodulifera Hincks, 1881b: 11. Monoporella nodulifera: Hincks, 1881b: 135, pl. 1, fig. 4; Harmer, 1926: 310, pl. 20, figs 21-23; Hayward, 1974: 374, figs 3a,b; Gordon, 1984: 53, pl. 16A,B. Monoporella fim briata carinifera Canu & Bassler, 1929: 157. Monoporella nodulifera is easily distinguished by its dark brown pigmented frontal membrane, blackishbrown D-shaped orifice, and three to five distal-oral spines. The concave cryptocyst has a median crest with a small round opesiule on either side, below the corners of the orificial rim. Originally described from the Bass Straits, Monoporella nodulifera has subsequently been recorded from the Mediterranean, Cape Verde Islands, Torres Straits, Molluccas, Philippines, China Sea, and Kermadec Ridge, New Zealand but it has has not been recorded from the Great Barrier Reef or the Coral Sea. Several fertile colonies were found encrusting coral rubble from Port Vila Harbour, Efate. FAMILY STEGINOWRELLIDAE HINCKS, GENUSTEGINOPORELLA SMITT, 1873 Type species: Membranipora magnilabris Busk, 1854.

19 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 53 Figure 6. A, Synnotum aegyptiacum. B, Steginoporella jellyae nom. nov. C, Synnotum aegyptiacum. D, Steginoporella jellyae nom. nov. Ancestrula. E, Thulamoporella gracilata sp. nov. Vicarious avicularium and primary orifice. F, Thulamoporella gracilata sp. nov. G, Smittipora codformis. Scale bars: A=0.20 mm; B = 0.50 mm; C = 0.50 mm; D = 0.25 mm; E = 0.20 mm; F = 0.40 mm; G = 0.25 mm.

20 54 K. J. TILBROOK ET AL. STEGINOPORELLA JELLYAE NOM. NOV. (Fig. 6B,D) Steganoporella lateralis: Harmer, 1900: 242, pl. 12, fig. 1; pl. 13, figs 19, 20, 27; Levinsen, 1909: 168, pl. 5, fig. 7; Harmer, 1926: 274, pl. 17, figs 5, 6, 8, 13 Pouyet & David, 1979: 783, text fig. 2. Not Steganoporella lateralis MacGillivray, 1895: 53, pl. 6, fig. 18. Material Holotype: NHM , Tahiti, collected by Miss Jelly, Hincks Coll. Other material examined: NHM , Kur Island, Kei Islands, 0-45 m; NHM , East of Palau, 5" 35' 32"S, 106" 33' 55"E, 110 feet; NHM ,76,308, Port Vila Harbour, Efate, Vanuatu, 4-5 m. Colony forming flat, encrusting sheets, horn-coloured when dried. Autozooids rounded distally, straight or concave proximally, generally parallel-sided. Gymnocyst most prominent as a smooth raised distal margin surrounding the operculum, apparently ending at the lateral opercular condyles, but may be seen as a thin band surrounding the entire cryptocyst. Cryptocyst occupying about half total length of autozooid; central portion flat, finely nodular, with numerous fine perforations; surrounded by a raised, non porous, tuberculate rim, particularly prominent at the distal edge, i.e. the proximal border of the opesia, which is generally concave. Frontal membrane thick, opaque. The rounded polypide tube is situated centrally and vertically, visible in frontal view; it has a more or less strongly flared distal margin with a pair of lateral 'horns', lacking the marginal flanges seen in other Steginoporella species. Measurements Holotype. means and standard deviations, mm (n= 25). Autozooid length 0.90 k0.08; width 0.50 f Etymology Steginoporella jellyae nomen n o w for Steginoporella lateralis auctt. non MacGillivray, Named for Miss Eliza C. Jelly in recognition of her contributions to bryozoology in the nineteenth century and as the collector of the holotype specimen. Steginoporella jellyae is characterized by having a complete distal edge to its cryptocyst, unlike other species such as Steginoporella magnilabris (Busk, 1854) in which the distal edge descends to the basal wall. S. jellyae appears initially to grow directionally in encrusting, bifurcating, bi- or triserial, fingers. Steginoporella jellyae has consistently been assigned to Steginoporella lateralis MacGilliway, 1895, a fossil species from the Tertiary of Victoria, Australia. By comparison of material with MacGilliway's (1895) excellent illustration it is obvious that the two species differ. The presence of a complete distal edge to the cryptocyst in S. jellyae is unique amongst described species of Steginoporella and makes identification from illustrations by authors easy. No dimorphic B zooids were observed in the material of Steginopodla jellyae from Vanuatu; however Harmer (1926) illustrated the two types and suggested that one of Levinsen's (1909) illustrations shows the same. Two unregistered specimens from the Solomon Islands, in the Soule Collection at the University of Southern California, have B zooids present: the operculum is darker in colour and wider than A zooids; 6-8 mandibular teeth, with four larger than the rest. The distal cryptocyst described in such species as S. magnilabris as an 'oral shelf' (Levinsen, 1909; Ryland & Hayward, 1992) is absent in S. jellyae. The basal wall of the polypide tube may sometimes be formed by the basal surface of the autozooid (Levinsen, 1909). Since Harmer (1926). this species has been recorded twice, by Brown (1956), from the Pliocene of South Australia and Gurgel & Vasseur (1974) from the Recent, Mozambique (though neither illustrated the species), surprising when areas within and neighbouring its known Pacific distribution have recently been studied (e.g. Winston & Heimberg, 1986; Scholz, 1991; Ryland & Hayward, 1992; Hayward & Ryland, 1995a). Several ancestrulae were found in the specimens studied. Only one primary zooid was ever found with no evidence of the secondary zooids described by Cook (1985), in Steginoporella buslzii Harmer, 1900 and S. magnila bris. A second as yet undescribed species of Steginoporella (NHM unknown locality, Hincks Collection.) also has a complete distal edge to its cryptocyst; however, the polypide tube, visible in frontal view, is situated asymmetrically and basally, it is wider, has a more strongly flared distal margin with a pair of lateral marginal flanges not seen in Steginoporella jel lyae. Several small colonies of Steginoporella jellyae were found associated with Antmpora granulifera, Monoporella nodulifera and Crepidacantha longiseta, encrusting coral rubble and overgrowing other bryozoans.

21 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 55 Harmer's (1900) records of this species from Torres Straits and Tahiti suggest that this species should be found on suitable reef substrata between these two outliers and its occurrence in Vanuatu and the Solomon Islands would confirm this. The presence of this species at Port Vila Harbour is the first record from this area. The records of Brown (1956) and Gurgel & Vasseur (1974) must be substantiated by examination of their material. FAMILY THALAMOPORELLIDAE LEVINSEN, 1909 GENUS THALAMOPORELLA HINCKS, 1887 Type species: Flustra mzieri Audouin, THALAMOPORELLA GRANULATA LEVINSEN l'halamoporella granulata, var. B Levinsen, 1909: 189, 190, pl. 6a, fig. la-f. Thalamoporellagranulata: Harmer, 1926: 297; Ryland & Hayward, 1992: 241, fig. lla,b; Soule, Soule & Chaney, 1992: 48, figs Levinsen (1909) reported three varieties of Thalarnoporella granulata, denoting the species now regarded as T granulata sensu strict0 as uar. B, from the Torres Straits. The other two varieties were also given full specific status by Harmer (1926), such that uar. A is now regarded as T stapifera Levinsen, 1909 and var. C as I: tubifera Levinsen, Although not found in our material from Vanuatu, it was found at three sites in Mele Bay, Efate, by Soule, Soule & Chaney (1992). With their records and that of Ryland & Hayward (1992), this species has a very wide distribution within the Indo-West Pacific, having been recorded from the Seychelles, Indonesia, the Philippines, Great Barrier Reef, Solomon Islands, Vanuatu, Fiji, Tonga, and from the Pliocene of Taiwan. THALAMOPORELLA GRACILATA SP. NOV. (Fig. 6E,F) Material Holotype: NHM , Erakor Island reef flat, Efate, Vanuatu. Paratypes: NHM , NHM , NHM , NHM , all same locality and collection data as holotype. Colony unilaminar, encrusting. Autozooids rectangular, distinct, separated by shallow grooves. Gymnocyst reduced, present as a narrow margin, most noticeable around the anter of the orifice. Cryptocyst with a flat, finely beaded surface, becoming more granular with ontogeny, reaching two-thirds the way around either side of the orifice, punctured frontally by irregularly spaced pores; a raised, beaded mural rim proximally and laterally. Opesiules irregularly oval, almost rectangular, generally of unequal size, cryptocyst descending to basal wall proximally and laterally. Orifice wider than long; anter raised, slightly wider than poster; angular, lateral condyles positioned at edge of cryptocyst. Avicularium shorter than autozooid, uncommon; rostrum slightly raised, constituting half total length, acute triangular, with smooth distal platform; cryptocyst imperforate, finely granular; single, central, pear-shaped opesia; two pointed condyles at junction of distal gymnocyst and proximal cryptocyst. Ovicellate zooids similar to autozooids; bivalved ovicell smooth, bulbous, thin-walled, with a triangular aperture and obvious median suture. Three types of spicules present: two sizes of almost straight compasses, small (60-80 km) and medium (2 130 pm), the larger spicules straighter than the smaller ones; one size of small closed calipers ( ~ 7 0 ~ ). Measurements Holotype. means and standard deviations, mm (n= 10). Autozooid length ; width 0.47 f0.03. Orifice length ; width 0.22k0.01. Etymology From gracilis, L.-slender, thin, alluding to its delicate appearance. The autozooids of Thalamoporellagracilata are similar to those of both T granulata and T tubifera, which differ from each other in the shape of the avicularia and the presence in T granulata of torquing i.e., twisting of the avicularium or its sibling zooid. However, T gracilata differs from both species, in the extension of the cryptocyst around the autozooidal orifice, and in the shape of the avicularia as both the latter have spatulate vicarious avicularia. The shape of the avicularia in T gracilata is more reminiscent of T stapifera, but it has opesiules far smaller relative to the autozooid size than does T gracilata. A second species of Thalamoporella was also found but the colony fragment consisted of only a few autozooids and was too small for adequate description. However, the colony fragment did include an avicularium, larger than the autozooids, with a subspatulate rostrum and two large cryptocystal opesia.

22 56 K. J. TILBROOK ET AL. It is hoped that further examples of this species will be found as a result of continuing work in the area. Several small colonies of Thalamoporella gracilata were found encrusting large pieces of coral rubble from Erakor Island reef flat only. GENUS LABIOPORELLA HARMER, 1926 Type species: Labiopora crenulata Levinsen, The species assigned to this genus require a thorough review, especially specimens assigned to Labioporella bursaria (MacGillivray, 1887). These include the Red Sea material described by Waters (1909) as L. bursaria, and Dumont (1981) as L. crenulata (Levinsen, 1909). Although species are generally distinguishable if vicarious avicularia are present, Harmer (1926) also used attributes of the polypide tube and cryptocystal laminae attachment to the basal wall as characters in his key. These characters should be fully detailed for each species in any future review, as it appears that even Harmer overlooked some of them. Two of the specimens listed by him under L. bursaria differ significantly enough from material of this species from Victoria, Australia (type locality) to be assigned to two separate species. Haswell's specimen (NHM ) from Holborn Island, Queensland is a specimen of L. spatulata Harmer, 1926, whereas the specimen from Siboga Stn 71 (NHM ) Makassar, SW Celebes, is another as yet undetermined species of Labioporella. Other criteria worth noting in a review of this genus are, as well as the size and shape of the vicarious avicularia, their association with surrounding autozooids, the size of the avicularian opesia, shape of the mandible, perforations of the cryptocyst, amount of autozooidal gymnocystal calcification, and the shape and ornamentation of the autozooidal cryptocyst. LABIOPORELLA SPATULATA HARMER (Fig. 7A) Labioporella spatulata Harmer, 1926: 283, pl. 21, figs 4-6. Labioporella bursaria: Ryland & Hayward, 1992: 243, fig. llc. Material Holotype: NHM , Paternoster Island, north of Sumbawa. Other material examined: NHM , Holborn Island, Port Denison, Queensland; NHM , NHM , Erakor Island reef flat, Efate, Vanuatu. Colony unilaminar, encrusting. Autozooids elongate, rectangular, separated by distinct grooves, x mm. Raised mural rim present, crenulate distally apart from the smoothly rounded middle portion of the distal terminal wall. Gymnocyst absent from autozooids; cryptocyst finely granular, occupying up to three-quarters of autozooid length, with a number of small pores in the middle region, flat, but dipping sharply adjacent to polypide tube. The frontally convex polypide tube has a transversely oval aperture. Vicarious avicularia as large as autozooids; cryptocyst finely granular, imperforate, with an oval foramen through which a transverse, medially thickened septum is visible, so dividing the cystid cavity in half; rostrum less than half length of the avicularium, squared, spatulate, with smoothly rounded gymnocystal calcification; thickened, proximolateral condyles present. Labioporella spatulata is characterized by the lightly granular autozooidal cryptocyst with few, relatively small, pores, by the squared spatulate avicularian rostrum, and the positioning of the oval avicularian foramen relative to the condyles. The small cryptocystal pores appear to be produced later in ontogeny as they are not present initially, but become more numerous with distance from the growing edge; the cryptocyst also thickens. Labioporella spatulata differs from L. bursaria in that the latter has an avicularium with a more rounded rostrum, and autozooids with a less-pronounced lesscrenulate mural wall. Although L. thornelyae Harmer, 1926 from Sri Lanka is similar to L. spatulata, it differs in having a far narrower spatulate avicularium, which appears to cause its sibling autozooid to be angled towards it; the autozooids of L. thornelyae have more numerous pores in the almost tuberculate cryptocyst. Labioporella cornuta Harmer, 1926, from south of Halmaheira (Djilolo), is easily distinguishable from those species already mentioned as it has clear, though minimal, gymnocystal calcification around the autozooids. Finally, in a Torres Straits specimen assigned to L. crenulata (Levinsen, 1909) by Harmer (1926, (the locality of the material described by Levinsen IS unknown), the avicularium affects the orientation of adjacent autozooids on each side; the rostrum is broadly spatulate and distally rounded and autozooids have very large pores through their granular cryptocyst. This is a new species. Ryland & Hayward (1992) based their determination

23 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 57 Figure 7. A, Labioporella spatulata. B, Cellaria punctata. C,D, Puellina egretta. Note the keel on each of the two ovicells at the top of the figure. D, primary orifice. E,F, Puellina vulgaris. F, primary orifice. Scale bars: A=0.40mm; B = 0.20 mm; C = 0.25 mm; D = 0.05 mm; E = 0.25 mm; F = 0.05 mm.

24 58 K. J. TILBROOK ETAL. of L. bursaria from Heron Island on the Haswell specimen (NHM ) of L. spatulata from Queensland, wrongly identified by Harmer (1926) as L. bursaria. Their material should therefore be assigned to L. spatulata. Labioporella spatulata was described from a single colony from Indonesia. It has subsequently been found to occur off the coast of Queensland at Heron Island (Great Barrier Reef') and at Vanuatu. On Vanuatu, this species was found encrusting both large and small pieces of coral debris from Erakor Island, Efate. SUPERFAMILY CELLARIOIDEA FLEMING, 1828 FAMILY CELLARIIDAE FLEMING, 1828 GENUS CELLARIA ELLIS & SOLANDER, 1786 CELLARIA PUNCTATA (BUSK) (Fig. 7B) Salicornia punctata Busk, 1852a: 366 (in part). Salicornia gracilis Busk, : 17, pl. 63, figs 14-16, pl. 65, fig. 2. Cellaria punctata: Harmer, 1926: 337, pl. 21, figs 14-16, text-fig. 13a; Hayward and Cook, 1983: 34, fig. 8B; Hayward and Ryland, 1995a: 545, fig. 7D. This species was discussed by Hayward & Cook (1983), when describing material from the Agulhas Shelf, eastern South Africa. Although Cellaria punctata was originally described from Queensland (Busk, 1852a), Hayward & Ryland (1995a) found only one small colony at Heron Island. Two fragmentary internodes were found in loose coral debris from Port Vila Harbour, Efate, Vanuatu. Harmer (1926), in his synonymy, extended the range of this species throughout the Indo-Malay region westwards to Sri Lanka and Zanzibar, and northwards to Japan. The presence of C. punctata in the Indian Ocean was confirmed by Hayward & Cook (1983) who recorded it from eastern South Africa and the Red Sea. SUPERFAMILY CRIBRILINOIDEA HINCKS, 1879 FAMILY CRIBRILINIDAE HINCKS, 1879 GENUS PUELLINA JULLIEN, 1886 Qpe species: Lepralia gattyae Landsborough, The European species of Puellina have been reviewed and revised by Bishop & Househam (1987). Species of Puellina are common in tropical reef environments but have usually been identified with poorly defined NE Atlantic taxa. All records from tropical regions need to be re-examined in the light of Bishop & Househam (1987) and it is possible that many undescribed species will be revealed. The two following species were only recently recognized from Heron Island, Great Barrier Reef. PUELLINA EGRETTA RYLAND & HAYWARD (Fig. 7C, 0) Cribrilaria aff. radiata Ristedt, 1985: 20, fig. 3e-h. Puellina egretta Ryland & Hayward, 1992: 244, fig. lld,e. Puellina egretta is distinguishable from I? vulgaris Ryland & Hayward, 1992 by having only five oral spines, more costae (c. 19), and avicularia with very long rostra produced from a pronounced cystid that appears almost vicarious. Puellina egretta also produces far larger colonies than f? uulgaris. Ryland & Hayward (1992) were unable to find ovicells in the material of f? egretta they described from Heron Island, however, several ovicellate colonies have been found in the material from Vanuatu. The ovicell of f? egretta is Type A sensu Bishop & Househam (1987) and longer than that seen in I! uulgaris; the two halves of the ectooecium clearly come together to form a raised midline suture which resembles a keel. Puellina egretta was the less common of the two species found only at Port Vila Harbour, Efate encrusting various sizes of coral debris. This species has been previously recorded from the Seychelles, the Philippines and Heron Island, Great Barrier Reef. PUELLINA VULGARIS RYLAND & HAYWARD (Fig. 7E, F) Puellina vulgaris Ryland & Hayward, 1992: 244, fig. 12b,c. Puellina uulgaris was twice as common in the Vanuatu samples as f? egretta. It is distinguished by seven oral spines, fewer costae (15-16), and generally smaller avicularia. However, larger dimorphic avicularia are often produced, which look similar to those of f? egretta but lack the characteristic pronounced cystid of that species. Colonies off? uulgaris are much smaller than

25 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 5s those of? egretta. The ovicells of I! vulgaris are m e C sensu Bishop & Househam (1987). Puellina vulgaris was described from Heron Island, Great Barrier Reef and this is only the second record of what appears to be quite a common species. It was found encrusting coral debris in Port Vila Harbour, Efate. GENUS FIGULARIA JULLIEN, 1886 '&pe species: Lepralia figularis Johnston, FIGULARIA PULCHERRIMA SP. NOV. (Fig. 8A) Material Holotype: , Iririki Island, Efate, Vanuatu, Colony encrusting, unilaminar, flat. Autozooids irregularly oval to rectangular, distinct, separated by deep grooves (Holotype, ~0.7 x x0.4 mm). Primary orifice longer than wide, sides almost parallel, distal border concave, anter constituting over 80% of total length, separated from the slightly concave, proximal border by very small lateral condyles that are almost hidden by the distal costae. Operculum deeper in colour than the almost translucent frontal membrane. No spines. Frontal gymnocystal calcification smooth; costal field comprising almost half length and onethird area of frontal shield. Costae comprise 8-10 flattened tubes, closely packed with a series of small lacunae between, each with a single large oval or teardrop-shaped proximal pelma (lumen pore) frontally; 4-5 distal uniporous septula and 7-8 in the lateral wall. Avicularia not seen. Ovicellate zooids with a dimorphic orifice, wider than long. Ovicell almost spherical, smooth ectooecial calcification with a longitudinal median suture and a pair of lateral fenestra (a second smaller pair may be present, appearing to be pinched off from larger fenestrae); it also bears a rounded frontal costate shield, similar in appearance to that of the autozooids, with 6-7 costae. Etymology The name is the superlative form of the Latin, pulcher, beautiful. Figularia pulcherrima is characterized by its distinctive costate frontal shield, but more especially by the ovicell, which is also costate. This feature is uniquc within the genus, suggesting that the ovicell could bc regarded as a distinct polymorph. Figularia species with this character perhaps ought to be segregated into a new genus. Figularia pulcherrima is very similar to l? jucunck Canu & Bassler, 1929, from the Philippines, in the structure of the costate shield, but there are great differences in the morphology of the ovicells of the two species. Those of l? jucunda do not possess the costate shield seen in l? pulcherrima; instead, there are three central fenestrae, two lateral and one transverse distomedial, as well as up to eight circular marginal fenestrae. These are similar to those seen in I? lepida Hayward, 1988, from Mauritius, which as well as surrounding the margin of the ovicell, cover the frontal wall of the autozooids, leaving only a very small costate shield proximal to the orifice. Figularia lepida also has vicarious avicularia, not seen in either of the other two species. Examination of the inside of the costal shield of Figulariapulcherrima shows a series of proximal pores that constitute the entrances to the costal lumina. A single small colony of this species was found encrusting a heavily bored shell from Iririki Island. SUPERFAMILY CATENICELLOIDEA BUSK, 1852a FAMILY CATENICELLIDAE BUSK, 1852a GENUS CATENEELLA DE BLAINVILLE, 1830 Type species: Eucratea contei Audouin, CATENEELLA TRIANG ULIFERA (HARMER) (Fig. 21F) Vittaticella triangulifera Harmer, 1957: 777, pl. 50, figs 7, 19. Colony erect, branching, jointed, to 6.5mm high in present material. Internodes consist of single autozooids, or bizooidal units, which comprise bifurcations; in proximal portions of the colony there may be 2 single autozooids between successive bifurcations, distally only 1 or none. Single autozooids slender, fusiform; average dimensions mm long, mm wide at the broad distal end. Primary orifice as wide as long, proximal edge shallowly concave, but arched frontally such that the orifice appears foreshortened. A small avicularium on each distolateral corner of the autozooid, level with and longer than the orifice; distal, apical portion of cystid projecting outwards at an oblique angle to the long axis of the

26 60 K. J. TILBROOK ET AL. autozooid, tapered and often produced as a cylindrical process, variably and unequally developed on all autozooids. Two narrow pore chambers (vittae) extend for most of the length of the autozooid, one on each side of the frontal shield, each with a single series of conspicuous pores. The proximal zooid of the bizooidal unit is as large as the single autozooid, the distal one slightly shorter; axial avicularium of the proximal zooid suppressed, distal zooid with the usual pair. No fertile autozooids present. Small colonies of Catenicella triangulifera, perhaps ten in all, were collected from beneath dead coral heads at Erakor reef flat. Catenicella Zvenusta MacGillivray figured by Gordon (1984: 67, pl. 22, fig. c) seems very similar to the material hefe assigned to C. triangulifera following comparison with Harmer s (1957) type material. According to Harmer (1957: 776), MacGillivray s species is distinguished especially by its ovicell, which bears a longitudinal frontal ridge with a line of pores on each side. In C. triangulifera the ovicell has a peripheral band of pores and a variable frontal foramen. Neither Gordon s (1984) specimens nor the present material included fertile zooids. In Harmer s (1957) original description he recorded this species from Indonesia and Malaysia as well as north to Japan and west to the Amirante Islands. Thus Catenicella triangulifera has an Indo-West Pacific distribution, reinforced by its occurrence in Vanuatu. GENUS VASIGNYELLA GORDON, 1989a Type species: Catenaria otophora Kirkpatrick, VASIGNYELLA OTOPHORA (KIRKPATRICK) Catenaria otophora Kirkpatrick, 1890: 17, pl. 5, figs la-c. Savignyella otophora: Harmer, 1957: 763, pl. 51, figs 19-21; Winston, 1986: 27. Vasignyella otophora: Gordon, 1989a 453, figs 13-15; Ryland & Hayward, 1992: 247, fig. 14a. Fragmentary colonies of this species were found at Erakor Island, in loose coral debris and on coral rubble in cryptic areas. It seemed to be associated with Savignyella lafontii as part of the cryptic bryozoan sward. Vasignyella otophora is probably common within the Ind*West-Pacific but it is particularly inconspicuous. Although most records are from the Indo-Malayan region, it has been recorded from the Red Sea (Dumont, 1981) and east to Western Samoa (Gordon, 1989a). FAMILY SAVIGNYELLIDAE LEVINSEN, 1909 SAWGNYELLA LEVINSEN, 1909 Type species: Eucratea lafontii Audouin, SAVIGNYELLA LAFONTII (AUDOUIN) (Fig. 8B) Eucratea lafontii Audouin, 1826: 242. Catenaria lafontii: Hastings, 1930: 732. Sauignyella lafontii: Harmer, 1957: 761, pl. 51, figs 11, 12; Hayward, 1988: 327; Gordon, 1989a: 453, figs 11) 12. The fragmentary specimens, in loose debris and within cavities in pieces of coral rubble from Poanangisu, are the same as those illustrated by Gordon (1989a) but lacked the avicularia or ovicells he described. It was found in association with Vasignyella otophora (Kirkpatrick, 1888) and Beania hexamicorum sp. nov. This inconspicuous species has been widely recorded. Lagaaij (1968) provided a comprehensive guide to its distribution, which extends throughout the Indo- Pacific Realm and the tropical and warm temperate Atlantic. He also reported Miocene specimens from the Gulf of Mexico and the Caribbean. It should be noted, however, that no type specimen exists and the taxon might prove to encompass more than one species. SUPERFAMILY HIPPOTHOIDEA BUSK, 1859 FAMILY HIPPOTHOIDEA BUSK, 1859 GENUS HIPPOTHOA LAMOVROUX, 1821 npe species: Hippothoa divaricata Lamouroux, Species of Hippothoa are fairly inconspicuous as they are small in size, hyaline in appearance, and cryptic in habit. The two species found in Vanuatu, H. calciophilia and H. flagellum, can be differentiated in several ways, including: H. flagellum is smaller (~0.25mm) than H. calciophilia (x0.35 mm). H. flagellum has only one pore chamber in each lateral wall compared to H. calciophilia which has two. Ovicellate zooids are shorter than autozooids in H. flagellum (similar size in H. calciophilia) and the orifice is markedly dimorphic. Evidence of Hippothoa colonies was present on a number of specimens other than those mentioned

27 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 61 Figure 8. A, Figularia pulcherrima sp. nov. Note ovicellate zooid, centre right. B, Savignyella lafontii. C, Chorizopora bmngniartii. D, Nimba saxatilis. E, Wostega maculata sp. nov. Primary orifice and zooeciule, above. F, Trypostega maculata sp. nov. G, Exochonella brasiliensis. Scale bars: A = 0.50 mm; B = 0.50 mm; C = 0.20 mm; D = 0.25 mm; E = 0.05 mm; F = 0.25 mm; G = 0.50 mm.

28 62 K. J. TILBROOK ET AL. below; owing to difficulty of identification these were not assigned to species. HIPPOTHOA CALCIOPHILIA GORDON (Fig. 9A,B) Hippothoa calciophilia Gordon, 1984: 108, text-fig. 10; pl. 42, figs A-C. Hippothoa calciophilia: Ryland & Hayward, 1992: 247, fig. 13a,b. Hippothoa calciophilia is very similar to H. distans MacGilliway, 1869, however, Gordon (1984) distinguished his new species on the basis of the distribution of communication pores, and the smooth ovicell, which is apically sculptured in H. distans. The wide geographical distribution of Hippothoa calciophilia was recorded by Gordon (1984): from New Zealand and the Great Barrier Reef, and from the eastern Pacific, including the Galapagos Islands, from Panama to Peru, as well as from the Bahamas. Ryland & Hayward (1992) record it from Heron Island, Great Barrier Reef. Several colonies were found encrusting small pieces of coral rubble at Port Vila Harbour and Erakor Island. HIPPOTHOA FLAGELLUM h Z0NI (Fig. 9C,D) Hippothoa flagellum Manzoni, 1870: 328. Hippothoa flagellum: Hayward & Ryland, 1979: 248, fig. 107; Gordon, 1984: 111, text-fig. 10D, pl. 43, figs C,D; Hayward, 1988: 326; Hayward & Ryland, 1995a: 547. This is one of the smallest known species of Hippothoa and favours biogenic carbonates as substrata (Hayward & Ryland, 1995a). Originally described from the Pliocene of Italy, H. flagellum seems to be one of the few truly cosmopolitan bryozoan species, occurring from subpolar regions to the tropics in shallow shelf habitats. In Vanuatu it was found on several small pieces of coral rubble and shell at Port Vila Harbour and Erakor Island. GENUS PLESIOTHOA GORDON & HASTINGS, 1979 m e species: Hippothoa gigerium Ryland & Gordon, PLESIOTHOA BUCARINA SP. NOV. (Fig. 9E) Material Holotype: , Erakor Island, Efate, Vanuatu, Colony comprising intergrown, branching chains of zooids. Autozooids and female zooids present. Male zooids and zooeciules not observed. Autozooids elongate, slender, broadest distally, tapered proximally, strongly convex (Holotype, x mm). Frontal shield thickly calcified, smooth, developing 3-5 umbones along midline; uncalcified windows present along the margins. Primary orifice orbicular, slightly longer than wide with a deep teardrop-shaped sinus; slight constriction in the anter distal to the condyles, which appear bifid through the translucent operculum. Generally four lateral pore chambers present, a distal pair appearing as raised, laterally compressed tubes. Female zooids similar in size to autozooids; ovicell not observed, but it appears to be budded from the raised distal pore chambers; female primary orifice differing from the autozooidal orifice, wider than long, semielliptical with a U-shaped sinus, proximal border convex on either side of sinus; condyles appear flat nok bifid. Etymology From bu- L.-prefix, large. carina L.-keeled. Named for its large prominent umbones. Plesiothoa bucarina, the largest-known Plesiothoa, is based on a single, small, fragmented colony that was found in coral debris from Erakor Island reef flat. The marginal windows are uncalcified and connect with the lateral pore-chambers as illustrated by Gordon C Hastings (1979) in I? trigemma. GENUS TRYPOSTEGA LEVINSEN, 1909 Type species: Lepralia venusta Norman, Osburn (1952) modified Levinsen s (1909) original diagnosis, which stated no avicularia, having noted spatulate opercula closing the apertures of the zooeciules in Trypostega claviculata (Hincks, ). He therefore suggested that zooeciules in T venusta were vestigial avicularia, being very minute with apparently no mandibles.

29 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 63 Figure 9. A,B, Hippothoa calciophiliu. B, note the proximo-lateral position of the zooeciules. C,D, Hippothoa pagellum. E. Plesiothoa bucurinu sp. nov. Scale bars: A = 0.12 mm; B = 0.18 mm; C = 0.12 mm; D = 0.40 mm; E = 0.36 mm.

30 64 K. J. TILBROOK ET AL. TRYPOSTEGA MACULATA SP. NOV. (Fig. 8E, F) Material Holotype: NHM , Port Vila Harbour, Efate, Vanuatu, October Paratype: NHM , same locality and collection data as holotype. Colony encrusting, forming broad, thin sheets. Autozooids oval to irregularly polygonal in shape, slightly convex, separated by distinct shallow grooves. Frontal shield smooth, hyaline, perforated by numerous, evenly spaced, round pores. Orifice keyhole-shaped, slightly longer than wide; deep, rounded anter, separated by short proximally pointing triangular condyles from shallower, semi-circular or rounded triangular poster; very slight raised rim around orifice. Ovicell hyperstomial, prominent, oval, slightly trilobed in appearance, the middle lobe largest and most prominent, as evenly perforated as the frontal shield; orifice of ovicellate zooids wider than long. Zooeciules (modified avicularia), appear distal to most autozooids and ovicells, calcification and perforation similar to autozooids; orifice small, mandible rounded or elliptical. Measurvments Holotype. means and standard deviations, mm (n= 25). Autozooid length 0.48 k0.07; width Orifice length 0.09 _+ 0.01; width 0.08 f Etymology From macula L.-spot, alluding to the large frontal pores seen in this species. The ovicells are polymorphs; the brood chamber is separated from the maternal zooid by an incomplete distal wall and is closed by the operculum of the maternal zooid. It is almost certain that Trypostega maculata has been confused with T. uenusta in the past; however, it differs from T. venusta in the shape of the primary orifice and the lack of a proximal oral umbo. The orifice of T uenusta has a sagittate poster with very slight lateral indentations proximal to the condyles, whereas the orifice of T. maculata is far more rounded and lacks the lateral indentations proximal to the condyles. Trypostega maculata is similar to T claviculata (Hincks, 1882b), both having very large frontal pores. However, they differ in the shape of the orifice: the poster of the primary orifice of T claviculata is far shallower and wider than that of T. maculata. Also, the shape of the zooeciular orifice differs in the two species; that of T claviculata has a far larger opening when compared to T maculata and bears a spatulate mandible. The most frequently recorded species of Trypostega, T. venusta has been accorded a wide distribution in warmtemperate and tropical seas, from all oceans, and it is possible that the occurrence of other, similar, species has been hidden, and that a re-examination of material attributed to T. venusta would uncover wrongly assigned specimens and even new species. Trypostega. maculata has only been found encrusting coral rubble in Port Vila Harbour. FAMILY CHORIZOFORIDAE VIGNEAUX, 1949 GENUS CHORIZOPORA HINCKS, 1879 Type species: Flustra bmngniartii Audouin, CHORIZOPORA BRONGNIARTII (AUDOUIN) (Fig. 8C) Flustra bmngniartii Audouin, 1826: 240; Savigny, [1817]: pl. 10, fig. 6. Chorizopora bmngniartii: Hayward & Ryland, , fig. 103; Gordon, 1984: 113, pl. 44C; Hayward, 1988; Ryland & Hayward, 1992: 249, fig. 13c. The thin, translucent colonies of this species make it inconspicuous on the coral rubble specimens examined for this study. However, Chorizopora bmngniartii is easily distinguished from other species of Chorizopora (e.g. Gordon, 1984) by the smooth or transversely wrinkled, but non-spinous, frontal walls of its autozooids. Chorizopora bmngniartii is widely distributed in both temperate and tropical shelf seas; in Vanuatu it was found on small pieces of coral rubble in Port Vila Harbour. SUPERFAMILY ARACHNOPUSIOIDEA JULLIEN, 1888 FAMILY ARACHNOPUSIIDAE JULLIEN, 1888 GENUS PORICELLA CAN, 1904 Type species: Poricella maconnica Canu, Thanks to the courtesy of Dr Juan J. Lopez Gappa (Argentine Museum of Natural Sciences, Buenos

31 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 65 Aires), it has been possible to ascertain that Poricella is a senior subjective synonym of Trernogasterina auctt., (not Canu, 1911). Dr Lopez Gappa scanned the type specimen of Trernogasterina pmblematica (and made photographs available to DPG and P.L. Cook), showing that it is not congeneric with the many species attributed to Tremogasterina by authors (see Cook, 1977). The species of Poricella (i.e. Tremogasterina auctt.) appear to have a worldwide distribution in shallow waters between 30" latitude north and south. The limited number of fossil and Recent species known were reviewed by Cook (1977). PORICELLA ROBUSTA (HINCKS) (Fig. 1OA) Lvpralia mbusta Hincks, 1884b: 360, pl. 13, fig. 4. Lt)pralia mbusta: Hincks, 1887: 131; Thornely, 1905: 119; Waters, 1909: 152, pl. 13, figs 13, 14. Twrnogasterina mbusta: Powell & Cook, 1967: 12, pl. 1, figs c,d; text-figs 1-4; Cook, 1977: 133, pl. 5, fig. E, pl. 8, figs B,C. Dvscript ion Colony encrusting, forming large multilaminar patches. Autozooids oval, convex, separated by deep grooves in early ontogeny; c x 0.60 mm. Orifice longer than wide, slightly narrower proximally, anter separated by two very inconspicuous condyles; 2-3 distal oral spines present. Frontal wall granular, with large marginal pores and 1-3 central foramina (which may be occluded). Interzooidal avicularia (equal in number to autozooids) large, asymmetrical or spatulate; crossbar complete; rostrum with finely serrated distal edges; directed distally or distolaterally. Ovicell imperforate, finely tuberculate, initially prominent but becoming immersed. 2-3 distal pore-chambers present. Rein arks Colonies are orange in life (Powell & Cook, 1967); rare vicarious avicularia have been noted in? mbusta but not described (Cook, 1977). Poricella mbusta has never been found at depths greater than 37 m. Originally described from the Mergui Archipelago (coast of Myanmar), it has subsequently been described from the Persian Gulf, Red Sea, Seychelles and Sri Lanka. However, of the three Indo-Pacific species of Poricella noted by Cook (1977) (as Tremogasterina), (I? cellcpomides, l? mbusta, l? spathulata),? mbusta has never before been recorded in any part of the Pacific. It was found encrusting discarded wall tiles in Port Vila Harbour. PORICELLA SPATHULATA (CANU & BASSLER) (Fig. 1OB) Hiantopora spathulata Canu & Bassler, 1929: 116, pl. 11, figs 13, 14. Arachnopusia spathulata: Harmer, 1957: 657, pl. 68, figs 22, 23, 25, 31. Tremogasterina spathulata: Cook, 1977: 136, text-figs lb, 5A, pl. 5A; Hayward, 1988: 296, pl. 5D; Scholz, 1991: 289, pl. 7, fig. 6; Ryland & Hayward, 1992: 249, fig. 13e. Large vicarious avicularia were present in a colony from Port Vila Harbour. Their rare occurrence in colonies was noted by Cook (1977); most lack them but when they are present within a colony they are numerous. Cook (1977) gave a very comprehensive guide to the distribution of Poricella spathulata (as Tremogasterina), showing that it is widely distributed in the tropical Indo-West Pacific from the Red Sea to the Great Barrier Reef. Poricella spathulata is very common on inner reef flats and was present in over half of the samples studied by Ryland & Hayward (1992) from Heron Island, Great Barrier Reef. It was the most commonly found bryozoan species in Vanuatu, in samples from Poanangisu, Iririki Island and Erakor Island, Efate, making these the most easterly records for this species. FAMILY EXECHONELLIDAE HARMER, 1957 GENUS EXECHONELLA DUVERGIER, 1924 Type species: Cyclicopora? grandis Duvergier, EXECHONELLA BRASILIENSIS CANU & BASSLER (Fig. 8G) Exechonella brasiliensis Canu & Bassler, 1928b: 72. Exechonel la brasiliensis: Winston & Heimberg, 1986: 15, figs 26,27; Winston, 1986: 19; Scholz, 1991: 289, pl. 7, fig. 1. Winston & Heimberg (1986) give a very comprehensive description and discussion of this species, having found a single colony in their samples from Indonesia. As its name suggests this species was originally described from Brazil; however, several colonies were found at both Iririki Island and Port Vila Harbour on small pieces of coral rubble and discarded wall tiles. This record firmly establishes the presence of this species in the Southwest Pacific area.

32 66 K. J. TILBROOK ET AL. SUPERFAMILY ADEONIDEA BUSK, 1884 FAMILY ADEONIDAE BUSK, 1884 GENUS REPTADEONELLA BUSK, 1884 m e species: Lepralia violacea Johnston, REPTADEONELLA FISSA (HINCKS) (Fig. lle,f) Microporella fissa Hincks, 1880b: 381, pl. 17, fig. 4. Adeona pomsa Canu & Bassler, 1929: 376, pl. 50, fig. 7. Adeona joloensis Bassler, 1936: 161 (nom. nov. pro Adeona pomsa Canu & Bassler, 1929). Reptadeonella joloensis: Harmer, 1957: 816, pl. 54, figs 1-3, text-fig. 86A-C; Ryland, 1974: 343; dhondt, 1986: 735; Winston, 1986: 24. Colony encrusting, forming unilaminar sheets that are often extensive. Autozooids irregularly polygonal or oval, convex, distinct, separated by shallow grooves, becoming less pronounced in later ontogeny ( x mm). Frontal shield finely granular but with some pronounced tuberculations; a single series (sometimes 2) of large, closely spaced, marginal pores becoming less distinct with ontogeny; a single round spiramen in depression at centre of zooid. Primary orifice semicircular, as is the secondary orifice, which surmounts a short peristome. A suboral avicularium always present, short, triangular, set medially on the sloping proximal side, directed distolaterally; no crossbar. Additional avicularia, similar in size and shape to the suboral avicularium, are often produced later in ontogeny, generally at the proximal end of the autozooid; variably orientated. Some autozooids bear an extremely large, sickle-shaped adventitious avicularium in place of the suboral avicularium, originating lateral to the midline between the spiramen and peristome, looping out distolaterally around the orifice, expanded proximally, narrowing distally; no crossbar. Gonozooids with a crescent-shaped orifice. The presence of the extremely large adventitious avicularium, figured by Hincks (1880b) which all but covers the frontal shield of the autozooid which bears it, is the most distinctive character of Reptadeonella fissa. Micmporella fissa was originally described by Hincks (1880b) from Indian Ocean specimens. His comprehensive description and clear illustration has seemingly been overlooked ever since. Adeona joloensis Bassler, 1936 (nom. nov. pro Adeona pomsa Canu & Bassler, 1929) from the Philippines, has been the name applied to all subsequent records of this species. Comparison of Hincks type material (NHM , , both Indian Ocean), Canu & Basslerk holotype (USNM 8170, Philippines), cotype (NHM , Philippines) of Adeona pomsa, and other material in the Natural History Museum, London with the material from Vanuatu shows that this is all one species, although there are certain minor differences among the specimens. In the series of type specimens, several adventitious avicularia may be present on the frontal shield in addition to the usual avicularium proximal to the orifice, between the orifice and the ascopore. The position, size, and orientation of the large sickle-shaped adventitious avicularia also differ within the material. In the type specimen they are large and robust and extend the zooid laterally and distally to the orifice without affecting the shape of the orifice itself. However, in material from Funafuti (NHM ) these avicularia are smaller, more gracile and more intimately connected to the orifice. This is also the case in the type of Adeona pomsa Canu & Bassler, Other material from the Philippines is intermediate in size to those described above, with the avicularia modifying the shape of its peristome. This unmistakable species has been noted from the Indian Ocean (Hincks, 1880b), Philippines (Canu & Bassler, 1929), and Great Barrier Reef (Ryland, 1974), and was found encrusting both large and small pieces of coral rubble from Poanangisu and Port Vila Harbour, Efate. Whilst Harmer s (1957) illustrations of material from the Loyalty Islands appear to match the species described above, his records from Singapore, China Sea and Malaysia must be reconsidered following examination of original specimens. REPTADEONELLA CELLULANUS SP. NOV. (Fig. 12A-C) Material Holotype: NHM , Erakor Island, Efate, Vanuatu. Paratypes: NHM , Port Vila Harbour, Efate, Vanuatu; NHM , Poanangisu, Efate, Vanuatu. Colony encrusting, unilaminar; covered by a thin membrane with a metallic purple sheen when dried. Autozooids irregularly polygonal, convex, distinct, separated by shallow grooves; irregularly arranged.

33 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 67 Figure 10. A, Poricella mbusta. B, Poricella spathulata. Note rare vicarious avicularium, right top and bottom. C, Drc,panophora indica. D, Drepanophora tuberculata. E,F, Drepanophora gutta sp. nov. F, primary orifice, note the anvil-like proximal denticle. Scale bars: A = 0.40 mm; B = 0.25 mm; C = 0.40 mm; D = 0.20 mm; E = 0.25 mm; F = 0.04 mm.

34 68 K. J. TILBROOK ET AL. Figure 11. A-C, l'brquatella duolamellata comb. nov. B, primary orifice, note the laterally facing sub-oral avicularium. C, Schizoporelloid ancestrula and peri-ancestrular autozooids. D, Reptadeonella novissima sp. nav. E,F, Reptadeonella fissa. Note crescent-shaped orifice of maternal zooid, right of centre. F, Note large, sickle-shaped adventitious avicularium. Scale bars: A = 0.40 mm; B = 0.05 mm; C =0.20 mm; D = 0.40 mm; E = 0.30 mm; F = 0.20 mm.

35 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 69 Figure 12. A-C, Reptadeonella cellulanus sp. nov. A, colony origin, note directional growth. B, developing autozooids at colony edge. C, primary orifice. Scale bars: A=0.45 mm; B=0.40 mm; C =0.08 mm. Frontal shield finely granular, with larger tuberculations produced later in ontogeny; a single series of marginal pores extending around the distal margin surrounding the peristome making it subterminal; a single, large, oval medial spiramen, in depression at centre of zooid. Primary orifice semicircular, with a slightly denticulate, concave, proximal border; secondary orifice, which surmounts a thin tubular peristome, is also semicircular; an avicularium is generally present, set medially, on the sloping proximal side of the peristome, i.e. distal to spiramen; acute rostrum directed distolaterally; mandible acutely triangular; no crossbar. Gonozooids or reproductive structures not known. Colony origin from a single ancestrula, the form of which is not known. Mcasurements Holotype: means and standard deviations, mm (n= 25). Autozooid length 0.65 & 0.07; autozooid width 0.34 f Et? 1 mology From cellulanus, L.-hermit, single ancestrula. solitary, alluding to the The apparent lack of an ancestrular complex in this species, as well as the denticulate proximal border to the primary orifice, warrant the erection of a new species. The colony origin is assumed to be from a single ancestrula, although the paratype specimen which shows the colony origin is equivocal in this regard. It is evident though, that the origin is not from a sixzooid ancestrular complex as seen in Reptadeonella violacea (Johnston, 1847), based on the positioning of the periancestrular zooids. Three small, apparently infertile, colonies of Reptadeonella cellulanus were found encrusting pieces of coral rubble from Erakor Island, Poanangisu and Port Vila Harbour, Efate. Material REFMDEONELLA NOVISSIMA SP. NOV. (Fig. lld) Holotype: NHM , Port Vila Harbour, Efate, Vanuatu. Colony encrusting, unilaminar. Autozooids rounded, polygonal, convex, distinct, separated by distinct grooves (z 0.8 x zz 0.5 mm). Frontal shield very finely granular, with larger obvious tuberculations; a single series of marginal pores extending around the distal margin and surrounding the orifice; the periphery of each zooid bordered by a margin of smooth calcification. A single large, round, medial spiramen in a depression towards proximal end of zooid. Primary orifice rounded with large, bluntly triangular, lateral condyles; operculum distinct; secondary orifice oval, surmounting a short, thin peristome. A large, adventitious avicularium always present, midproximal to orifice, between orifice and spiramen; with expanded semicircular proximal area; rostrum setiform, directed

36 70 K. J. TILBROOK ETAL. distolateral to orifice around which it may curve slightly; no crossbar. There also appears to be a number of other pores, visible in frontal view, surrounding the avicularium-these may be marginal pores associated with the avicularium cystid. Gonozooids not known. Communication via septula at base of vertical walls. Etymology From novus L.-new; of this genus. so-named as the newest species The most characteristic feature of Reptadeonella novissima is the size of the frontal adventitious avicularium and the presence of condyles in the primary orifice. Large avicularia are also known in Reptadeonella fissa (Hincks, 1880b). Some of the specimens of R. joloensis (Canu & Bassler, 1929), a junior synonym of R. fissa, particularly those from Singapore, discussed by Harmer (1957), may be attributable to R. novissima. Primary orifical condyles are known only in Reptadeonella levinseni (Borg, 1940; see Cook & Chimonides, 1981). Thus, R. levinseni and R. novissima make a distinct grouping within Reptadeonella sensu stricto. One very small colony fragment was found in debris recovered from Port Vila Harbour, Efate. SUPERFAMILY LEPRALIELL~IDEA VIGNEAUX, 1949 FAMILY ROMANCHEINIDAE JULLIEN, 1888 GENUS ESCHAROIDES MIWE EDWARDS, 1836 Type species: Cellepora coccinea Abildgaard, ESCHAROIDES LONGIROSTRIS DUMONT Eschamides longimstris Dumont, 1981: 160, fig. 1A. Eschamides longimstris: Hayward, 1988: 293; Hayward & Ryland, 1995a: 548, fig. 8A. This species was originally described from the Sudanese Red Sea but has subsequently been recorded from Mauritius and the Great Barrier Reef. Colonies are very small, rarely exceeding 2-3 mm2 and generally comprising less than a dozen autozooids (Ryland & Hayward, 1992), and seem restricted to cryptic habitats in coral rubble, which might explain the paucity of records. The presence of several colonies in coral rubble from Poanangisu greatly increases its range eastwards. It may yet be found to occur widely throughout the Indo-West Pacific. FAMILY LEPRALIELLIDAE VIGNEAUX, 1949 GENUS CELLEPORARIA LAMOUROUX, 1821 Type species: Cellepora cristata Lamark, Species of this genus, often characterized by large colony size and conspicuous pigmentation, are common in tropical and subtropical seas. Many species have been reported from shallow reef habitats but unfortunately their identification is often very difficult as many are still to be named and those already described are relatively poorly known and inadequately illustrated. CELLEPORARIA PILAEFERA (CANU& BASSLER) (Fig. 13A-C) Holoporella pilaefera Canu & Bassler, 1929: 422, pl. 60, figs 2-6. Celleporaria pilaefera: Hayward, 1988: 343, pl. 16, figs e,f. Colony encrusting, forming a multilaminar sheet. Recumbent autozooids broad, rectangular to irregularly polygonal; frontal shield convex, with a single series of large, conspicuous marginal pores, coarsely nodular in later ontogeny. Primary orifice wider than long, distal border almost straight but arched frontally; short, bluntly tapered condyles present in proximolateral corners; no oral spines, no peristome. A small adventitious avicularium present medially on the proximal edge of the orifice, rostrum elliptical, orientated perpendicular to frontal plane, facing laterally; cystid developing a short apical spike, or sometimes a more substantial column, especially prominent in some autozooids. Vicarious avicularia sparse, distinctive: rostrum narrowest at the crossbar, spoon-shaped and deeply cupped distally. Ovicell asymmetrically developed, typically occupying the distal and one lateral border of the orifice, elongate and projecting high above the orifice; frontal surface flat and membranous, the cap-like calcified portion thick and finely granular. Vertical walls with small uniporous septula. Measurements ( ), means and standard deviations, mm (n = 20). Autozooid length , width 0.36 & Orifice length 0.12 k0.007, width 0.15 ko.01. Small colonies of this species were found in sampies from Erakor and Iririki Islands, Port Vila Harbour,

37 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 71 Figure 13. A-C, Celleporaria pilaefera. A, primary orifice. B, large, raised vicarious avicularium. C, colony view. D-F, Celleporaria inaudita sp. nov. D, colony view. E, primary orifice. F, large, raised vicarious avicularium. Scale bars: A = 0.06 mm; B = 0.18 mm; C = 0.36 mm; D = 0.28 mm; E = 0.05 mm; F = 0.16 mm.

38 72 K. J. TILBROOK ET AL. and Poanangisu, encrusting dead Acrwpora and fragments of tile. It is identical to the Mauritian species attributed to Celleporariapilaefera by Hayward (1988), but it should be noted that C. pilaefera (Canu & Bassler) remains to be adequately redescribed and figured. Harmer (1957) described C. pilaefera from three Siboga stations and listed further material from the Indian Ocean and the Red Sea. However, his account differs in some points from that of Canu & Bassler (1929) and consequently is excluded from the present synonymy until all specimens have been reexamined. Celleporaria pilaefera is similar to C. columnaris (Busk, 1881) but may be distinguished from that species by its proportionately wider orifice, smaller condyles, and conspicuous marginal pores. Further, it lacks the variety of small frontal adventitious avicularia so numerous in C. columnaris. CELLEPORARIA VAGANS (BUSK) Cellepora vagans Busk, 1881: 343; Busk, 1884: 198, pl. 29, fig. 10; pl. 35, fig. 11. Celleporaria vagans: Harmer, 1957: 671, pl. 42, fig. 4; text-fig. 55; Ryland & Hayward, 1992: 256, fig. 16a-c. Celleporaria vagans is most easily characterized by its primary orifice, which contains three thin, pointed denticles, evenly spaced proximally just within its rim. It was the least common of the Celleporaria species found, with only two colonies recorded on small pieces of coral rubble from Erakor Island. Ryland & Hayward (1992) state that although this species is known from Hawaii, Great Barrier Reef, Torres Straits, Indonesia, and the eastern Indian Ocean (albeit from relatively few specimens) they note that it may actually differ from Busks (1881, 1884) material from Crozet Island, and advocate its re-examination. CELLEPORARIA INAUDITA SP. NOV. (Fig. 13Ll-F) Material Holotype: NMH , Erakor Lagoon, Efate, 2 October Paratype: NMH , collection data as for holotype. Other material examined: NHM , NHM , NHM , NHM , NHM , Erakor Lagoon, Efate, 2 October 1990; NHM , NHM ,112, NHM , NHM ,119, NHM ,314, NHM , Port Vila Harbour, Efate. Colony encrusting, multilaminar, developing small nodules on coral substrata and spreading mounds on tile fragments. Autozooids recumbent, broadly oval to hexagonal, convex, with indistinct boundaries. Frontal shield finely nodular, vitreous, with small and rather indistinct marginal pores. Primary orifice slightly wider than long; proximal border gently convex in early ontogeny and slightly arched frontally, thickening in later ontogeny and developing two short, pointed cusps with a shallow, variably developed concavity between. Degree of development of the cusps also varies between autozooids, and in a very few specimens one is replaced by three very much smaller denticulations. No oral spines, no peristome. A small, median suboral avicularium present; rostrum elliptical, its distal rim finely denticulate, orientated perpendicular to the plane of the orifice and facing laterally; apex of cystid thickened in later ontogeny but not developing as a spike, rostra1 rim always distinct close to its apex. Vicarious avicularia sparse; rostrum acute to frontal plane, 0.37 mm long, broadest at the crossbar, narrowly spatulate. the distal rim coarsely denticulate; crossbar without columella, palate with very large foramen. Ovicell a shallow cap-like structure occupying the distal and lateral borders of the orifice; frontal surface flat and membranous, distal portion thickly calcified, nodular. Vertical walls with small multiporous septula. Measurements Holotype, means and standard deviations, mm (n= 20). Autozooid length 0.54 ko.09, width 0.37 & Orifice length 0.13&0.008, width Etymology From inauditus, L.-unheard Celleporaria inaudita is reminiscent of Celleporaria aperta (Hincks, 1882a) but may be distinguished by its primary orifice, with usually two differently sized small cusps, not defining the rather regular sinus seen in C. aperta. It also lacks oral spines, while C. aperta has two to four, often well developed. Finally, the vicarious avicularium of C. aperta has a lanceolate rostrum, quite unlike the narrowly-waisted rostrum seen in C. inaudita. Celleporaria mauritiana Hayward, 1988 is probably referable to C. aperta. of.

39 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 73 Celleporaria inaudita was the commonest Celleporaria species collected. Mound-shaped colonies up to 2 cm diameter were collected from both Erakor and Iririki Islands, Port Vila Harbour, and Poanangisu. Diagnosis GENUS DREPANOPHORA HARMER, 1957 Colony encrusting, unilaminar. Autozooids oval, convex, distinct, primary orifice large, the proximal border with a lyrula, associated with lateral sinuses and/or straight or curved denticles, or condyles. Peristome long, thin and tubular, or shorter and more robust, bearing a single avicularium either on the inside of the peristome or on the outer edge, facing transversely or frontally. The avicularium may produce a ridge down the inside of the peristome, or give rise to the projecting denticle. Spines present or absent. Frontal shield imperforate except for large marginal pores. Ovicell prominent, globular, recumbent on distal autozooid, a foramen on either side of midline. Qpe species: Rhynchozoon incisor Thornely, This genus was introduced by Harmer (1957) and the above diagnosis is based on his account. However, the traditional understanding of Drepanophora has been amended here to include the presence of oral spines (see D. gutta sp. nov.). Of the three species Harmer (1957) attributed to his new genus only two remain, D. incisor (Thornely, 1905) and D. corrugata (Thornely, 1905). The assignment of Drepanophora longiuscula Harmer, 1957 to Drepanophora has been questioned (see discussion of Torquatella duolamellata (Scholz, 1991) comb. nov.? below). The genus Drepanophora now also includes, D. tuberculata (Osburn, 1914), D. mgickae (Brown, 1958), D. uerrucosa Winston & Heimberg, 1986, D. indica Hayward, 1988, and D. gutta sp. n ov. DREPANOPHORA INDICA HAYWARD (Fig. 10C) Drepanophora indica Hayward, 1988: 338, pl. 14, fig. e. Colony encrusting, unilaminar. Autozooids oval, convex, distinct, separated by deep grooves. Primary orifice longer than wide, almost pear-shaped, wider distally; proximal border with a single transversely orientated avicularium on one side, rostrum sharply hooked, and a slight constriction opposite. Peristome thin, low, incomplete proximally. Frontal shield nodular, with large marginal pores. Ovicell prominent, globular, recumbent on distal autozooid, a large elliptical foramen, angled to frontal plane, either side of midline. Winston & Heimberg (1986) described Drepanophora uerrucosa from Indonesia; Hayward (1988) distinguished it from D. indica using criteria of the ovicell and orificial avicularium. Drepanophora indica was originally described from Mauritius; a single ovicellate colony was found at Poanangisu, Efate and was compared with Haywards (1988) type specimen. DREPANOPHORA TUBERCULATA OSBURN (Fig. 1OD) Rhynchozoon tuberculatum Osburn, 1914: 200, textfig. 9. Rhynchozoon tubemulatum: Osburn, 1940: 442; 1947: 39; 1952: 461, pl. 54, fig. 6. Drepanophora tuberculata: Cook, 1968: 205; Cook, 1985: 70, 180, text-fig. 47, pl. 20, fig. F. Colony encrusting, unilaminar. Autozooids oval, convex, distinct, separated by deep grooves. Primary orifice longer than wide, wider distally, proximal border with a single transversely orientated denticle below a single frontally facing avicularium on the edge of the thin entire peristome; avicularium rostrum acute triangular. Frontal shield slightly nodular, imperforate except for large marginal pores. Ovicell prominent, globular, recumbent on distal autozooid, a large elliptical foramen, perpendicular to frontal plane, either side of midline. Drepanophora tubemulata is very similar to D. indica, but it appears less robust and the frontal shield is less granular, the peristome is taller and thinner, and it has a frontally facing, rather than transversely facing, avicularium. Also, the ovicell foramina are angled to the frontal plane differently; perpendicular in D. tuberculata and more acute in D. indica. Cook (1985) described and figured the ancestrula of D. tubemulata from her Ghanaian material. Drepanophora tuberculata appears to have an almost circum-tropical distribution: Gulf of Mexico, West Indies, West Africa, Southwest and eastern Pacific.

40 74 K. J. TILBROOK ET AL. Found on coral rubble from Iririki Island and Poanangisu, Efate, this was the commonest of the Drepanophora species and was found associated with Zbbrquatella duolamellata (Scholz, 1991) comb. nov. DREPANOPHORA GUTTA SP. NOV. (Fig. 1 OE,F) Material Holotype: NHM , Port Vila Harbour, Efate, Vanuatu, October Colony encrusting, unilaminar. Autozooids oval, convex, distinct, separated by shallow grooves. Primary orifice longer than wide, wider distally, the proximal border with a deep, narrow, anvil-shaped lyrula. Peristome very long, tubular, with a notch distally that accommodates 2 distal oral spines; a single frontally facing avicularium present on the rim of the peristome, rostrum acute triangular. Frontal shield very slightly nodular, imperforate other than for 5 or 6 large marginal pores, generally proximal. Ovicell prominent, globular, recumbent on distal autozooid, a small teardrop-shaped foramen, perpendicular to frontal plane, either side of midline, distally situated. Oral spines not seen in brooding zooids. Measurements Holotype, means and standard deviations, mm (n= 30). Autozooidal length 0.33 & 0.04; width 0.22 f Etymology From guttum, L.-a narrow-necked flask; alluding to the appearance of the peristome. Drepanophora gutta is included in the genus Drepanophora because it has a large peristome with an associated avicularium, an entire frontal shield with marginal pores, a globular ovicell with a pair of foramina, and an orificial lyrula. However, it differs from all other species assigned to this genus in the fact that it possesses two oral spines, the taxonomic significance of which is far outweighed by the features just mentioned. Drepanophora gutta appears close to Rhamphostomella von Lorenz, 1886, as described by Gordon (1984). Drepanophoragutta was represented by a single colony found encrusting a small piece of coral debris from Port Vila Harbour, Efate. SUPERFAMILY SMI'ITINOIDEA LEVINSEN, 1909 FAMILY BITECTIPORIDAE MACGILLIVRAY, 1895 GENUS METROPERIELLA CANU & BASSLER, 1917 m e species: Schizoporella lepralioides Calvet in Jullien & Calvet, METROPERIELLA MONTFERRANDII (AUDOUIN) (Fig. 15F) Flustra montferrandii Audouin, 1826: 240. Codonellina montferrandii: Harmer, 1957: 1049, pl. 69, figs 25, 26, 30 (cum syn.); Gordon, 1984: 77, pl. 26A. Material examined NHM , NHM ,59, Port Vila Harbour, Efate, 4-5 m; NHM ,61, Torres Straits. Colony encrusting, forming small unilaminar patches; pale pink when dried. Autozooids irregularly polygonal, convex, distinct, separated by grooves ( x mm). Primary orifice oval, longer than wide; broad, deep anter separated from shallow poster by small triangular condyles. Frontal shield evenly perforated by numerous small pores. No oral spines. A single, median adventitious avicularium, acute rostrum raised slightly, directed proximally or obliquely laterally. Ovicell prominent with thin-walled calcification and scattered pores of various sizes. This species has no type material and the plate figure of Savigny [1817] in Audouin (1826) is somewhat idealized. It has been accorded an almost global distribution in warm-temperate to tropical waters except the Atlantic Ocean. Re-examination of all material attributed to M. montferrandii, with the designation of a neotype specimen from the Egyptian Red Sea, would stabilize the species and also aid in clarifying the debate over possible conspecificity with M. lepralioides (see Gordon, 1984, 1994 for discussion). The small fragments of material from Port Vila Harbour, Vanuatu, differ from Red Sea specimens in having a shorter, wider orifice and a longer, more acute suboral avicularium. Although certain Red Sea specimens have larger, proximally pointing, spatulate frontal avicularia in place of the more normal median avicularia, these were not seen in the Vanuatu material but were present in specimens from the Torres Strait.

41 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 75 FAMILY WATERSIPORIDAE VIGNEAUX, 1949 GENUS WATERSIPORA NEVIANI, 1895 WATERSIPORA SUBOVOIDEA SENSU W R (Fig. 18F) Dakaria subovoidea (dorbigny) Harmer, 1957 (in part): 1022, pl. 69, fig. 12. Watersipora subovoidea: Dumont, 1981: 636. 'Watersipora subouoidea'sensu Harmer: Soule & Soule, 1976: 308, pl. 3, figs 4-6; Winston & Heimberg, 1986: 17, figs There has been much discussion about the taxonomic status of this genus and species attributed to it, Watersipora subovoidea in particular (e.g., Ryland, 1974; Soule & Soule, 1976; Winston & Heimberg, 1986; Gordon, 1989b; Soule et al., 1995). All species attributed to Watersipora need to be reviewed before any synonymies can be resolved. Watersipora su bovoidea has been recorded from the Red Sea and Indonesia, but the true extent of its geographical range will remain unclear until the genus has been reviewed and revised (Winston & Heimberg, 1986). Iridescent black dried colonies were found on a shell at Vila Waterfront, Erakor Lagoon, Efate. FAMILY SMI'ITINIDAE LEVINSEN, 1909 GENUS PARASMITTINA OSBURN, 1952 Type species: Lepralia jeffreysi Norman, Colonies of species of Parasmittina are often small, but they may be abundant in shallow reef habitats where their diversity is usually high (Ryland & Hayward, 1992). Species are generally distinguished by features of the primary orifice, such as the size and shape of the lyrula and condyles, denticulation of the distal margin, numbers of distal oral spines, and the shape, size, and orientation of adventitious avicularia. PARASMITTINA GALERITA RYLAND & HAYWARD (Fig. 14A,B) Parasmittina galerita: Ryland & Hayward 1992: 270, fig. 21d-f. Parasmittina latiavicularia: Ryland & Hayward 1992: 270, fig. 22a-c. Colony encrusting, multilaminar. Autozooids rectangular to irregularly polygonal, convex, separated by thickened sutures; x mm. Frontal shield coarsely nodular, with a single series of large marginal pores. Primary orifice longer than wide; lyrula short, rectangular, with straight distal edge and cusped corners, occupying about half proximal orifice width; condyles rounded, downcurved, finely denticulate. 1-2 distal oral spines present in early ontogeny only. Peristome consisting mostly of prominent paired lateral lappets, deeply concave proximally. Avicularia polymorphic, numerous, mostly lateral-oral, single or paired, with acutely triangular, finely denticulate rostrum, this directed medially and ascending the side of the peristomial lappet; similar avicularia often present distal to ovicell; similar-sized avicularia with parallelsided, distally rounded rostrum typically developed along margins of zooids, proximolateral to peristome and proximally directed. Enlarged avicularia frequent: proximal portion lateral to orifice, rostrum proximally directed, extending along most or all of the margin of the zooid, broadly and asymmetrically spatulate, with upturned, irregularly toothed rim. Ovicell about as wide as long, with numerous small frontal pores; peristome forms a thin rim curving above the ovicell aperture. Parasmittina galerita is very similar to Parasmittina latiauicularia (Kirkpatrick, 1888) with which it has been confused previously. Examination of Kirkpatricks holotype specimen, however, shows that the two species differ in several aspects: autozooids in I! latiavicularia are 50% larger than in l? galerita; while the primary orifices are almost identical in size and shape, the denticulate condyles seen in both species are smaller in l? galerita; the characteristic paired lateral-oral avicularia of l? latiavicularia are more acute and constant whereas in? galerita they are shorter and often paired with an oval, shoe-shaped avicularium; consequently the peristome is larger in l? latiauicularia and entire. Parasmittina galerita has large spatulate avicularia, not seen in I! latiavicularia. The frontal shield in l? latiavicularia is finely granular, in l? galerita coarsely nodular; finally, the ovicells differ, with l? latiauicularia having smaller, more separated pores compared to the larger more closely packed pores of l? galerita. Parasmittina latiavicularia is thus restricted to Mauritius. Another very similar species, l? leviavicularia Soule & Soule, 1973, may prove to be synonymous with l? galerita but the type material has not been examined. If the two species are found to be the same then the known range of l? galerita is extended but the Soules' name has taxonomic precedence.

42 76 K. J. TILBROOK ET AL. Described from Heron Island, off Queensland, Australia, Ryland & Hayward (1992) noted that one of Harmer's (1957) specimens of Smittinaparsevalii (Audouin, 1826) from Makassar Strait was the same as their new species. Several colonies of Parasmittina galerita were found on coral rubble at Iririki Island, reinforcing its presence in the western Pacific. PARASMITTINA HASTINGSAE SOULE & SOULE (Fig. 14C,D) Parasmittina hastingsae Soule & Soule, 1973: 417, fig. 9C,D. Parasmittina hastingsae: Winston & Heimberg, 1986: 26, figs 59-63; Ryland & Hayward, 1992: 268, fig. 21a-c; Hayward & Parker, 1994: 63, fig. 4A-F. Smittina trispinosa var. spathulata Hincks, 1884c: 284, pl. 9, fig. 4. Colony encrusting, multilaminar. Autozooids oval to irregularly polygonal, convex, separated by distinct sutures; x mm. Frontal shield coarsely nodular, with large and conspicuous marginal pores. Primary orifice rounded, about as wide as long; lyrula short, straight-edged, occupying about half proximal orifice width; condyles thin, acute, downcurved. Spines variable: 1-3, or often absent, present in early ontogeny only, bases later obscured. Peristome developed as prominent flared, lateral lappets, rim deeply embayed proximally, incomplete distally except in ovicelled zooids. Avicularia sporadic, infrequent in present material but displaying polymorphism characteristic of the species: most commonly with an acute triangular rostrum (of varying length), randomly directed, generally one lateral-oral with others found anywhere on frontal shield; the second, oval, shoe-shaped, generally proximally directed, lateral-oral; or gigantic, seemingly in place of the oval avicularium, with a swollen cystid, rostrum parallel-sided, linguiform, sharply serrated distally, proximally directed. Ovicell about as wide as long, with numerous irregular frontal pores; peristome forms a low ridge frontally, an ooecial cover develops distally. Parasmittina hastingsae is characterized especially by its primary orifice, which has a shallow quadrate lyrula and very small, triangular, lateral condyles. In some specimens, including part of the type series and material from Heron Island, a proportion of the autozooids in the colony develops obscure denticulation on the mid-distal portion of the orifice rim. The only slight difference between the colony from Vanuatu and a paratype specimen of Parasmittinu hastingsae (NHM , Hawaii) is that the gigantic adventitious avicularia on the Vanuatu specimen are longer than those found on the paratype specimen from Hawaii. Described from Hawaii, where it was reportedly particularly common, this species has been found subsequently from Komodo, Indonesia (Winston & Heimberg, 1986), Victoria, Australia (Hincks, 1884b) and Heron Island, Great Barrier Reef (Ryland & Hayward, 1992), where again it was reported as occurring abundantly on the reef flat. One small colony of Parasmittina hastingsae was found on a piece of coral rubble from Poanangisu, Efate. PARASMITTINA SERRULA SOULE & SOULE (Fig. 14 E, F) Parasmittina serrula Soule & Soule, 1973: 386, fig. 3D-F. Parasmittina serrula: Winston, 1984: 23, fig 45; Gordon, 1984: 96, pl. 35B,C; Scholz, 1991: 325, pl. 20, figs 1,2,4; Ryland & Hayward, 1992: 272, figs 23e,f, 24a; Gordon & d'hondt, 1997: 21, fig. 27. Parasmittina serrula is readily identified by the morphology of the orifice and associated structures, and by two distinctive types of avicularia. The primary orifice is slightly wider than long and has a variably denticulated distal rim; the lateral condyles are rounded and downcurved, with finely toothed edges visible with high-power light microscopy. The anvil-shaped lyrula is broad basally, tapering towards a curved distal edge, the corners of which are generally sharply produced. There are three or four stout distal oral spines, five or six in periancestrular zooids. The peristome is well developed proximally and laterally but has a deep midproximal notch. The most frequently occurring adventitious avicularia are elongate, narrow and parallel-sided, rounded at tip, lateral-suboral, single or paired, of similar shape but uneven lengths, and proximally directed characteristically, one of these lies on its side, displaying a downcurved, finely denticulate rostra1 rim. Other enlarged giant avicularia are also diagnostic: the slender crossbar is situated adjacent to the orifice, the rostrum extending to the proximal end of the autozooid the rostrum is narrow proximally, slightly flared laterally, with a deeply toothed rim. Some colonies lack one or other of the avicularia from large areas. Ovicells are siightiy wider than long, raised, recumbent on the frontal shield of

43 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 77 Figure 14. A,B, Parasmittina galerita. A, note the large spatulate adventitious avicularium, left. B, primary orifice, note the denticulate oral condyles. C,D, Parasmittina hastingsue. C, colony growing edge. D, primary orifice, note the smooth oral condyles. E,F, Parasmittina serrula. E, note the denticulate rostrum to the adventitious avicularium, centre. F, primary orifice. Scale bars: A =0.25 mm; B = 0.05 mm; C = 0.50 mm; D = 0.05 mm; E = 0.15 mm; F = 0.05 mm.

44 78 K. J. TILBROOK ET AL. the distal autozooid, with about 20 small, tubiform pores frontally, the orificial peristome encroaching onto its frontal surface in later ontogeny, forming a complete rim above its aperture. Parasmittina serrula is the most widely recorded of the species of Parasmittina found in Vanuatu. Described originally from Hawaii, P. serrula has been reported also from Belize and Jamaica (Winston, 1984), the Great Barrier Reef (Ryland & Hayward, 1992), the Philippines (Gordon & dhondt 1997), and the Kermadec Ridge (Gordon, 1984). As in Hawaii (Soule & Soule, 1973) it was by far the commonest species of Parasmittina in Vanuatu, found at Poanangisu, Iririki Island, and Port Vila Harbour, Efate. GENUS PLEUROCODONELLINA SOULE & SOULE, 1973 Type species: Pleumcodonellina lahainae Soule & Soule, PLEUROCODONELLINA SIGNATA (WATERS) Smittia signata Waters, 1889: 17, pl. 3, figs 4-6. Smittina signata: Harmer, 1957: 928, pl. 63, figs Parasmittina signata: Winston & Heimberg, 1986: 21, figs Pleumcodonellina signata: Ryland & Hayward, 1992: 273, fig. 24b. Young zooids have one or two oral spines. No large spatulate avicularia have been found on colonies from Vanuatu, which, although small, still bear ovicells. Quite common in the shallow reef environments sampled at Heron Island, Great Barrier Reef, Pleumcodonel lina signata has an almost circumtropical distribution from the Indo-West Pacific and Mauritius, East Africa, West Africa, and the Caribbean. In Vanuatu it was found at Iririki Island and Port Vila Harbour, Efate. GENUS SMInOrDEA OSBURN, 1952 Type species: Smittoidea pmlifica Osburn, SMITlVIDEA PACIFICA SOULE & SOULE (Fig. 18E) Smittoideapacifica Soule & Soule, 1973: 380, fig. 1E-H. Smittoidea pacifica: Winston, 1986: 28; Ryland & Hayward, 1992: 268, figs 24e,f. Harmer (1957) included specimens of this species in his description of Smittoidea levis (Kirkpatrick, 1890) according to Soule & Soule (1973). Distri butwn Described from the Hawaiian and Galapagos Islands, Smittoiclea pacifica is also known from the Caribbean and Venezuela (Winston, 1986), and from Heron Island, Great Barrier Reef (Ryland & Hayward, 1992). It was found encrusting coral rubble and shell at Port Vila Harbour and Iririki Island, Efate. FAMILY TEUCHOWRIDAE NEVIANI, 1895 TORQUATELLA GEN. NOV. Diagnosis Colony encrusting, multilaminar. Autozooids with convex, cryptocystidean frontal shield, perforated by a number of large pores, predominantly around the margin. Primary orifice suborbicular; anter large, smoothrimmed, rounded, separated from the wide, shallow poster by small, squared, lateral condyles. Flared peristome bears a small, rounded, suboral avicularium on inner edge, laterally or frontolaterally directed. No spines. Ovicell hyperstomial, globular; single, oval, transverse, frontal foramen through smooth ectooecium exposing an entire, striated, endooecium. Interzooidal communication by multiporous septula. Qpe species: Osthimosia duolamellata Scholz, Etymology From torquata, L.-adorned with a collar; -ellus, L.- diminutive, small, alluding to the flared orificial peristome. The wide sinus and lack of vicarious avicularia, seen in other species of Osthimosia (see Gordon, 1989b), suggest that Osthimosia duolamellata Scholz (1991) is incorrectly assigned to Osthimosia Jullien, Tbbrquatella gen. nov. is introduced for its reception. Several other previously described species may be included in Tbrquatella, including Reptescharellina cornuta Gabb & Horn, 1862, originally found in the Pleistocene of Santa Barbara, California (see Soule et al., 1995 for further discussion), Schizoporella dissimilis Osburn, 1952, from the Gulf of Califoria and Galapagos Islands, and an undescribed species from Pearson Island, South Australia (in South Australian Museum), photographed by the late Shane Parker. In the pseudoporous shield and peristomial rim, Torquatella somewhat resembles Lagenicella Cheetham

45 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 79 & Sandberg (see Gordon, 1984) which, however, has a more perforated shield. For this reason, our new genus is provisionally included in the Teuchoporidae. TORQUATELLA DUOLAMELLATA (SCHOLZ) COMB. NOV. (Fig. lla,b) Osthimosia duolamellata Scholz, 1991: 302, pl. 11, figs 1-6. Colony encrusting, multilaminar. Autozooids elongate polygonal, quincuncially arranged in early ontogeny, though obscured later in ontogeny by secondary calcification and irregularly orientated frontal budding; c x Frontal shield convex, granular, with up to a dozen fairly large pores, predominantly around the margin. Primary orifice suborbicular; anter large, smooth, rounded, separated from the wide, shallow poster by small, squared, lateral condyles. Peristome flared, with a small, rounded proximal avicularium on inner surface, laterally or frontolaterally directed. Ovicell hyperstomial, globular, with a single, oval, transverse, frontal foramen. In his original description Scholz (1991) described the ancestrula as tatiform, pointing out that it is often difficult to observe, being obscured by the five periancestrular zooids. It is in fact schizoporelloid, with six oral spines around the D-shaped orifice. Initially, the primary orifice sits within a slightly raised rim, however, this is superseded by a large chimney-like flared peristome. The peristome is connected to the parent autozooid via a multiporous septulum and seems to develop from the cystid producing the proximal oral avicularium (seen on the inner surface of the peristome), and originating from one of the two most distolateral pores. The avicularian cystid also appears to cover most of the autozooidal frontal shield in secondary calcification. This happens early in ontogeny. The ovicell seems to be formed in a similar way, as a polymorph which evaginates from the distal edge of the oral area (distal wall) of the maternal zooid. It rests on the frontal shield of the distal autozooid producing the distal part of the peristome and being incorporated into the secondary calcification of the distal autozooid.?brquatella duolamellata is superficially similar to Drepanophora longiuscula Harmer, Both have a schizoporelloid ancestrula surrounded by five periancestrular zooids. The primary orifices are almost identical, as is the development of a flared peristome, and accompanying avicularium, from a distolateral pore. However, D. longiuscula has an ovicell with two fenestrae, while T duolamellata has only one. More significantly, however, the frontal shield in D. longiuscula is entire with only about six areolar septular pores whereas T duolamellata has up to a dozen pseudopores, mainly marginally placed (though some of these might function as septular pores). These two criteria, especially the latter, question their congeneric status. Gordon (1993b) doubted the assignment of Harmer s (1957) species to Drepanophora but in the absence of any other material this problem remains unresolved. Examination of the inside of the frontal shield of T duolamellata shows that this is not umbonuloid, i.e. no ring scar is present. Scholz (1991) described the growth and ecology of this species. Found on coral rubble from Poanangisu Erakor and Iririki Islands, T duolamellata was first described from the fringing reef flats of Cebu, the Philippines. SUPERFAMILY SCHIZOFQRELLOIDEA JULLIEN, 1883 FAMILY SCHIZOPORELLIDAE JULLIEN, 1883 GENUS SCHIZOPORELLA HINCKS, 1877 Qpe species: Schizoporella unicornis Johnston, SCHIZOPORELLA DUNKERI (REUSS) (Fig. 15A) Cellepora dunkeri Reuss, 1848: 90, pl. 10, fig. 27. Schizopowlla unicornis form longimstris Hincks, 1886: 266, pl. 10, fig. 2. Schizopowlla longimstris: Hayward, 1976: 320, fig. 1; Hayward & Ryland, 1979 (in part): 173, not fig. 73. Schizopowlla dunkeri: Hayward & Ryland, 1995b: 39, pls 2, 3. Colony encrusting, often extensive. Autozooids large, rectangular, convex, distinct, separated by deep grooves. Frontal shield evenly perforated by numerous round pores, each in a shallow depression; a suboral umbo may be present. Primary orifice as wide as long, almost circular; narrow, slit-like median sinus, often with a rounded, proximal broadening; condyles broad and flat. No oral spines. Adventitious avicularium single, lateral to the orifice and level with the proximal border or sinus; cystid large and prominent; rostrum abruptly tapered distal to complete crossbar, with a slender, acuminate tip; mandible acutely triangular, laterally or distolaterally orientated. Ovicells prominent, globular, regularly perforated.

46 80 K. J. TILBROOK ET AL. Schizoporella dunkeri, described from the Miocene deposits of the Vienna Basin of Austria (Reuss, 1848), shows a relict Tethyan distribution. It is known from the Pliocene Coralline Crag of Britain and Pliocene deposits in Belgium and The Netherlands (Bishop & Hayward, 1989), is extant from the Canary Islands, Madeira, and the Isles of Scilly, and is common throughout the Mediterranean, especially the Aegean and Adriatic. It purportedly also occurs in the Red Sea (Hayward & Ryland, 1995b). Our record of two small colony fragments of Schizoporella dunkeri from debris at Erakor Island, Vanuatu is the most easterly and most tropical record to date. SCHIZOPORELLA ERRATA (WATERS) (Fig. 15B) Lepralia errata Waters, 1878: 11, pl. 1, fig. 9. Schizoporella errata: Hayward & Ryland, 1979: 170, fig. 68; Brock, 1985: 46, fig. 2C; Gordon & Mawatari, 1992: 31, pl. 9, fig. B. Colony encrusting, unilaminar or multilaminar, or developing erect, foliaceous, branched or cylindrical growth; whitish-pink to reddish-brown in life. Autozooids rectangular or lozenge-shaped, flat or slightly convex, separated by shallow grooves, in linear series; in multilaminar colonies, more disorientated in frontally budded additional layers. Frontal shield evenly perforated by numerous round pores, each in a shallow depression. Primary orifice slightly wider than long, appearing almost circular; with broad and shallow U- shaped median sinus. No oral spines. Adventitious avicularium single, situated lateral to the orifice and level with the proximal border or sinus; cystid prominent, mandible acute triangular, angled to frontal plane, distolaterally orientated. Ovicells prominent, globular, regularly perforated. Schizoporella errata is a well-known fouling species with many records from harbour installations or the hulls of ships. Compared with many other fouling community species S. errata displays a rather catholic choice of substratum, from pier pilings and shell, to algae and mangroves. Gordon & Mawatari (1992) document its distribution in New Zealand and the Pacific, dating the first record in New Zealand at about The increase in cruise-liner traffic to Vanuatu over the last 20 years or so may have promoted the introduction of this species (as well as others) from other localities in the Southwest Pacific area. Schizoporella errata is a widespread warm temperatesubtropical species. Originally described from Naples, Italy, it is found throughout the Mediterranean, West Africa, eastern Canada, eastern coasts of the Americas, from North Carolina through the Caribbean to Brazil. Pacific coast of North America, Red Sea, Persian Gulf, South Australia, and New Zealand. It was found several times in Port Vila Harbour, on old wall tiles. GENUS ESCHARINA MILNE EDWARDS, 1836 Type species: Eschara uulgaris Moll, ESCHARINA PESANSERIS (SMI~) (Fig. 17A,B) Hippothoa pes anseris Smitt, 1873: 43, pl. 7, figs 159, 160. Escharina pesanseris: Harmer, 1957: 998, pl. 67, figs 12-14,18,19; Winston, 1984: 26, figs 53-55; Scholz, 1991: 311, pl. 14, fig. 4, Ryland & Hayward, 1992: 264. Escharina pesanseris is a well-known species that apparently has a circumglobal distribution within warm-temperate and tropical shallow waters. It is generally rather inconspicuous, with most colonies comprising a dozen or fewer zooids, many of which may be ovicellate. It was quite common at Port Vila Harbour on coral rubble, and was also found at Iririki Island. GENUS STYLOPOMA CANU AND BASSLER, 1920 Type species: Eschara spongites Pallas, This speciose genus has a pan-tropical distribution in shallow, generally reefal, habitats. A review of the Indo-Pacific species of Stylopoma appears in the companion paper (pp. 1-34). STYLOPOMA NOVUM TILBROOK (Fig. 15C) Stylopoma nouum Tilbrook, 2001: companion paper, p. 24, Fig. 9D-F. Colony encrusting, multilaminar, often extensive. Autozooids hexagonal or irregularly polygonal, flat or slightly convex, separated by distinct grooves;

47 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 81 Figure 15. A, Schizoporella dunkeri. B, Schizoporella errata. C, Stylopoma nouum. D, Stylopoma velatum. E, Stylopoma uilaensis. F, Codonellina montferrandii. Scale bars: A = 0.50 mm; B = 0.40 mm; C = 0.60 mm; D = 0.50 mm; E = 0.40 mm; F = 0.40 mm.

48 82 K. J. TILBROOK ET AL. c x 0.42 mm. Primary orifice wider than long, with a smooth distal border; sinus deep, U-shaped, occupying half of the straight proximal border; condyles smooth, small and rounded, inconspicuous with a frontal lip. Frontal shield evenly and densely perforated with large round pores; marginal pores and lateral walls distinct. Adventitious avicularium, single, proximally latero-oral; rostrum bluntly triangular, slightly acute to frontal plane, distolaterally directed. Vicarious avicularia sporadic, larger than autozooids, with large spatulate rostrum. Ovicells globular, prominent, calcification similar to frontal shield, aperture an inverted C-shape, labellum spade-shaped with a conspicuous median suture line. Stylopoma nouum is characterized by its wide sinus, inconspicuous smooth condyles and its extremely elaborate ovicellular labellum. Stylopoma nouum is very similar to S. thornelyae Livingstone, 1926 from the Great Barrier Reef but they differ in their orifices. Stylopoma nouum has a smooth anter while S. thornelyae has a denticulate anter. The ovicellular labellum in S. thornelyae is far straighter than in S. novum. Stylopoma nouum was the least common of the three commonest Stylopoma species found in Vanuatu; it was found encrusting small pieces of coral rubble in shallow water around Iririki Island, Efate. A small colony has also been found in material from Cleveland Bay, Great Barrier Reef. STYLOPOMA VELATUM TILBROOK (Fig. 150) Stylopoma duboisii: Ryland & Hayward, 1992: 265, fig. 20a,b. Stylopoma velatum Tilbrook, 2001: companion paper, pp Fig. 11D-F. Colony encrusting, unilaminar to multilaminar. Autozooids hexagonal or irregularly polygonal, convex, separated by distinct grooves; c x 0.36 mm. Primary orifice D-shaped, wider than long with a straight proximal border, with a deep, slit-like sinus; condyles denticulate, occupying entire proximal edge of orifice. Frontal shield perforated by large round pores and larger marginal pores. Adventitious avicularium single, lateral-oral; rostrum bluntly triangular, slightly acute to frontal plane, distolaterally directed. Vicarious avicularia sporadic, autozooid-sized, with large spatulate rostrum; lacking a palate, but with a complete, slender crossbar. Ovicells globular, very densely perforated, aperture oval, labellum incomplete. Stylopoma velatum is characterized by its D-shaped orifice and slit-like sinus. It differs from S. duboisii (Audouin, 1826), to which Heron Island specimens were attributed by Ryland & Hayward (1992), in having a more semicircular orifice, distinctly wider than long, and condyles which are shorter and more markedly denticulate. The ovicells also differ, the ovicell aperture in S. uelatum is oval and the labellum split whereas in S. duboisii the aperture is D-shaped and the labellum entire. There is a lack of adventitious avicularia on some colonies from Vanuatu, but there are still rare vicarious avicularia. This was the commonest of the three common Stylopoma species found encrusting coral debris on Erakor and Iririki Islands, Vanuatu. It has also been recorded from the Great Barrier Reef to Tahiti (companion paper, pp ). STYLOPOMA VILAENSIS TILEIROOK (Fig. 15E) Stylopoma vilaensis Tilbrook, 2001: companion paper, pp Fig. 12A-C. Colony an extensive, often multilaminar, encrusting sheet. Autozooids irregularly polygonal or hexagonal, slightly convex, separated by distinct grooves; c x 0.41 mm. Frontal shield evenly perforated by very small round pores; the marginal pores and lateral walls are conspicuous. Primary orifice wider than long, D-shaped, the straight proximal border with a V- shaped median sinus; condyles smooth, rectangular, distinct, occupying most of the proximal border each side of the sinus and tapering off medially into it, One adventitious avicularium proximolateral to the orifice, acute to frontal plane, distolaterally directed; mandible short, triangular. Additional adventitious avicularia sometimes found elsewhere on frontal shield. Vicarious avicularia also present, as large as autozooids, with a broadly spatulate mandible distally directed. Ovicells prominent, flattened, densely porous with a slit-like oval aperture and entire labellum. Stylopoma vilaensis is characterized by the shape of its orifice, almost Y-shaped when viewing the sinus and condyles together, and the limited number of pores in its frontal shield.

49 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 83 Stylopoma vilaensis has been found at only two localities in Vanuatu (Iririki Island and Port Vila Harbour), on pieces of coral debris. It has also been reported from Indonesia (companion paper, p. 31). STYLOPOMA VIRIDE (THORNELY) Schizopomlla viridis Thornely, 1905: 116, plate, fig. 3. Stylopoma viride: Harmer, 1957 (cum syn.): 1036, pl. 74, figs Stylopoma viride: Winston & Heimberg, 1986: 21, figs 48-50; Ryland & Hayward, 1992: 265, fig. 2d; Tilbrook, 2001: companion paper, pp. 12, 14, 15. Fig. 5A-C. Stylopoma grandis Canu & Bassler, 1929: 316, pl. 37, figs 1-3. Stylopoma viride is perhaps the most easily recognized of the Indo-Pacific Stylopoma species (see companion paper, pp. 12, 14, 15). Not only is it generally green in colour, as its name suggests, but its zooids are amongst the largest known in the genus (c x 0.70mm). It forms massive multilaminar colonies up to 100 cm2 in area. A single small colony of Stylopoma viride was found, from Poanangisu, Efate. FAMILY LACERNIDAE JULLIEN, 1888 GENUS NIMBA JULLIEN IN JULLIEN & CALVET, 1903 Diagnosis Colony encrusting, uniserial. Frontal shield of autozooids with marginal pores. Orifice with sinus, bordered by a raised peristome. Avicularia adventitious, rare or absent. Ovicells hyperstomial, not closed by the zooidal operculum. Type species: Nimba praetexta Jullien in Jullien & Calvet, NIMBA SAXATILIS HAYWARD & RYLAND (Fig. 80) Nimba saxatilis Hayward & Ryland, 1995a: 558, fig. 11D,E. Colony encrusting; autozooids in uniserial chains which branch at irregular intervals. Autozooids elongate, oval, broadest at the distal end; x mm. Primary orifice about as wide as long; poster broadly concave, separated from anter by small, knob-like condyles. Peristome well developed, forming an erect tube completely encircling the orifice, its rim thin and evenly flared. Frontal shield convex, thick, nodular; bordered by a single or double series of small marginal pores. No spines or avicularia found. The structure of the ovicell is still not known. Nimba saxatilis appears to be rather common on coral rubble, but is easily overlooked because of its cryptic habitat, uniserial growth form, and hyaline calcification. Described from Heron Island, Great Barrier Reef, Nim ba saxatilis was found in Port Vila Harbour, Efate. This species is probably commoner in shallow reef habitats than these records suggest. ECHINOVADOMIDAE FAM. NOV. Diagnosis Colony encrusting, unilaminar. Autozooids with perforate cryptocystidean frontal shield. Primary orifice suborbicular, longer than wide; anter deep, with smooth inner edge separated from a shallow concave poster by proximally incurved condyles. Large flared peristome of fused lobes or processes. No spines. No avicularia. Ovicell hyperstomial, globular, embedded in frontal shield of distal autozooid; ectooecium membranous; spinose calcareous endooecium growing frontally from both sides of the orifice, the two sides meeting and joining during ontogeny, leaving two foramina of which the proximal one is larger; no other perforations; not closed by maternal zooid. Ancestrula not observed. Etymology From echinatus, L.-spiny, prickly; ovum, L.-egg; domus, L.-house, alluding to the appearance of the ovicell in the type species of the genus. ECHINOVADOMA GEN. NOV. Diagnosis With the characters of the family. Type species: Echinovadoma anceps gen. et sp. nov. This genus and species are named for the very characteristic ovicell, the endooecium of which appears to grow frontally from either side to meet at the midline distally, leaving a single rounded median distal opening, retained through to completion of the ovicell which sutures in a proximal direction. The endooecial

50 84 K. J. TILBROOK ETAL. calcification is covered in spinose processes, often large, which become less obvious with addition of calcification in ontogeny. The ovicell is unique among cheilostomes. The occurrence of a spinose endooecium with membranous ectooecium is reminiscent of that seen in some species of Characodoma Maplestone, 1900, e.g. C. glabra Gordon & dhondt, The ovicell of Echinouadoma has large foramina instead of small perforations, however, and the frontal shield is pseudoporous, which precludes close association with the Cleidochasmatidae. The primary orifice is similar in appearance to that seen in Characodoma Maplestone, Yrbozoon Gordon, 1989b, Anchicleidochasma Soule, Soule & Chaney, 1991, Calyptooecia Winston, 1984 and Fedorella Silh, 1947, all cleidochasmatids, but these genera possess avicularia, not seen in Echinouadoma. The species illustrated by Gordon & dhondt (1997: fig. 128) as Characodoma sp., has a wide orificial poster, a tuberculate and perforate frontal shield, and no avicularia and so may perhaps belong to Echinouadoma. Lepraliella granulata Canu & Bassler, 1929, from the Philippines, also possesses attributes, such as the form of the ovicell, which are characteristic of Echinouadoma; however, L. granulata possesses adventitious avicularia. The combined characters of the frontal shield and ovicell preclude inclusion of Echinouadoma in any known cheilostome family. In consequence, a new family, Echinovadomidae, is established here. Two further species of Echinouadoma have been found in material, from the Banda Sea, donated to the Natural History Museum by Dr Gerhard Cadee. These as yet undescribed species, Echinouadoma sp. 1 (NHM ) and E. sp. 2 (NHM ) are similar to E. anceps gen. et sp. nov.in the structure of their ovicells (distal foramen) and their frontal shields. Echinouadona sp. 1, however, has fewer and larger frontal pores, a less spinose ovicell, and a peristome producing approximately five large, frontally facing, processes. Echinovadoma sp. 2, in contrast, has a smoother frontal shield with more frontal pores than E. anceps, as well as a smoother ovicell and a peristome that produces five lobes (cf. three in E. anceps). A fuller description of these two species appears in Tilbrook (in press). ECHINOVADOMA ANCEPS GEN. ET SP. NOV. (Fig. 16C-E) Material Holotype: NHM , Poanangisu, Efate, Vanuatu. Paratypes: NHM ,216, NHM , NHM , NHM , NHM , same locality data as holotype. Autozooids small, distinct, irregularly polygonal, separated by shallow grooves. Frontal shield slightly convex, tuberculate, perforated by numerous small pores that are not enlarged along the margins. Primary orifice suborbicular, longer than wide; anter deep, wide, with smooth inner edge separated from the shallow, concave, narrower poster by small, proximally incurved, condyles. Large, flared peristome developed proximally and laterally, as one midproximal and two lateral lobes, appearing tripartite; not developed distally. Ovicell hyperstomial, globular, embedded in frontal shield of distal autozooid; ectooecium membranous; endooecium very spinose initially, becoming less so when the two sides meet and join, separating two foramina, the proximal one larger; not closed by maternal zooid. Measurements Holotype, means and standard deviations, mm (n == 20). Autozooid length 0.41 k0.04; width 0.35 ko.04. Orifice length ; width 0.11 ko.00. Etymology From anceps, L.-two-sided, velopment of the ovicell. alluding to the de- Echinovadoma anceps has only been found encrusting coral at Poanangisu, Efate. FAMILY F'ETRALIELLIDAE HARMER, 1957 GENUS MUCROPETRALIELLA STACH, 1936 Qpe species: Lepralia ellerii MacGillivray, MUCROPETRALIELLA CAPRICORNENSIS SP. NOV. (Fig. 16F) Material Holotype: NHM , Erakor Island reef flat, Efate, Vanuatu. Paratype: NHM , collection data as for holotype. Colony developing loosely encrusting, unilaminar sheets, deep brown in colour when dried. Autozooids rectangular to irregularly polygonal, convex, distinct, showing obvious lateral walls. Frontal shield thickly calcified, densely perforated by large, round pores.

51 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 85 Figure 16. A, Crepidacantha carsioseta. B, Crepidacantha longiseta. C-E, Echinovadoma anceps gen. et sp. nov. C. colony view. D, note the spinose ovicell endooecium and two, foramina. E, primary orifice. F, Mucropetraliella cupricornensis sp. nov. Scale bars: A = 0.25 mm; B = 0.25 mm; C =0.25 mm; D = 0.10 mm; E = 0.05 mm; F = 0.50 mm.

52 86 K. J. TILBROOK ET AL. Primary orifice wider than long, subquadrate, with a wide shallow lyrula and bluntly triangular lateral condyle-like denticulations. A suboral mucro present, with a single avicularium, laterally facing and frontally orientated; rostrum oval, mandible elongate, semielliptical. Often a short spinose process is produced on the mucro, on the opposite side to the avicularium. Two pairs of oral spines are seen in younger zooids, a pair at each distolateral corner. A low peristome may surround the distal and lateral borders of the orifice, incorporating a pair of small lateral-oral avicularia, similar in shape to that found on the mucro, located nearer to the proximal than the distal border, laterally directed. Futher avicularia may be produced distally, similar to those already described, as the peristome is established and encompasses the spines. Later in ontogeny, adventitious avicularia, similar to those described above, may be produced anywhere on the frontal shield, but more generally around the margins of the zooids, variably orientated. Ovicell globular, as wide as long, recumbent on distal zooids; endooecium very finely perforate, appearing dimpled; ectooecium not apparent. Measurements Holotype, means and standard deviations, mm (n= 15). Autozooid length 1.11 ko.09; width 0.65 k0.06. Etymology Named for the proximity of the type locality to the Tropic of Capricorn. Mucmpetraliella capricornensis is characterized by the wide, shallow lyrula, bluntly triangular lateral denticulations, and two pairs of distal oral spines. The lateral denticulations, although incorporated into the orificial rim, do not appear to function as condyles, i.e. function to articulate the operculum; in dried material of this and other Mucmpetraliella species, the operculum always sits below the lateral denticles. Gordon & d Hondt (1997) described a species from New Caledonia that they attributed to Mucmpetraliella philippinensis (Canu & Bassler, 1929), noting the presence of four oral spines, not described in Canu & Bassler s (1929) original description. Having examined cotype material of both M. philippinensis and M. falcifera (Canu & Bassler, 1929) (and found them to be the same species) it is apparent that Gordon & d Hondt (1997) were mistaken in their assignation as Canu 8z Bassler s (1929) species is devoid of spines. Mucmpetraliella loculifera Harmer, 1957 differs from M. capricornensis in having four equally spaced oral spines, sharp processes on the lateral corners of the wide shallow lyrula, and sharply triangular proximally pointing lateral denticles. Mucmpetraliella loculifera is also characterized by the presence of occlusor laminae in the primary orifice, not seen in M. capricornensis. Mucmpetraliella bennetti (Livingstone, 1926) and M. serrata (Livingstone, 1926), both from the Great Barrier Reef, differ from M. capricornensis in having six or more oral spines, a smaller lyrula with sharp processes, and sharply triangular lateral denticles. Mucmpetraliella magnifica (Busk, 1884) from Hawaii has four oral spines, but these are equidistant and the lyrula is far narrower than in M. cap rico rnensis. Only two colonies of Mucmpetraliella capricornensis were found; a small, infertile, colony fragment and a larger fertile colony. Both were found encrusting small pieces of coral rubble at Erakor Island. SINUPETRALIELLA STACH, 1936 Type species: Petralia litoralis Livingstone MS in Hastings, SINUPETRALIELLA LITORALIS (LIVINGSTONE MS IN HASTINGS) (Fig. 19A) Petralia litoralis Livingstone MS, Hastings, 1932: 438, text-fig. 15, pl. 1, fig. A. Sinupetraliella litoralis: Harmer, 1957: 706, pl. 45, figs 11-13; Ryland & Hayward, 1992: 276, fig. 25a. Petraliella litoralis: Ryland, 1974: 342. Only one very small fragment of a colony was found in debris from Vila Harbour, but the shape of the orifice is very distinctive in this species, and the present material may be confidently assigned to it. Sinupetraliella litoralis has been found on the Great Barrier Reef, Australia and from Indonesia. This record is the most easterly to date. FAMILY MICROWRELLIDAE HINCKS, 1877 GENUS MICROPORELLA HINCKS, 1877 Type species: Eschara ciliata Pallas, 1766.

53 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 87 MICROPORELLA ORIENTALIS HARMER (Fig. 19C,D) Micmporella orientalis Harmer, 1957: 962, pl. 62, figs 25-28, 38. Micmporella orientalis: Ristedt & Hilmer, 1985: 137, pl. 3, fig. 3; Ryland & Hayward, 1992: 279, fig. 25e,f. Colony encrusting, unilaminar, forming broad sheets. Autozooids with D-shaped primary orifice, with a straight, finely denticulate proximal edge. Three distal oral spines present in early ontogeny. Frontal shield nodular, densely perforated by small round pores. Ascopore with a slightly raised thickened rim and coarsely denticulate, crescentic lumen. A single adventitious avicularium on each autozooid, proximolateral to ascopore; rostrum abruptly tapered and narrowest distally; mandible setiform or variably broadened, and typically with a pair of short, hooked, lateral processes close to the base. Ovicell more or less spherical, nodular, perforated by minute pores; brooding autozooids develop a medially peaked, thickened peristomial rim distal to the ascopore, fused on each side with the ovicell. Micmpomlla orientalis was found on discarded tiles and an alga in Port Vila Harbour. Originally described from Indonesia, M. orientalis was recorded from the Great Barrier Reef by Ryland & Hayward (1992), who commented that although this species probably occurs throughout the western Pacific and perhaps the Indian Ocean, the wide geographic distribution accorded it by Harmer (1957) should not be accepted without first scrutinizing his specimens and records. FAMILY LANCEOPORIDAE HARMER, 1926 GENUS CALYPTOTHECA HARMER, 1957 Type species: Schizoporella nivea wasinensis Waters, CALYPTOTHECA RUPICOLA HAYWARD & RYLAND Calyptotheca rupicola Hayward & Ryland, 1995a: 555, figs 9F, 10A,B. Colony an encrusting, unilaminar sheet. Autozooids rectangular to irregularly polygonal, flat, separated by thick raised sutures; x mm. Primary orifice broader than long; proximal border shallowly concave between short, rounded condyles. Frontal shield densely perforated, pores interspersed with smooth, rounded tubercles, thickening with ontogeny. A curved band of especially large tubercles extends around the proximal and lateral borders of the orifice, constituting a distinct peristomial rim. Avicularia rare, proximolateral to orifice, on slightly inflated cystid; rostrum long and bluntly triangular, medially directed. Ovicell recumbent on distally succeeding autozooid, not crossed by sutures; slightly broader than long, distinctly flattened; perforation and tuberculation uniform with frontal shield. Three species of Calyptotheca were found at Heron Island, Great Barrier Reef; Calyptotheca rupicola Hayward & Ryland, 1995, C. tenuata Harmer, 1957, and C. thornelyae Dumont, 1981 (Ryland & Hayward, 1992; Hayward & Ryland, 1995a) but only C. rupicola has so far been found in Vanuatu. Four small colonies were found in the samples from Erakor and Iririki Islands on coral rubble; this is only the second record of C. rupicola. FAMILY GIGANTOPORIDAE BASSLER, 1934 GENUS COSNIOPSIS CANU & BASSLER, 1927 '&pe species: Cosciniopsis coelatus Canu & Bassler, All species have very thick, often strikingly pigmented, frontal membranes which keep their colour when dried. The colours range from orange-brown, through grey to purple, depending on the species. This genus would benefit from a thorough review as certain species assigned to it appear to differ from the concept of the type species. For example, Cosciniopsis globosa Harmer, 1957 has basal pore-chambers rather than uniporous septula. COSCINIOPSIS LONCHAEA (BUSK) (Fig. 1 7C,D) Lepralia lonchaea Busk, 1884: 146, text-fig. 43. Lepralia vestita Hincks, 1885: 256, pl. 9, fig. 9. Cosciniopsis fusca Canu & Bassler, 1927: 22, pl. 4, figs 3, 4. Gephymphora mstrigera Canu & Bassler, 1929: 278, text-fig. ll4j-0, pl. 29, figs 6-8. Cosciniopsis lonchaea: Harmer, 1957: 1083, pl. 72, figs 16, 17, 19, 20; Ryland & Hayward, 1992: 258, fig. 17b. Hippaliosina auriculata Hayward, 1988: 321, pl. 9, figs e,f.

54 88 K. J. TILBROOK ET AL. Colony encrusting, forming small unilaminar patches; light to dark brown when dried. Autozooids oval to irregularly polygonal, strongly convex, distinct, separated by deep grooves; x mm. Primary orifice oval, longer than wide, deep anter separated from shallow poster by small triangular condyles, surrounded by a peristome. Frontal shield tuberculate, perforated by numerous tiny pores. Adventitious avicularia single or paired, lateral-oral, very close to peristome, the raised rostrum directed distomedially towards orifice. Ovicell globular with tuberculate calcification and pores identical to those of the frontal shield. Autozooid vertical walls with basal series of septula. Ryland & Hayward (1992) stated that most autozooids in colonies from Heron Island had a single, lateraloral avicularium, although in some zooids this was paired, on each side of the orifice, and in others an identical avicularium occurred on the distal edge of the peristome. Many colonies from Vanuatu have very few autozooids with lateral-oral avicularia whereas some may have autozooids with one, two, or even three. In some instances the distal avicularium appears on the distal edge of the ovicell. Having examined and compared the type specimens of Cosciniopsis lonchaea and Hippaliosina auriculata Hayward, 1988, there is no doubt as to the conspecificity of the two specimens. Winston (1986) illustrated a species she described as Gephymphora rubra Osburn, 1940, from Jamaica. Her material appears, however, not to be Gephymphora Busk, 1884, as it lacks the characteristic proximal peristomial spiramen. It is extremely similar to C. lonchaea and, if not conspecific, is at least congeneric. However, Cook (1985) synonymises Gephymphora rubra Osburn, 1940 with Aptonella violacea Canu & Bassler, 1928b from Brazil, going on to discuss problems associated with these genera. Having examined the material she assigned to A. violacea it is obvious that it differs from that illustrated by Winston (1986) as G. rubra, which also lacks the peristomial spiramen. Aptonella violacea differs from C. lonchaea in a number of ways: it is purple in colour when dried, C. lonchaea is brown; the poster of the primary orifice is wider and the condyles more proximally positioned than in C. lonchaea; and the lateral oral avicularia (when present) in A. violacea are more proximal and often proximomedially directed, unlike those seen in C. lonchaea which are positioned lateral to the centre of the orifice and disto-medially directed. Despite these differences it is felt that these two species may be congeneric with Cosciniopsis, a genus known for its strikingly pigmented species, taking precedence over Aptonella. Cosciniopsis lonchaea has a known distribution in the tropical shallow seas of the Indo-West Pacific, from the Red Sea in the west to Tahiti and Hawaii in the east. Although uncommon, the occurrence of (3. lonchaea on coral rubble, from Erakor Island and Port Vila Harbour, was not unexpected. FAMILY HIPFOWDINIDAE LEVINSEN, 1909 GENUS HIPPOPODINA LEVINSEN, 1909 m e species: Lepralia feegeensis Busk, HIPPOPODINA FEEGEENSIS (BUSK) (Fig. 18A) Lepralia feegeensis Busk, 1884: 144, pl. 22, figs 9, 9a, 9b. Hippopodina feegeensis: Powell, 1967: 169, pl. 2, fig. 11; Hayward, 1988: 319; Ryland & Hayward, 1992: 256, fig. 17a; Tilbrook, 1999: 451, fig. la-f. Tilbrook (1999) recently redescribed this species and re-examined material attributed to it by previous authors. Four species of Hippopodina could be recognized from the material, including two new to science, dstinguished primarily by the shape of the primary orifice, the shape and size of avicularia, and the morphology of the ancestrular complex. Hippopodina feegeensis has a circumtropical distribution in warm shallow waters. It often forms very broad encrusting sheets on all kinds of substrata (Powell, 1969; Ryland & Hayward, 1992) and was considered by Hayward (1988) and Ryland & Hayward (1992) to be one of the most abundant bryozoans occurring on coral rubble, and to be one of the primary colonising species. However, only fragmentary specimens of H. feegeensis were found in loose debris from Iririki Island and Port Vila Harbour, despite its abundance on the Great Barrier Reef and Fiji. It appears that the relative abundances of H. feegeensis and H. iririkiensis (the former dominating, 20:1), noted by Scholz (1991) in the Philippines are reversed in Vanuatu.

55 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 89 Figure 17. A,B, Escharina pesanseris. A, colony (uncleaned). B, primary orifice, note the small rostra for the ducks foot -shaped lateral-oral avicularia. C,D, Cosciniopsis lonchaea. C, colony, growing edge. D, primary orifice. E,F, Thornelya fuscina sp. nov. E, ovicellate zooid, top centre. F, primary orifice. Scale bars: A = 0.25 mm; B = 0.05 mm; C =0.50mm; D=0.05mm; E=0.40mm; F=O.lOmm.

56 90 K. J. TILBROOK ET AL. HIPPOPODINA IRIRIKIENSIS TILBROOK (Fig. 18B) Hippopodina iririkiensis Tilbrook, 1999: 454, fig. 2a,c,e. Tilbrook (1999) distinguished Hippopodina iririkiensis from H. feegeensis (Busk, 1884) using orificial and avicularium characters and especially the occurrence, in the former, of a triad ancestrular complex compared with the tetrad complex seen in H. feegeensis. The form of the ancestrular complex is not known in the other species from Vanuatu, H. viriosa Tilbrook, Hippopodina iririkiensis is widely distributed in the tropical Indo-West Pacific, extending to the Red Sea and the Mediterranean. Originally described from Iririki Island, Vanuatu, numerous colonies of this species were collected on all types of substratum, including shell, tile, and coral rubble from Iririki Island, Erakor Island, and Port Vila Harbour. Hippopodina iririkiensis was by far the most spatially dominant species found in Vanuatu and one of the commonest. HIPPOPODINA VIRIOSA TILBROOK (Fig. 18C) Hippopodina viriosa Tilbrook, 1999: 455, fig. 2b,d,f. Tilbrook (1999) recorded Hippopodina viriosa from Vila Waterfront, and further fragmentary specimens were found in Port Vila Harbour, on tile. Although this was the least common species of Hippopodina in Vanuatu, it appears to have an almost circumglobal tropical distribution, from the Great Barrier Reef and Vanuatu to Singapore (Harmer, 1957), India, Africa (Osburn, 1940), Colombia, and the West Indies (Osburn, 1940). GENUS THORNELYA HARMER, 1957 Diagnosis Colony encrusting, unilaminar. Autozooid primary orifice bell-shaped, proximal border concave; lateral condyles distinct; oral spines present. Frontal shield with large marginal pores and small scattered frontal pores. Adventitious avicularia small, often numerous, usually associated with the orifice or ovicell. Ovicell prominent, recumbent on succeeding autozooid, irregularly rounded, surface densely perforated; closed by autozooid operculum.?stpe species: Lepralia ceylonica Thornely, THORNELYA FUSCZNA SP. NOV. (Fig. 17E,F) Material Holotype: NHM , Port Vila Harbour, Efate, Vanuatu. Paratype: NHM , Poanangisu, Efate, Vanuatu. Other material: NHM (as Thornelya sp.), Falefuti, Funafuti, Tuvalu, 146 m; NHM (as Thornely ceylonica), Sri Lanka, Thornely Coll.; NHM (as Thornelya ceylonica), New Guinea, Siboga Stn 131, 32m. Colony encrusting, unilaminar. Autozooids rectangular to irregularly polygonal, slightly convex, separated by distinct grooves. Primary orifice elongate-oval, slightly thickened around smooth edges; anter occupies 90% of orifice, flaring laterally at junction with poster, proximal border shallowly concave; condyles large, rounded-triangular; 3-4 oral spines present in younger autozooids, disappearing in later ontogeny. Frontal shield evenly perforated by round pores, immersed in thickened frontal calcification giving an almost tuberculate appearance. Adventitious avicularia small, lateral-oral, single or paired; small cystid very close to oral rim, the rostrum acute to frontal plane, directed mid-distally towards orifice; mandible acutely triangular. In some autozooids a third avicularium may be present on the proximal edge of the distal autozooid, identical to the lateral-oral avicularia, rostrum perpendicular to frontal plane, directed towards orifice of preceding autozooid. Ovicells globular, slightly wider than long, evenly perforated as in the autozooids, recumbent on, or slightly immersed in, the frontal shield of the distal zooid; closed by maternal operculum. Measurements Holotype, means and standard deviations, mm (n- 30). Autozooid length 0.56 k0.07; width 0.46 k0.07. Orifice length 0.15 ko.00; width 0.13 fo.oo. Etymology From fuscina, L.-three-pronged fork, alluding to the three perioral avicularia seen in some autozooids.

57 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 91 Figure 18. A, Hippopodina feegeensis. B, Hippopodina iririkiensis. C, Hippopodina viriosa. D, Robertsonidra novella. E, Smittoidea pacifica. F, Watersipora subovoidea. All scale bars =0.50mm.

58 92 K. J. TILBROOK ET AL. Thornelya fuscina is characterized by its orifice shape, narrower than in other species of Thornelya, and also by the three perioral avicularia, fewer than in most other Thornelya species. Some of the material attributed by Harmer (1957) to Thornelya ceylonica belongs to T fuscina (e.g. NHM ). The primary orifice in T ceylonica is far wider, and the poster much shallower, than in T fuscina and the orifice is surrounded by many avicularia. Thornelya ceylonica also has 6-8 oral spines, compared to three or four in T fuscina. While Harmer s specimens from Sri Lanka and New Guinea clearly belong to T fuscina, the specimen from Tuvalu (NHM ) differs subtly in that one of the pair of lateral-oral avicularia is longer than the other, less acute to the frontal plane, and distally directed. The oral spines also appear to remain evident longer. Only two colonies of Thornelya fuscina were found in Vanuatu, encrusting coral debris at Poanangisu and Port Vila Harbour, Efate. Thornelya fuscina occurs from Sri Lanka in the west to Tuvalu in the east, and undoubtedly occurs elsewhere within the Southwest Pacific. FAMILY INCERTAE SEDIS GENUS ROBERTSONIDRA OSBURN, 1952 Type species: Schizopodla oligopus Robertson, ROBERTSONIDRA NOVELLA RYLAND & HAYWARD (Fig. 180) Robertsonidra novella Ryland & Hayward, 1992: 261, fig. 19a. Colony encrusting, unilaminar. Autozooids oval to hexagonal, strongly convex, separated by distinct grooves; x mm. Primary orifice transversely oval, the proximal edge with a short U-shaped median sinus flanked by small condyles; one or two short spines present distally, their bases persisting, particularly in ovicellate zooids; a short umbo present, lateral to the sinus. Frontal shield regularly tuberculate, with a single series of round marginal pores. Adventitious avicularia present on most autozooids, laterally suboral; rostrum elongate-triangular, directed proximolaterally. A second, shorter avicularium may occur lateral to the sinus, acute to the frontal plane, directed laterally. Ovicell wider than long, globular, tuberculate, with minute perforations be-. tween tubercles. Originally described from Heron Island, Australia, single colonies of Robertsonidra novella were collected at Iririki Island and Port Vila Harbour on small pieces of coral rubble. SUPERFAMILY NIAMILUIFOROIDEA CANU & BASSLER, 1927 FAMILY CREPIDACANTHIDAE LEVINSEN, 1909 GENUS CREPIDACANTHA LEVINSEN, 1909 Type species: Flustra poissonii Audouin, Flustra poissonii Audouin, 1826 has no type material, a considerable taxonomic problem, especially as it has been designated type species of Crepidacantha Levinsen, This could be rectified by the selection of a neotype specimen from the Egyptian Red Sea, which would not only stabilize the species but also the genus. CREPIDACANTHA CARSIOSETA WINSTON & HEIMBERG (Fig. 16A) Crepidacantha carsioseta Winston & Heimberg, 1986: 27, figs Crepidacantha carsioseta: Ryland & Hayward, 1992: 277, fig. 25d. Crepidacantha carsioseta differs from C. longiseta Canu & Bassler, 1928a in its avicularia which are frontolaterally placed and transversely orientated, thus pointing laterally, whereas those of C. longiseta are lateral-oral, very long, and proximally orientated. The shapes of the primary orifices are similar although the proximal border of C. carsioseta is far more convex than in C. longiseta. This distinctive species was only recently described from Indonesia by Winston & Heimberg (1986), and subsequently reported from Heron Island by Ryland & Hayward (1992). Crepiducantha carsioseta was mainly found on small pieces of coral debris from Poanangisu, Efate. CREPIDACANTHA LONGISETA CANU & BASSLER (Fig. 16B) Crepidacantha longiseta Canu & Bassler, 1928a: 135, pl. 21, figs 3, 4. Crepidacantha longiseta: Hayward, 1988: 313.

59 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 93 Figure 19. A, Sinupetraliella litoralis. B, Reteporella graeffei. C,D, Micruporella orientalis. C, ovicellate zooids. D, detail of primary orifice, ascopore and avicularium. E,F, Monoporella nodulifera. E, note ovicellate zooid, centre. F, primary orifice and prominent opesiules. Scale bars: A = 0.40 mm; B = 0.25 mm; C = 0.40 mm; D = 0.05 mm; E = 0.40 mm; F =0.10 mm.

60 94 K. J. TILBROOK ETAL. Crepidacantha longiseta is readily distinguishable from C. carsioseta by the positioning of its avicularia. However C. crinispina Levinsen, 1909, also from the western Pacific (see Ryland & Hayward, 1992) has similarly placed lateral-oral avicularia, but these are shorter, and the primary orifices also differ between the two species; the primary orifice in C. longiseta has a far flatter proximal border than that seen in C. crinispina. Further differences are seen in the ovicell: the horse-shoe-shaped uncalcified area of ectooecium being more frontally positioned and smaller in C. longiseta than in C. crinispina. Crepidacantha longiseta was the commoner of the two species from Vanuatu, being found on both small and large pieces of coral rubble, particularly in Port Vila Harbour, Efate. It has also been reported from Brazil, the Caribbean, Mauritius, and the western Indian Ocean (Brown, 1954) and appears to have a circumglobal distribution. SUPERFAMILY CELLEWROIDEA JOHNSTON, 1838 FAMILY CELLEPORIDAE JOHNSTON, 1838 GENUS TURBICELLEPORA RYLAND, 1963 Type species: Cellepora comnopus Wood, TURBICELLEPORA AMPLA (KIRKPATRICK) Schizoporella ampla Kirkpatrick, 1888: 76, pl. 7, fig. 4. Stephanosella bernardii Harmer, 1957: 1051, pl. 74, figs !ILrbicellepora ampla: Hayward, 1988: 341, figs 6d,e; Ryland & Hayward, 1992: 284, fig. 27c,d. A single, small, infertile colony of this species was found encrusting a small piece of coral rubble at Erakor Island. Originally described from Mauritius in the Indian Ocean, it has subsequently been noted in the Indo-Malayan region and the Great Barrier Reef. The present record extends the known range eastwards into the Coral Sea. FAMILY PHIDOLOWRIDAE GABB & HORN, 1862 FODINELLA GEN. NOV. Diagnosis Colony encrusting to calyciform. Autozooids subrectangular to variable in shape, the frontal shield imperforate; two to six areolar pores along margin. Autozooids most distinct in early ontogeny, the boundaries becoming increasingly indistinct through secondary calcification and frontal budding. Orifice longer than wide; relatively broad, rounded anter with a denticulate rim, separated from the narrower, shallower, smooth poster by well developed lateral condyles. Suboral umbo may be present. Distal oral spines present; spine bases seen at the entrance to the ovicells. Adventitious avicularia present, may be lacking in some zooids, or developing in later ontogeny. Ovicells recumbent, smooth, imperforate, with a wide rounded, concave, proximal aperture, arching above the primary orifice; not closed by autozooid operculum; subimmersed in later ontogeny. Type species: Eschamides spinigera Philipps, Etymology From fodina, L.-mine; -ellus, L. diminutive, small, alluding to the pitted appearance of colonies. When erecting their new genus, Lifuella, Gordon & dhondt (1997) suggested that all tropical species previously assigned to Hippoporella Canu, 1917 which possessed a beaded or denticulate distal rim should probably be reassigned to Stephanollona Duvergier, 1921 (the senior subjective synonym of Bmdiella Uttley & Bullivant, 1972). However, the type species Stpphanollona spinifera Duvergier, 1921 (and all described species of Bmdiella) possess a labellum across the ovicellular opening, e.g. Hippoporella labiata Hayward & Cook, Fodinella gen. nov. is accordingly introduced for three species having in common a noncleithridiate primary orifice, a beaded distal orificial rim and a widely open ovicellular aperture. Both Lepralia calyciformis Philipps, 1900 (assigned to Lifuella by Gordon & dhondt, 1997) and Lepralia tuberculata, Philipps, 1899 are here referred to this genus, following examination of their holotype specimens. It is noteworthy that all of Philipps' (1900) species were originally described from the Loyalty Islands, southwest of Vanuatu and thus all known species of Fodinella gen. nov. are from a limited area of the southwest Pacific. FODINELLA SPINIGERA (PHILIPPS) COMB. NOV. (Fig. 20A,B) Eschamides spinigera Philipps, 1900: 400, 448, pl. 43, fig. 12. Hippoporella spinigera: Harmer, 1957 (in part): Not Hippoporella spinigera: Hayward & Cook, , figs 19C,D.

61 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 95 Colony encrusting often calyciform. Autozooids subrectangular to variable in shape, frontal shield thick, imperforate, porcellanous, smooth except for up to 6 marginal pores; x mm. Autozooidal boundaries become indistinct in later ontogeny. Primary orifice longer than wide, widest distally; anter rounded, with denticulate rim, separated from shallowly concave poster by prominent, proximally directed, lateral condyles. Seven to eight distal oral spines present on all autozooids. A stout suboral umbo developing in later ontogeny, partly overhanging the primary orifice, sinuses on either side uneven in depth. Avicularia large, parallel-sided, slightly spatulate; sporadic, orientation variable, often becoming obscured in later ontogeny; crossbar complete with a very small columella; a small triangular opesia, pointing distally at proximal end of avicularium; rostrum raised into a thin wall distally, denticulate at flattened distal end. Ovicell subimmersed in later ontogeny, imperforate with a wide, rounded proximal opening; not closed by the zooidal operculum; spine bases visible within the ovicellular opening. Rounded, elongate oval porechambers high on lateral and distal walls. Some older, larger colonies appear pinkish-brown when dried, younger colonies more translucent white. Fodinella spinigera is similar to the species described and illustrated by Gordon (1984) as Lepraliella multidentata (Thornely, 1905) from New Zealand, which however, was not Thornely s species. Fodinella spinigera differs from that species in having seven or eight oral spines rather than six, and a greater number of areolar septular pores of which the lateral and distal ones are less oval than those proximally; the orifice differs in shape too. Gordon s (1984) species is certainly congeneric with R spinigera and rather similar to Schizotheca mucmnata Powell, 1967, also described from New Zealand, another species here tentatively referred to Fodinella. Hayward & Cook (1983) mistakenly identified Lifuella porelliformis (Kirkpatrick, 1888) as l? spinigera (see below) in their material from the eastern coast of Africa, due to Harmer s (1957) confused identifications. Autozooids of R spinigera may be substantially deeper than longer, depending on the contour of the substratum. Colonies can become reproductive when consisting of less than twenty autozooids. Philipps (1900) described a minute avicularium in Eschamides spinigera on the inner side of the peristome, not seen in the specimens examined. Fodinella spinigera was found on small pieces of coral rubble from Port Vila Harbour, Efate. It was originally described from the nearby Loyalty Islands. Dumont s (1981) record of this species from the Red Sea should be investigated further. GENUS LIFUELLA GORDON & D HONDT, 1997 Diagnosis Colony encrusting to calyciform. Autozooids porcellanous, the frontal shield imperforate, with or without small areolar septular pores. Primary orifice with a broadly arcuate, smooth-rimmed anter, separated from the poster by proximally directed condyles. Articulated oral spines present in both infertile and fertile autozooids. Adventitious avicularia, when present, frequently lateral-oral; with complete crossbar. Ovicell subimmersed, the ectooecium imperforate, usually covered by a smooth layer of secondary calcification; the ovicellular entrance usually widely open, not closed by the autozooidal operculum. Type species: Lepralia multidentata Thornely, Gordon & dhondt (1997) introduced this genus to include the tropical to warm-temperate species previously attributed to Hippoporella Canu, 1917 (named for Lepralia hippopus Smitt, 1873), which have a primary orifice without a denticulate distal rim. This followed concerns expressed by Hayward & Cook (1983) over the congeneric assignment of tropical species with the boreal type species. Species which should be included in this genus are Lepralia multidentata Thornely, 1905, Mucmnella porelliformis Kirkpatrick, 1888, Mucmnella articulata Philipps, 1900 and possibly Lepraliella?mooraboolensis (MacGillivray, 1895) sensu Gordon (1984). Under their original description of this genus, Gordon & dhondt (1997) assigned Lepralia calyciformis Philipps, 1900 to Lifuella, but examination of the type material shows it to be a species of Fodinella gen. nov. (see above) as it possesses a denticulate anter (readily seen in Gordon & d Hondt, 1997, fig. 145). LIFUELLA MULTIDENTATA (THORNELY) (Fig. 200) Lepralia multidentata Thornely, 1905: 120, pl. (unnumbered), fig. 9. Lepralia purpurea Thornely, 1905: 120, pl. (unnumbered), fig. 13. Lepralia multidentata: Hastings, 1966: 64. Hippoporella multidentata: Harmer, 1957: 1099, pl. 73, figs 9-12; Dumont, 1981: 636. Not Lepraliella multidentata: Gordon, 1984: 123, pls 50E, 51A.

62 96 K. J. TILBROOK ET AL. Figure 20. A,B, Fodinella spinigera comb. nov. A, autozooids and ovicellate zooids. B, note the denticulate anter in primary orifice. C, Lifuellaporelliformis. D, Lifuella multidentata. Scale bars: A=0.50mm; B=0.05 mm; C =0.20mm; D=O.ZOmrn. Colony encrusting, sometimes calycifonn. Autozooids appearing porcellanous when dried, the frontal shield imperforate, slightly tuberculate in younger autozooids, with 5-7 small areolar pores; separated by shallow grooves in early ontogeny, later less distinct owing to secondary calcification. Primary orifice bellshaped, slightly longer than wide, with a broadly arcuate smooth-rimmed anter, separated from the shallow poster by small, proximally directed condyles. Generally 4-5 articulated oral spines present in both infertile and fertile autozooids. Suboral umbo present, often becoming very tall and pointed in later ontogeny and even bifurcating. Adventitious avicularia present; small, triangular, single or paired, lateral-oral and frequently distal-oral; with complete crossbar. Dried material ranges from translucent white to pinkish purple. Autozooids of Lifiella multidentata change greatly in appearance through ontogeny, so much so that were the growing edges not present in some specimens one might easily assign younger and older colonies to different species. The young colonies found at Vanuatu were small and encrusting, the autozooids distinct and devoid of avicularia. However, lectotype and paralectotype material (designated by Hastings, 1966) has autozooids that are far less individually distinct, with an abundance of avicularia and very tall umbones which in some cases (NHM ) are bifurcate. The lectotype specimen (NHM , L.R. Thornely Coll.) is a large colony with a growing edge, which allows comparison with the older parts of the colony. The specimen illustrated by Gordon (1984) had, by his own description, a denticulate anter and is there-

63 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 97 fore not a species of Lifuella, rather a species of Fodinella gen. nov. Two young colonies of this species were found from Iririki Island. This record extends the range of the species from the Red Sea and Sri Lanka through the Philippines to the south west Pacific Ocean. LIFUELLA PORELLIFORMIS (KIRKPATRICK) (Fig. 20C) Mucmnella porelliformis Kirkpatrick, 1888: 81, pl. 9, fig. 1. Mucmnella serratilabris O Donoghue, 1924: 48, pl. 3, fig. 18. Hippoporella spinigera: Harmer, 1957 (in part): 1100; Hayward & Cook, 1983: 79, fig. 19C,D. Hippoporellaporelliformis: Hayward, 1988: 322, pl. 10, figs a,b. Not Hippoporella porelliformis: Scholz, 1991: 324, pl. 19, fig. 4, pl. 7, fig, 2. Colony encrusting, forming small, raised, circular, silvery patches. Autozooids small, oval to hexagonal, rather broad; separated by shallow grooves, obscured by calcification later in ontogeny, c mm long. Primary orifice wider than long, almost D-shaped, blunt lateral condyles present. Eight long, slender, closely grouped distal oral spines. Peristome developed proximally as a prominent umbo with a substantial mucro, broad, quadrate, a finely serrated lip on its distal edge. Frontal shield covered by smooth tuberculations, imperforate except for 3-5 relatively inconspicuous marginal pores. Paired adventitious avicularia, lateral to orifice; mandible acutely triangular, directed laterally or slightly disto-laterally. Additional avicularia sometimes present on frontal shield; most frequently elongate, situated on the midline of the autozooid, the slender triangular mandible directed proximally or proximolaterally; more rarely one or two avicularia, similar to oral type but larger, situated elsewhere on the frontal shield, orientation varying. Basal pore-chambers present. Ovicell prominent, hyperstomial, recumbent on succeeding autozooid, smooth, spherical, with a large semicircular orifice, often developing a small umbo. Rein arks Lifuella porelliformis is a very distinctive species, with its D-shaped primary orifice; eight long, slender, spines, prominent umbo, substantial quadrate mucro with a finely serrated lip on its distal edge, and paired lateral-oral triangular avicularia. Harmer (1957) seems to have confused subsequent authors by describing Eschamides spinigera Philipps, 1899 while illustrating a specimen which is undoubtedly Kirkpatricks (1888) Mucmnella porelliformis. The two species are similar, but the suboral umbo is far more pronounced in L. porelliformis and has denticulations on the mucro, not seen in Philipps species, which produces a more pronounced proximal sinus and has a denticulate anter (see generic description of Fodinella gen. nov., above). The position and shape of the avicularia are also characteristic for L. porelliformis. Lifuella porelliformis does closely resemble Mucmnella articulata Philipps 1899 (here assigned to Lifuella, see above) both in overall colony form and in autozooid morphology. However, the latter has a more elongate orifice, only six, long, slender spines (which flare out at their ends), paired lateral avicularia, which are shoe -shaped, and no denticulations on the distal edge of its suboral mucro. Neither of the Lifuella porelliformis colonies found from Vanuatu showed the additional frontal-shield avicularia seen in the type specimen, described by Hayward & Cook (1983) and illustrated by Hayward (1988). Lifuella porelliformis was found encrusting small pieces of coral rubble in Port Vila Harbour, Efate. Its wider distribution extends to Mauritius and South Africa. GENUS METACLEIDOCHASMA SOULE, SOULE & CHANEY, 1991 &pe species: Metacleidochasma dimorphum Soule, Soule & Chaney, Some specimens of the type species may include zooids with large dimorphic orifices. The ovicellular labellum is formed by the central suturing of two projections from the lateral margins of the ovicellular hood. These extend across the ovicell aperture, leaving a narrow slit-like opening above (no lateral incisions are produced as seen in Schedochleidochasma Soule, Soule & Chaney, 1991 and Plesiochleidochasma Soule, Soule & Chaney, 1991); the ovicell thus becoming immersed. METACLEIDOCHASMA PLANULATA (CANU & BASSLER) (Fig. 21 C-E) Hippoporina planulata Canu & Bassler, 1929: 321. Metacleidochasma ovule Soule, Soule & Chaney, 1991: 480, pl. 1, fig. 7, pl. 4, figs 1-2.

64 98 K. J. TILBROOK ET AL. Metacleidochasmaplanulata: Gordon & d'hondt, 1997: 59. Autozooids in early ontogeny generally bear 4 (but can vary from 3-5) oral spines which are soon obscured by succeeding autozooids. Only 2 frontal pores appear, situated near the margin on either side of the autozooid at its widest point. Soule et al. (1991) describe the rare presence of a large, orifice-sized, broadly oval, adventitious avicularium, proximal to the orifice. Ovicell globose, imperforate, with a slit-like opening and completely fused labellum; ovicells become immersed, indistinct, only the slit above the labellum visible. Tatiform ancestrula with 9 spines around a large, distally positioned, rounded, membranous frontal area; 3 spines evenly spaced around the proximal half of the frontal membrane, the remainder around the distal half. None of the specimens of Metacleidochasma planulata found in Vanuatu had the frontal avicularia described by Soule et al. (1991), and noted by Gordon & d'hondt (1997) in the type specimen, but not noted by Canu & Bassler (1929). Originally described from the Philippines by Canu & Bassler (1929), the species was redescribed from Hawaii as Metacleidochasma ovule by Soule et ul. (1991), who noted its further distribution in the Maldives, Thailand, Tonga, and French Polynesia. Its occurrence in Port Vila Harbour and at Iririki Island, is thus not surprising. GENUS PLESIOCHLEIDOCHASMA SOULE, SOULE & CHANEY, 1991 m e species: Lepralia porcellana var. normani Livingstone, PLESZOCHLEIDOCHASMA NORMANI (LIVINGSWNE) Lepralia porcellana var. normani Livingstone, 1926: 92, pl. 8, fig. 1. Plesiocleidochasma normani: Soule, Soule & Chaney, 1991: 474, pl. 3, fig. 3, pl. 4, fig. 4; Hayward & Ryland, 1995a: 565. Schedocleidochasma porcellanum: Ryland & Hayward, 1992: 287, fig. 27e. Plesiocleidochasma normani appears to be widely distributed in the western Pacific. Soule et al. (1991) recorded it from many of the Southwestern Pacific island groups including Vanuatu, as well as Indonesia, Tonga, French Polynesia, and Hawaii. Ryland & Hayward (1992) reported it (as Schedocleidochasma porcellanum, see Hayward & Ryland, 1995a) to be very common at over 20 of the sites they sampled at Heron Island, Great Barrier Reef. It was not found in the material examined for the present study, however. GENUS RETEPORELLA BUSK, 1884 Qpe species: Reteporella jlabellata Busk, RETEPORELLA GRAEFFEI (KIRKENPAUER) (Fig. 19B) Retihornera graeffei Kirkenpauer, 1869: 30. Reteporella graeffei: Harmer, 1934: 573, pl. 35, figs 12-15, pl. 38, figs 13-15; text-figs 25E, 31, 32; Ryland, 1984: 74, figs 38.11, ; Ryland & Hayward, 1992: 285, figs 28c,d, 29a. Reteporella graeffei is a common member of shallow reef faunas throughout the tropical western Pacific from Queensland and the Great Barrier Reef to Japan, ranging westwards through Indo-Malaysia to Sri Lanka and East Africa. It should be noted that the type locality is Fiji, and thus the described geographical range may include some spurious records, referring to species other than Reteporella graeffei sensu stricto. In Vanuatu a small fragmentary sample was found at Poanangisu in loose coral debris. GENUS SCHEDOCLEIDOCHASMA SOULE, SOULE & CHANEY, 1991 Type species: Schedocleidochasma pornellaniforme Soule, Soule & Chaney, SCHEDOCLEIDOCHASMA PORCELLANIFORME SOULE, SOULE & CHANEY (fig. 21A,B) Schedocleidochasma porcellaniforme Soule, Soule & Chaney, 1991: 481, pl. 1, fig. 6, pl. 5, figs 3, 4. Autozooids bear three oral spines in early ontogeny which are later obscured by succeeding autozooids. Primary orifice with a large, almost circular, smooth anter and a small, deep, arrow-shaped poster. Avicularia, small, single or paired, directed distolaterally; rostrum acute triangular, with slightly raised acuminate tip; crossbar complete, with short columella.

65 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 99 Figure 21. A,B, Schedocleidochasma porcellaniforme. A, note the ovicell labellum, centre left. B, primary orifice, note the smooth anter and arrowhead-shaped poster. C-E, Metachleidochasma planulata. D, primary orifice, note the denticulate anter and rounded poster. E, Ancestrula (top left) and two peri-ancestrular autozooids. F, Catenicella triangulifera. Scale bars: A=0.20mm; B=0.05mm; C=0.25mm; D=O.O5mm; E=O.lOmm; F=0.25mm.

66 100 K. J. TILBROOK ET AL. Schedocleidochasma porcellaniforme resembles Lepraliaporcellana Busk, 1860 which Soule et al. (1991) included in this genus. In their opinion, S. porcellaniforme has larger avicularia and ovicells than Busks species and a slightly wider and shallower poster to the primary orifice. In their original description of Schedocleidochasma porcellaniforme from American Samoa, Soule et al. (1991) noted its presence from the Maldive Islands, Thailand, Indonesia, Solomon Islands, Fiji, Tonga, French Polynesia, Hawaiian Islands, and Tizard Reef (China Sea). They also found this species in Vanuatu, on reefs around Mele Bay, Efate. Our study has found colonies encrusting coral rubble from Poanangisu and Iririki Island, Efate. GENUS RHYNCHOZOON HINCKS, 1895 Type species: Lepralia bispinosa Johnston, Rhynchozoon species are common in all tropical reef habitats. However, they are notoriously difficult to identify; the most useful taxonomic characters being the size and shape of the primary orifice, particularly the proximal sinus, and the morphology of the proximal suboral avicularium. Certain characters of the frontal avicularium and ovicell may also be useful. Many new species have been described in recent years (e.g. Ryland & Hayward, 1992). RHYNCHOZOON FEROCULA HAYWARD (Fig. 22A,B) Rhynchozoon ferwcula Hayward 1988: 332, pl. 13, figs a-c. Very small colonies only of this species were found in samples from Iririki Island, Port Vila Harbour and Poanangisu, but none had developed the domed form seen in the Mauritian type material. However, the distinctive primary orifice, with its border of four spines, is characteristic: the anter is circular, c. 0.1 mm diameter, with blunt, widely spaced denticulations and small, rounded condyles proximolaterally; the short, rounded sinus is defined by a pair of medially curved processes arising frontal to the condyles. The peristome is most pronounced proximally and variably produced into short processes. There is no uncinate process and no suboral avicularium; small adventitious avicularia, with triangular rostrum, thin crossbar and lacking a palate, develop on the proximal frontal shields of zooids and tend to be directed towards adjacent peristomes. The ovicell is broader than long, with a very short, broad labellum; it has a semielliptical frontal tabula, exposing smooth entooecium, and develops a coarsely granular ooecial cover. Colonies from Iririki Island consisted of ten or fewer autozooids, with the ancestrula still visible. The ancestrula is oval with a transversely elliptical opesia occupying half its frontal surface; the opesial rim has few, very widely spaced, knob-like denticulations, and there are eight evenly spaced spines. Originally described from Mauritius (Hayward, 19881, these records from Vanuatu represent only the second occurrence of the species. RHYNCHOZOON SPLENDENS HAYWARD (Fig. 22C-E) Rhynchozoon splendens Hayward, 1988: 335, pl. 13, figs f,g. Rhynchozoon splendens: Ryland & Hayward, 1992: 294, figs 31f, 32a,b. Rhynchozoon rwstratum: Winston & Heimberg, 1986: 38, figs 95-98; Scholz, 1991: 317, pl. 18, figs 1, 2, 5. Colony unilaminar to multilaminar; autozooids oval to hexagonal, commonly x mm. Frontal shield convex, with large and conspicuous marginal pores; vitreous, initially smooth but developing large, rounded nodules early in ontogeny. Primary orifice about as wide as long, 0.12 mm diameter; distal denticulations small and widely spaced, sinus short, U- shaped, with prominent thickenings laterally, below indistinct condyles. No oral spines. Peristome completely encircling orifice, partially obscuring it; rim produced into short, knobbed processes, proximally with an eccentrically placed U-shaped notch, adjacent to a short, tapered uncinate process projecting into the peristome lumen. A prominent lateral-oral avicularium incorporated into the peristome in some autozooids; its proximal portion continuous with the uncinate process, rostrum triangular, hooked distally, directed distolaterally. Frontal adventitious avicularia present in later ontogeny, 1, rarely 2 per autozooid, characteristic: proximolateral to peristome, directed distolaterally, with elongate, diamond-shaped outline. Ovicell slightly broader than long, with extensive frontal tabula; aperture straight edged, 2 short lateral notches defining a short, broad labellum; developing a nodular ooecial cover.

67 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 101 Figure 22. A,B, Rhynchozoon splendens. A, ovicellate zooid, top right. B, primary orifice, note the wide, shallow proximal sinus. C-E, Rhynchozoon femcula. C, ovicellate zooids. D, primary orifice, note the small, rounded proximal sinus. E, aperture of ancestrula. Scale bars: A = 0.40 mm; B = 0.05 mm; C = 0.25 mm; D = 0.05 mm; E = 0.05 mm.

68 102 K. J. TILBROOK ET AL. Rhynchozoon splendens is readily recognized by its orbicular orifice, slightly wider than long, a short U- shaped sinus, grossly nodular frontal shield and diamond-shaped frontal avicularia; the latter were not seen in the Vanuatu material. Rhynchozoon splendens appears widespread throughout the Indo-West Pacific. It was originally described from Mauritian reef flats and was found to be common in similar habitats at Heron Island, Great Barrier Reef (Ryland & Hayward, 1992). The species has been recorded from Indonesia, the Philippines and Australia, with the records from Iririki and Erakor Islands, Efate the most easterly to date. CONCLUSION This first account of the cheilostomatous Bryozoa of Vanuatu includes 92 species, of which 20 are new species and a further one, Beania cf. mirabilis, cannot be confidently assigned to any established taxon. This proportion of previously undescribed species is not unusual for faunistic studies of Southwest Pacific bryozoans (e.g. Ryland & Hayward, 1992; Hayward & Ryland, 1995a; Gordon & dhondt, 1991, 1997), and it is certain that the total number of species recorded does not accurately represent the true taxonomic diversity of the Cheilostomatida of Vanuatu. The collections forming the basis of this study were made during two periods, at just four shallow water locations on the island of Efate; the majority of the islands comprising Vanuatu remain unsampled. Collecting was limited by access to the shore, and was conducted by wading at low tide and by snorkelling. The range of habitats sampled was consequently narrow, and the substrata were restricted to coral rubble, occasional shell fragments, and harbour trash. In the light of these limitations, therefore, the number of new taxa recorded is perhaps, after all, unusual but simply reflects the minimal attention sessile invertebrates, other than corals, have attracted from taxonomic specialists in the Southwest Pacific generally. The habitats sampled in this survey were all in proximity to Port Vila and other tourist centres, and must be described as highly perturbed. Samples collected for bryozoan specimens were equally encrusted by sponges, foraminiferans and algae, and showed evidence of continuous grazing by fish, and perhaps molluscs. However, natural perturbations resulting from earthquakes, volcanic eruptions, and consequent sea level changes are a long established feature of the reef habitats of Vanuatu (Done & Navin, 1990). The scale of the present survey is inadequate to enable speculation regarding any relationship between the frequency of natural environmental disturbance and the taxonomic and ecological characteristics of sessile epifaunas. Of the 20 new taxa described here, 16 are presently known only from Vanuatu, but it is unlikely that they will prove to be endemic to the unstable reef habitats of these islands and further surveys elsewhere in the eastern Coral Sea and on the Great Barrier Reef will probably show all to have wider geographic distributions. Similarly, it is unlikely that few if any of the species described here will be limited to the shallow, perturbed environments in which they were found, but instead will have been collected at the limits of their ecological ranges. Several exceptions, however, should be noted. Poricella spathulata was especially abundant in these collections, represented by 57 specimens, and from all four localities. It has been collected, again abundantly, in harsh upper shore habitats at both Mauritius and Heron Island (Hayward, 1988; Hayward & Ryland, 1995a) and is perhaps adapted through extreme ecological tolerances to such habitats. Antmpora granulifera, Crepidacantha longiseta, Escharina pesanseris and the recently recognized Hippopodina iririkiensis were also common at three, at least, of the localities sampled, and the first three species seem to have a geographical and ecological distributions similar to that of Poricella spathulata. It is not possible to give an initial ecological and biogeographical evaluation of the shallow bryozoan fauna of Vanuatu, given such a proportion of newly recognized taxa, and the almost total lack of information on the biology and ecology of all 92 species, but it is probable that it consists predominantly of eurytopic species. Nevertheless, it should be noted that 45 species occurred only at one of the four localities sampled and only 20 were found at three of four localities; 29 of the species were represented by ten or more specimens while 31 were represented by three or fewer. It is worth noting that a number of species are recorded here for only the second time, Puellina egretta, Nimba saxatilis, Calyptotheca rupicola, Beania cookae sp. nov., Beania klugei, Bugula scaphula sp. nuv., Drepanophora indica, Rhynchozoon femcula, with the geographic ranges of the last four species being greatly extended eastwards from the Indian Ocean. The taxonomic diversity of the cheilostomatous Bryozoa of Vanuatu will probably prove to be at least twice the figure noted here, but further sampling, at other localities and in other habitats, is needed before it can be reasonably estimated. Comparison with other tropical bryozoan faunas is presently premature. The fauna is predominantly Indo-West Pacific in character with a high proportion of species, 27, apparently known only in the Southwest Pacific, and it has in common with other Southwest Pacific faunas so far studied, a

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74 108 K. J. TILBROOK ET AL. genus pseudoplanktonic on molluscs and algae. Znvertebrate Biology Thornely LFL Report on the Polyzoa collected by Professor Herdman. at Ceylon. in Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar 4 (Suppl. Report 26): Tilbrook KJ The species of Antmpom Norman (Bryozoa: Cheilostomatida). with the description of a new genus in the Calloporoidea. Records of the South Australian Museum 31: Tilbrook KJ of Hippopodina feegeensis and three other species of Hippopodina Levinsen (Bryozoa: Cheilostomatida). Jouml of Zoology. London 217: Tilbrook KJ First record of the bryozoan genus Stylopoma from the Mediterranean Sea. Journal of the Marine Biological Association. UK 80: Tilbrook KJ Indo-West Pacific species of the genus Stylopoma Levinsen (Bryozoa: Cheilostomatida). Zoological Journal of the Linnean Society 131: Tilbrook KJ. (in press). New bryozoan family from the Indo-Pacific shows unexpected diversity. Journal of the Marine Biological Association. UK. Uttley GH. Bullivant JS Biological results of the Chatham Islands 1954 Expedition. Part 7. Bryozoa Cheilostomata. Memoirs. New Zealand oceanographic Institute Vigneaux M %vision des Bryozoaires neogenes du Bassin daquitaine et essai de classification. Memoires de la Societe geologique de Fmnce 28: Waters AW The use of opercula in the determination of the cheilostomatous Bryozoa. Proceedings of the Manchester litemry and philosophical Society 18: Waters AW Bryozoa from New South Wales. Part IV. Annals and Magazine of Natural History (6) Waters AW Reports on the Marine Biology of the Sudanese Red Sea. from collections made by Cyril Crossland. M.A., BSc., F.Z.S.; together with collections made in the Red Sea by Dr. R. Hartmeyer - XI1. The Bryozoa. Part 1 - Cheilostomata. Journal of the Linnean Society, Zoology 31: Waters AW The marine fauna of British East Africa and Zanzibar. from collections made by Cyril Crossland. M.A., B.Sc., F.Z.S., in the years Bryozoa- Cheilostomata. Proceedings of the Zoological Society of London Winston JE Marine bryozoans (Ectoprocta) of the Indian River area (Florida). Bulletin of the American Museum of Natural History 173(2): Winston JE Shallow-water bryozoans of Carrie Bow Cay. Belize. American Museum Novitates 2799: WinstonJE An annotated checklist of coral-associated bryozoans. American Museum Novitates Winston JE. Heimberg BF Bryozoans from Bali. Lombok and Komodo. American Museum Novitates 2847: 149. Wood SV Descriptive catalogue of the zoophytes of the Crag. Annals and Magazine of Natural History APPENDIX INDEX OF SPECIES (NEW SPECIES IN BOLD) Aetea?australis fide Gordon... Aetea ligulata Busk... Jellyella tuberculata (Bosc) Biflustra reticulata... Conopeum papillorum... Antmpora gmnulifera (Hincks)... Antmpora minor (Hincks)... Pamntmpom penelope Tilbrook... Corbulella corbula (Hincks)... Crasaimarginatella corniculata... Crasaimarginatella maxillaria... Cranosina comnata (Hincks)... Nellia tenuis Harmer... Bugula acaphula... Brettiella culmoaa... Beania klugei Cook... Beania petiolata Harmer... Beania coohe... Beania hexumicorum Beania cf mirabilis Harmer... Scrupocellaria spatulata (dorbigny)... Synnotum aegyptiacum (Audouin)... Smittipora cordiforrnis Harmer... Monoporella nodulifem Hincks... Steginoporella jellyae nom. nov... Thalamoporella granulata Levinsen... Thalamoporella gracilata... Labioporella spatulata Harmer... Cellaria punctata (Busk)... Puellina egretta Ryland & Hayward... Puellina vulgaris Ryland & Hayward... Figularia pulcherrima... Catenicella triangulifera Harmer... Vasignyella otophom (Kirkpatrick)... Savignyella lafontii (Audouin)... Hippothoa calciophilia Gordon... Hippothoa flagellum Manzoni... Pleaiothoa bucarina... Trypoatega maculata... Chorizopora bmngniartii (Audouin)... Poricella mbusta (Hincks)... Poricella spathulata (Canu & Bassler)... Exechonella brasiliensis Canu & Bassler... Reptadeonella fissa (Hincks)... Reptadeonella cellulanue... Reptadeonella noviaaima... Eschamides longimstris Dumont... Celleporaria pilaefera (Canu & Bassler)... Celleporaria vagans (Busk) Celleporaria inaudita... Drepanophora indica Hayward... Drepanophora tuberculata Osburn... Drepamphra gutta... Metmperiella montferrandii (Audouin)... Watersipora subouoidea sensu Harmer... Parasmittina galerita Ryland & Hayward... Parasmittina hustingsae Soule & Soule... Parasmittina serrula Soule & Soule... Pleurocodonellina signata (Waters)... Smittoidea pacifica Soule & Soule... Tbrquatella duolamellata (Scholz) comb. nov $

75 CHEILOSTOMATOUS BRYOZOA FROM VANUATU 109 Schizoporella dunkeri (Reuss)... Schizoporella ermta (Waters)... Escharina pesanseris (Smitt)... Stylopoma novum Tilbrook Stylopoma velatum Tilbrook... Stylopoma vilaensis Tilbrook... Stylopoma viride (Thornely)... Nimba saxutilis Hayward & Ryland... Echinouadoma anceps gen. et sp. nov... Mucropetraliella capricornensis... Sinupetmliella litoralis Hastings... Micmporella orientalis Harmer... Calyptotheca rupicola Hayward and Ryland... Cosciniopsis lonchuea (Busk)... Hippopodina feegeensis (Busk)... Hippopodina iririkiemis Tilbrook Hippopodina viriosa Tilbrook Thornelp fuacina Robertsonidra novella Ryland & Hayward Crepidacantha carsioseta Winston & Heimberg Crepidacantha longiseta Canu & Bassler Thrbicellepora ampla (Kirkpatrick) Fodinella spinigera (Philipps) comb. nov Lifuella multidentata (Thornely) Lifuella porelliformis (Kirkpatrick) Metacleidochasma planuluta (Canu & Bassler) Plesiochleidochasrna normani (Livingstone) Retepowlla graeffei (Kirkenpauer) Schedocleidochasma pornellaniforme Soule. Soule & Chaney Rhynchozoon ferocula Hayward Rhynchozoon splendens Hayward