CHEILOSTOMATOUS FROM VANUATU 79

& Sandberg (see Gordon, 1984) which, however, has a fenestrae, while T. duolamellata has only one. More more perforated shield. For this reason, our new significantly, however, the frontal shield in D. lon- is provisionally included in the Teuchoporidae. giuscula is entire with only about six areolar septular pores whereas T. duolamellata has up to a dozen pseudopores, mainly marginally placed (though some TORQUATELLA DUOLAMELLATA (SCHOLZ) COMB. NOV. of these might function as septular pores). These two (Fig. 11A,B) criteria, especially the latter, question their congeneric Osthimosia duolamellata Scholz, 1991: 302, pl. 11, figs status. Gordon (1993b) doubted the assignment of 1–6. Harmer’s (1957) species to Drepanophora but in the absence of any other material this problem remains Description unresolved. Examination of the inside of the frontal shield of T. Colony encrusting, multilaminar. Autozooids elongate duolamellata shows that this is not umbonuloid, i.e. polygonal, quincuncially arranged in early ontogeny, no ring scar is present. though obscured later in ontogeny by secondary cal- Scholz (1991) described the growth and ecology of cification and irregularly orientated frontal budding; this species. c. 0.36×0.21 mm. Frontal shield convex, granular, with up to a dozen fairly large pores, predominantly around the margin. Primary orifice suborbicular; anter Distribution large, smooth, rounded, separated from the wide, shal- Found on coral rubble from Poanangisu Erakor and low poster by small, squared, lateral condyles. Peri- Iririki Islands, T. duolamellata was first described stome flared, with a small, rounded proximal from the fringing reef flats of Cebu, the Philippines. avicularium on inner surface, laterally or fronto- laterally directed. Ovicell hyperstomial, globular, with a single, oval, transverse, frontal foramen. SUPERFAMILY SCHIZOPORELLOIDEA JULLIEN, 1883 FAMILY JULLIEN, 1883 GENUS SCHIZOPORELLA HINCKS, 1877 Remarks In his original description Scholz (1991) described the Type species: Johnston, 1847. ancestrula as tatiform, pointing out that it is often Schizoporella dunkeri (Reuss, 1848). Tilbrook, Hayward & Gordon, 2001, p.79, fig.15A. CHEILOSTOMATOUS BRYOZOA FROM VANUATU 81 difficult to observe, being obscured by the five peri- ancestrular zooids. It is in fact schizoporelloid, with SCHIZOPORELLA DUNKERI (REUSS) six oral spines around the D-shaped orifice. (Fig. 15A) Initially, the primary orifice sits within a slightly Cellepora dunkeri Reuss, 1848: 90, pl. 10, fig. 27. raised rim, however, this is superseded by a large Schizoporella unicornis form longirostris Hincks, 1886: chimney-like flared peristome. The peristome is con- 266, pl. 10, fig. 2. nected to the parent autozooid via a multiporous sep- Schizoporella longirostris: Hayward, 1976: 320, fig. 1; tulum and seems to develop from the cystid producing Hayward & Ryland, 1979 (in part): 173, not fig. 73. the proximal oral avicularium (seen on the inner sur- Schizoporella dunkeri: Hayward & Ryland, 1995b: 39, face of the peristome), and originating from one of the pls 2, 3. two most distolateral pores. The avicularian cystid also appears to cover most of the autozooidal frontal shield in secondary calcification. This happens early Description in ontogeny. The ovicell seems to be formed in a similar Colony encrusting, often extensive. Autozooids large, way, as a polymorph which evaginates from the distal rectangular, convex, distinct, separated by deep edge of the oral area (distal wall) of the maternal zooid. grooves. Frontal shield evenly perforated by numerous It rests on the frontal shield of the distal autozooid round pores, each in a shallow depression; a suboral producing the distal part of the peristome and being umbo may be present. Primary orifice as wide as long, incorporated into the secondary calcification of the almost circular; narrow, slit-like median sinus, often distal autozooid. with a rounded, proximal broadening; condyles broad Torquatella duolamellata is superficially similar to and flat. No oral spines. Adventitious avicularium Drepanophora longiuscula Harmer, 1957. Both have a single, lateral to the orifice and level with the proximal schizoporelloid ancestrula surrounded by five peri- border or sinus; cystid large and prominent; rostrum ancestrular zooids. The primary orifices are almost abruptly tapered distal to complete crossbar, with a identical, as is the development of a flared peristome, slender, acuminate tip; mandible acutely triangular, and accompanying avicularium, from a distolateral laterally or distolaterally orientated. Ovicells prom- 80 K. J. TILBROOK ET AL. pore. However, D. longiuscula has an ovicell with two inent, globular, regularly perforated.

Distribution Distribution Schizoporella dunkeri, described from the Miocene de- Schizoporella errata is a widespread warm temperate- posits of the Vienna Basin of Austria (Reuss, 1848), subtropical species. Originally described from Naples, shows a relict Tethyan distribution. It is known from Italy, it is found throughout the Mediterranean, West the Pliocene Coralline Crag of Britain and Pliocene Africa, eastern Canada, eastern coasts of the Americas, deposits in Belgium and The Netherlands (Bishop & from North Carolina through the Caribbean to Brazil, Hayward, 1989), is extant from the Canary Islands, Pacific coast of North America, Red Sea, Persian Gulf, Madeira, and the Isles of Scilly, and is common South Australia, and New Zealand. It was found sev- throughout the Mediterranean, especially the Aegean eral times in Port Vila Harbour, on old wall tiles. and Adriatic. It purportedly also occurs in the Red Sea (Hayward & Ryland, 1995b). Our record of two small GENUS ESCHARINA MILNE EDWARDS, 1836 colony fragments of Schizoporella dunkeri from debris at Erakor Island, Vanuatu is the most easterly and Type species: Eschara vulgaris Moll, 1803. most tropical record to date.

ESCHARINA PESANSERIS (SMITT) SCHIZOPORELLA ERRATA (WATERS) (Fig. 17A,B) (Fig. 15B) Hippothoa pes anseris Smitt, 1873: 43, pl. 7, figs 159, Lepralia errata Waters, 1878: 11, pl. 1, fig. 9. 160. Schizoporella errata: Hayward & Ryland, 1979: 170, Escharina pesanseris: Harmer, 1957: 998, pl. 67, figs fig. 68; Brock, 1985: 46, fig. 2C; Gordon & Mawatari, 12–14,18,19; Winston, 1984: 26, figs 53–55; Scholz, 1992: 31, pl. 9, fig. B. 1991: 311, pl. 14, fig. 4, Ryland & Hayward, 1992: 264.

Description Remarks Colony encrusting, unilaminar or multilaminar, or de- veloping erect, foliaceous, branched or cylindrical Escharina pesanseris is a well-known species that growth; whitish-pink to reddish-brown in life. Auto- apparently has a circumglobal distribution within zooids rectangular or lozenge-shaped, flat or slightly warm-temperate and tropical shallow waters. It is convex, separated by shallow grooves, in linear series; generally rather inconspicuous, with most colonies comprising a dozen or fewer zooids, many of which in multilaminar colonies, more disorientated inFigurfront-e 15. A, Schizoporella dunkeri. B, Schizoporella errata. C, Stylopoma novum. D, Stylopoma velatum. E, Stylopoma may be ovicellate. It was quite common at Port Vila ally budded additional layers. Frontal shield evenlyvilaensis. 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; Harbour on coral rubble, and was also found at Iririki perforated by numerous round pores, each in a shallowF=0.40 mm. depression. Primary orifice slightly wider than long, Island. appearing almost circular; with broad and shallow U- shaped median sinus. No oral spines. Adventitious GENUS STYLOPOMA CANU AND BASSLER, 1920 avicularium single, situated lateral to the orifice and level with the proximal border or sinus; cystid prom- Type species: Eschara spongites Pallas, 1766. inent, mandible acute triangular, angled to frontal plane, distolaterally orientated. Ovicells prominent, Remarks globular, regularly perforated. This speciose genus has a pan-tropical distribution in shallow, generally reefal, habitats. A review of the Remarks Indo-Pacific species of Stylopoma appears in the com- panion paper (pp. 1–34). Schizoporella errata is a well-known fouling species with many records from harbour installations or the hulls of ships. Compared with many other fouling STYLOPOMA NOVUM TILBROOK community species S. errata displays a rather catholic (Fig. 15C) choice of substratum, from pier pilings and shell, to Stylopoma novum Tilbrook, 2001: companion paper, algae and mangroves. Gordon & Mawatari (1992) docu- p. 24, Fig. 9D–F. ment its distribution in New Zealand and the Pacific, dating the first record in New Zealand at about 1960. The increase in cruise-liner traffic to Vanuatu over the Description last 20 years or so may have promoted the introduction Colony encrusting, multilaminar, often extensive. Au- of this species (as well as others) from other localities tozooids hexagonal or irregularly polygonal, flat or in the Southwest Pacific area. slightly convex, separated by distinct grooves;