A preliminary annotated checklist of the marine algae and seagrasses of the Wallis Islands (French Overseas Territory of Wallis and Futuna), South Pacific

Antoine De Ramon N&#39;Yeurt

Cette deuxieme publication dans une serie monographique sur la flore marine de la Polynesie francaise donne une distribution detaillee des especes de Chlorophycees qui sont presentes dans ces iles. Un total de 81 taxons sont presentes, dont 23 sont nouveaux pour la Polynesie francaise, 1 (Halimeda heteromorpha) represente une nouvelle espece et 2 (Microdictyon sp. et Struveopsis sp.) representent de nouveaux taxons. La flore a le plus d'affinites avec l'archipel voisin des iles Cook (Index de SΦrensen de 0.43), mais seulement avec 44 % des especes en commun avec ce groupe d'iles, tandis que 69 et 64 % des especes sont en commun avec les iles Fidji et la Micronesie, respectivement. Environ 75 % de la flore est en commun avec l'Ocean Indien et l'Ocean Pacifique en general; 3 especes (ou environ 4 % de la flore) semblent restreintes a la Polynesie francaise.

Occurrence and distribution of various species of benthic marine algae in Fiji and Rotuma were investigated, with the resulting floras being subjected to comparison indices (Jaccard) both between themselves and with respect to other neighboring floras in the Southwest Pacific. The Rotuman flora was found to be rather impoverished (106 species) with respect to the Fijian flora (314 species), and the Nauruan and Rotuman flora were most similar, as were the Fijian and Micronesian floras. Dispersal mechanisms for algae in the region were considered, with special attention to the major ocean currents. These currents tend to favor movements of algal species from more northerly locations toward Fiji, and dispersal in the opposite direction seems unlikely. This is assumed to explain the similarities of the Rotuman flora with the Nauruan and Micronesian floras. Peculiar distribution patterns of algae were examined, with one species (Meristotheca procumbens P. Gabrielson & Kraft) being found ...

CSIRO PUBLISHING www.publish.csiro.au/journals/asb Australian Systematic Botany 17, 367–397 A preliminary annotated checklist of the marine algae and seagrasses of the Wallis Islands (French Overseas Territory of Wallis and Futuna), South Pacific Antoine D. R. N’YeurtA,C and Claude E. PayriB A Laboratoire Terre-Ocean, Université de la Polynesié Française, BP 6570 Faa’a 98702, Tahiti, French Polynesia. UMR 7138, Systématique, Adaptation, Evolution, Equipe Biodiversité Marine Tropicale, IRD-Nouméa – BPA5, 98848 Nouméa cedex, New Caledonia. C Corresponding author; email: nyeurt@upf.pf B Abstract. A total of 194 species of marine algae (14 Cyanobacteria, 41 Chlorophyta, 11 Heterokontophyta and 128 Rhodophyta), as well as three species of seagrasses, represent the first published records for the isolated island of Wallis, South Pacific. The flora has its strongest affinities with Fiji and Rotuma, followed by Samoa and French Polynesia. The lack of diverse habitats and its geographical location are invoked to explain the relatively low species richness compared with localities such as Fiji and Samoa. The flora has a typically tropical component dominated by encrusting coralline red algae, the calcified green algal genera Halimeda, and assemblages of Cyanobacteria. Normally ubiquitous species such as Halimeda discoidea, and the brown algal genera Hydroclathrus, Colpomenia, Rosenvingea, Asteronema, and Chnoospora are notably absent from the island, perhaps due to seasonality and the lack of suitable habitats. The minute epiphytic red alga Acrochaetium kurogii is reported for the first time outside of its type locality in Japan, while two as yet unidentified species of red algae (Gracilaria sp. and Sebdenia sp.) could represent new taxa if further useful material is found. SB0327 MAa. rDi.nRe.pNal’YnsteufrotamndWaCl. iE.sPayri Introduction The Wallis Islands (Fig. 1) include Wallis Island, also known as Uvea Island, and 19 small islets that are located in the south-western Pacific between 13°18′S and 176°124′W, two-thirds of the way between Hawaii and New Zealand (Fig. 2). The Wallis Islands form an archipelago that also includes the islands of Futuna and Alofi. Wallis Island is a volcanic high island, of generally elongate shape, which represents the emerged portions of ancient craters. It is 15 km wide in a north–south direction, and 7.8 km long in an east–west direction. Emerged land area comprises 79.12 km2, with 129 km of coastline for a total reef-lagoon area of 219.5 km2. The highest point is 155 m tall Mt Lulu; the island’s geology is composed of volcanic tufs and basalts. The island was formed in several stages, the last being in the middle Pleistocene. The human population is approximately 10000, distributed in villages mainly on the eastern coast of the island. There is no previous information in the literature concerning the marine algae or seagrasses of Wallis Island, although there exists a report on the lagoons and reefs of Wallis and Futuna (Salvat 1982) with an excellent treatment of the corals, but which unfortunately only hints at the © CSIRO 26 August 2004 existence of two genera of marine algae. A few years later, a study was made of the terrestrial flora of Wallis, including mangroves (Morat and Veillon 1985). At the request of the Environmental Service of the French Overseas Territory of Wallis and Futuna, a survey was undertaken from 24 May to 8 June 2002, so as to establish a preliminary inventory of the coral fauna and benthic macroalgae/seagrass flora, and determine the ecological characteristics of the coral reef and lagoon habitats of Wallis (Payri et al. 2002). This paper deals with the marine benthic algae and seagrasses encountered in that study. Scattered algal floras and checklists exist for neighbouring islands such as Rotuma and Fiji (N’Yeurt 1996a, 1996b; N’Yeurt 2002; South and Skelton 2003, 2004), Samoa (Skelton and South 1999, 2002a, 2000b), the Solomon Islands (Womersley and Bailey 1970), Nauru (South and Yen 1992), Phoenix Group, Kiribati (South et al. 2001), the Cook Islands (Chapman 1977; N’Yeurt in prep.) and French Polynesia (Payri and N’Yeurt 1997; Payri et al. 2000). Recent monographs on the Lord Howe Island flora (e.g. Allender and Kraft 1983, Kraft 2000) and Norfolk Island (Millar 1999) have also proved very useful in identifying species occurring in more northerly islands such as Fiji, Rotuma and Wallis. Other areas of interest such as 10.1071/SB03027 1030-1887/04/040367 368 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Fig. 1. Map of the Wallis Islands, showing the 42 collecting stations. See Table 1 for co-ordinates. Tonga, Tokelau, Vanuatu, and much of Kiribati still remain largely unknown phycologically. The biogeographical implications of algal distribution in the region has been discussed by Millar (1990) and N’Yeurt and South (1997). Recently, Littler and Littler (2003) published a useful illustrated guide to South Pacific algae, adding many new records mainly to the Fijian flora, but also some for American Samoa, the Solomon Islands, the Cook Islands, French Polynesia and Papua New Guinea. Unfortunately, exact localities of collections are not mentioned in that publication. Materials and methods All material was collected using SCUBA, snorkeling or reef-walking. The position of each of the 42 collecting stations (Table 1) was determined with a Garmine 12 portable GPS receiver. Station numbers in the text correspond to the localities mentioned in Table 1. The 42 sites studied have been clustered into five large groups according to geomorphological criteria (fringing reef, barrier reef, lagoon, pinnacle and outer reef slopes; see Table 2 in the Results section). At the base station on land, specimens were sorted out and systematically photographed with a digital camera (Nikon Coolpix E-995, Nikon Corporation, Toyko, Japan) before processing. Microphotographs were taken with an Olympus C-4000Z digital camera (Olympus Optical Co. Marine plants from Wallis Fig. 2. Australian Systematic Botany 369 Map showing the position of the Wallis Islands (arrow) in relation to neighbouring island groups. (Europa) GMBH, Hamburg, Germany) mounted on an Olympus BH-2 microscope (Olympus Optical Co. Ltd., Tokyo, Japan). Voucher herbarium specimens were pressed by standard techniques, and representative parts of thalli and turf algae were stored in 4% buffered formalin in sealed plastic bags packed in a light-proof container, for shipment and later anatomical examination in the laboratory. Specimens are housed at the Phycological Herbarium of the Université de Polynésie française in Tahiti (UPF), with ‘WSS’ referring to slide collections. Results Geomorphology There is an almost continuous reef system around the main island of Uvea, composed of a barrier reef 63 km long encircling a lagoon 24 km wide in a north–south direction and 15 km long in an east–west direction, and fringing reefs extending up to the barrier reefs in the northern and western parts of the island. The lagoon communicates with the open sea through four passes, one in the South and three on the Western part of the barrier reef. Average depth of the lagoon varies between 10 and 20 m, with maximum depths of 47 m and 52 m in the Eastern and Southern parts, respectively. A total of 19 islets surround the main island, some of volcanic or coralline origin are present on the Eastern barrier reef, and volcanic islets with fringing reefs are present in the Eastern and Southern parts of the lagoon. Maximum tidal fluctuations are of 2 m, and water exchange in the lagoon occurs through the passes and over the barrier reefs. Systematic listing of species The taxonomy adopted generally follows that of Silva et al. (1987, 1996) supplemented by de Reviers (2003), and updated by other sources where necessary. To conserve space, type locality and basionym information are not given as these can be readily found in the sources cited above and in the text. For similar reasons, figures for species are omitted, but many of these can be found in Payri et al. (2002), upon request to the authors. Cyanobacteria This section mostly follows the classification of Velasquez (1962) and Silva and Pienaar (2000). Chroococcales Dermocarpellaceae Sphaenosiphon olivaceus Reinsch, 1875: 17. (= Dermocarpa olivacea (Reinsch) Tilden; Desikachary 1959: 174, pl. 33 figs 13, 14; Nagarkar 1998: 533). Vouchers: WSS 06 St. 5, WSS 53 St. 29, WSS 88 St. 1, 127 St. 8 370 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Table 1. List of collecting stations around Wallis, as shown in Fig. 1 Station no. Latitude Longitude 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 13°17'30.3"S 13°17'8.04"S 13°18'20.2"S 13°22'11"S 13°22'6"S 13°22'0.84"S 13°21'30"S 13°15'52.4"S 13°16'26.3"S 13°16'28.2"S 13°17'28.3"S 13°17'46.9"S 13°16'15.8"S 13°17'4.38"S 13°17'11.6"S 13°15'3.3"S 13°15'20.6"S 13°15'22.2"S 13°16'15"S 13°16'2.04"S 13°16'25.1"S 13°20'0.9"S 13°19'0.3"S 13°19'34.6"S 13°21'22.4"S 13°21'28"S 13°21'9.6"S 13°21'5.55"S 13°22'41.4"S 13°21'39.8"S 13°11'17.8"S 13°13'1.38"S 13°11'6.84"S 13°16'5.4"S 13°10'5.1"S 13°14'17.4"S 13°11'14.2"S 13°13'0.06"S 13°14'25.6"S 13°23'10.9"S 13°22'50"S 13°13'07.4"S 176°15'35.9"W 176°16'0.84"W 176°15'34"W 176°13'26.1"W 176°13'15.5"W 176°13'12.5"W 176°12'40"W 176°9'40.36"W 176°07'29"W 176°07'30"W 176°08'30.3"W 176°08'17.8"W 176°08'18.9"W 176°10'5.52"W 176°07'29"W 176°15'5.52"W 176°15'22.3"W 176°15'17.6"W 176°15'14.3"W 176°14'28.7"W 176°14'33.4"W 176°16'5.28"W 176°16'14.8"W 176°16'16.4"W 176°13'27.8"W 176°12'51.68"W 176°13'10.9"W 176°13'07.71"W 176°14'34.1"W 176°16'6.96"W 176°12'7.62"W 176°11'5.64"W 176°12'0.54"W 176°09'24.4"W 176°11'5.1"W 176°15'3.36"W 176°13'6.72"W 176°14'24"W 176°15'6.78"W 176°12'51.8"W 176°12'43.4"W 176°11'14.0"W Comments: this characteristically dumbell-shaped alga is commonly found as epiphytic colonies on macroalgae such as Laurencia, Hypnea. Nostocales Habitat Lagoon Outer slope Lagoon Fringing reef islet Fringing reef islet Fringing reef islet Fringing reef coastline Fringing reef coastline Inner barrier reef Inner barrier reef Lagoon Lagoon Lagoon Fringing reef coastline Inner barrier reef Outer slope Inner barrier reef Inner barrier reef Inner barrier reef Pinnacle Pinnacle Outer slope Inner barrier reef Inner barrier reef Pinnacle Fringing reef coastline Fringing reef coastline Fringing reef coastline Outer slope Outer slope Inner barrier reef Fringing reef coastline Inner barrier reef Fringing reef coastline Outer slope Outer slope Inner barrier reef Inner barrier reef Inner barrier reef Pass Lagoon Fringing reef coastline Scytonemataceae Scytonema polycystum Bornet et Flahault, 1887: 90. Silva and Pienaar 2000: 26, figs 48, 88–89. Vouchers: WSS 20 St. 26, 60 St. 31, 156 St. 7, 183 St. 40 Nostocaceae Oscillatoriales Nostoc calcicola Brebisson in Meneghini, 1842: 121. Velasquez 1962: 341, pl. 8 fig. 100. Vouchers: WSS 58 St. 31 Oscillatoriaceae Rivulariaceae Calothrix confervicola (Dillwyn) C. Agardh, 1824: 70. Velasquez 1962: 353, pl. 10 fig. 125. Vouchers: WSS 123 St. 22 Lyngbya majuscula (Dillwyn) Harvey, 1833: 370. Dawson 1954: 380, fig. 3d; Velasquez 1962: 319, pl. 4 fig. 68; Littler and Littler 1997: 122, fig. 182; N’Yeurt 2001: 692; Albert et al. 2002. Vouchers: WSS 88, 230 St. 1, 33 St. 2; UPF 1648, 1679 St. 3; WSS 158 St. 7, 125, 128 St. 8; UPF 1615 St. 16, 1407 St. 20; WSS 224 St 20; UPF 1511 St. 29, 1450 St. 30; WSS 57, 58 St. 31, 235 St. 35; UPF 1740 St. 36, Marine plants from Wallis Australian Systematic Botany Table 2. Distribution of the species within the five major geomorphological zones OS: outer slope (stations 2, 16, 22, 29, 30, 35, 36), BR: barrier reef (9, 10, 15, 17, 18, 19, 23, 24, 31, 33, 37, 38, 39), L: Lagoon (1, 3, 11, 12, 13, 41), P: Pinnacles (20, 21, 25, 40), FR: fringing reef (4, 5, 6, 7, 8, 14, 26, 27, 28, 32, 34, 42) Rhodophyta Chlorophyta Heterokontophyta Cyanobacteria Magnoliophyta Total OS BR L P FR 67 13 05 05 00 90 068 031 004 012 000 115 067 021 009 008 001 106 13 07 01 03 00 24 080 030 010 007 003 130 1517, 1518 St. 41; WSS 226 St. 14.1; UPF 1603 St. 4; WSS 06 St. 5; UPF 1532 St. 10, 1498 St. 11 Comments: this species is found at most sites, its colour ranging from greenish to brownish-black. Lyngbya polychroa (Meneghini) Rabenhorst, 1847: 83. Velasquez 1962: 316, pl. 3 fig. 64 (‘Lynbya sordida’); Littler and Littler 1997: 183. Vouchers: UPF 1686 St. 3; WSS 206 St. 31 Comments: this species (often reported under the synonym L. sordida Gomont, see Silva et al. 1996) is distinguished from Lyngbya majuscula by the smaller average diameter of its filaments, which are between 16 and 30 µm as opposed to 20–60 µm in L. majuscula (Velasquez 1962). The sheath is also thinner, and cell contents appears less dense, in L. polychroa. Oscillatoria bonnemaisonii (P. Crouan et H. Crouan) P. Crouan et H. Crouan, 1860: 371. Velasquez 1962: 288, pl. 1 fig. 16. Voucher: WSS 206 St. 31; UPF 1522 St. 41 Oscillatoria margaritifera (Kützing) Gomont, 1892b: 216. Dawson 1954: 380, fig. 3f; Velasquez 1962: 293, pl. 2 fig. 23. Voucher: WSS 206 St. 31 Phormidiaceae Phormidium nigroviride (Thwaites) Komárek, 1988: 405. Anagnostidis et Comments: = Oscillatoria nigroviride Thwaites; Dawson 1954: 380, fig. 3g; Velasquez 1962): 293, pl. 2 fig. 24 . Vouchers: WSS 59 St. 31; UPF 1519 St. 41 Phormidium papyraceum Gomont, 1890: 355. Velasquez 1962: 310, pl. 3 fig. 57. Vouchers: UPF 1553, WSS 85 St. 1; UPF 1707 St. 20; WSS 227 St. 14.1 Comments: forms thin sheets on sandy substrata in the lagoon. Filaments interwoven, 3–4 µm in diameter, with prominent transverse walls. When dry, specimens become translucent with a papery texture. 371 Spirulina subsalsa Oersted, 1842: 566, pl. 7 fig. 4. Velasquez 1962: 284, pl. 1 fig. 13; Nagarkar 1998: 534, fig. 16. Vouchers: WSS 57, 206 St. 31 Comments: this characteristically coil-shaped minute Cyanobacteria is often found mixed with other algae in turf populations. Symploca atlantica Gomont, 1892a: 109, pl. 2 fig. 5. Velasquez 1962: 312; Littler and Littler 1997: 123, fig. 184. Vouchers: UPF 1617 St. 16; WSS 213 St. 23, 216 St. 36, 138 St. 4 Comments: forming erect fascicles; filaments 4–5 µm in diameter. Symploca hydnoides (Harvey) Kützing, 1849: 272. Dawson 1954: 380, fig. 3o, p; Velasquez 1962: 312, pl. 3 fig. 59; Islam and Aziz 1982: 650, pl. 2 figs 13–16, pl. 7 figs 34–35; Payri et al. 2000: 293; Payri et al. 2002: 53, pl. 6 fig. 1; Abed et al. 2003: 867, fig. 7a–d. Vouchers: WSS 88 St. 1, 124 St. 8; UPF 1568 St. 16, 1437 St. 25; WSS 62 St. 31; UPF 1527 St. 41, 1287 St. 4, 1479 St. 5, 1541 St. 12 Comments: this species forms characteristic erect fascicles with a clay-like texture; filaments 6–7 µm in diameter. Symploca laeteviridis Gomont, 1892b: 109, pl. 2 figs 6–8. Velasquez 1962: 311, pl. 3 fig. 58; Nagarkar 1998: 539, fig. 25. Voucher: WSS 30 St. 1; UPF 1490 St. 5 Comments: this species is distinguished from S. hydnoides by the smaller diameter of the filaments (3–4 µm, never reaching 5 µm) and the absence of adhesion of filaments into erect fascicles. Chlorophyta Ulvales Ulvaceae Ulva clathrata (Roth) C. Agardh, 1811: 23. (= Enteromorpha clathrata (Roth) Greville; Dawson 1954: 384, fig. 6d–e; Egerod 1974: 134, fig. 4; Kraft 2000: 525, fig. 8; N’Yeurt 2001: 693, figs 2, 3a–b; Skelton and South 2002a: 160, fig. 22A–B; 2002b: 8, pl. 5 figs 29–33, pl. 6 fig. 34; Abbott and Huisman 2004: 46, fig. 5A–C). Vouchers: WSS 60 St. 31 Comments: based on culture and molecular studies, Shimada et al. (2003) suggested that Japanese records of Enteromorpha should be transferred to Ulva. Most species previously listed under the genus Enteromorpha Link (1820) have been transferred to Ulva by Hayden et al. (2003) based on their own molecular studies. Ulva clathrata is readily distinguished by multiseriate laterals ending in bulbous expansions. Ulva flexuosa Wulfen subsp. paradoxa (Dillwyn) Bliding, 1963: 79. (= Enteromorpha flexuosa (Wulfen) J. Agardh subsp. paradoxa; Bliding 1963: 79, figs 42a–g, 45a–f; 372 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Egerod 1974: 132, figs 1–3; N’Yeurt 1996: 365, fig. 16; Kraft 2000: 525, fig. 7F; N’Yeurt 2001: 697, fig. 46). Vouchers: WSS 228, 229 St. 3 Comments: this subspecies is characterised by its longer uniseriate laterals (up to 2 mm long) and regularly transversely arranged cell rows. Cladophorales Cladophoraceae Chaetomorpha crassa (C. Agardh) Kützing, 1845: 204. Sartoni 1992: 299, fig. 4E; Littler and Littler 1997: 93, fig. 125; N’Yeurt 2001: 701, fig. 15. Vouchers: UPF 1336, 1337, 1338 St. 8 Chaetomorpha linum (O.F. Müller) Kützing, 1845: 204. Egerod 1974: 135, fig. 9; Littler and Littler 1997: 93, fig. 126; 2003: 198. Vouchers: UPF 1583; WSS 112 St. 8 188, figs 30–33; Kraft 2000: 578, fig. 27A–B; Payri et al. 2000: 76; Littler and Littler 2003: 202; Abbott and Huisman 2004: 89, fig. 29A. Vouchers: UPF 1554, 1657, WSS 166 St. 1, 1286 St. 4, 1330 St. 9, 1305 St. 12, 1297 St. 13 Dictyosphaeria versluysii Weber-van Bosse, 1905: 144. Egerod 1952: 351, figs 1a, 2h–k; Sartoni 1992: 319, figs 13B, 14A; Leliaert et al. 1998: 190, figs 37–39; Payri et al. 2000: 76; Payri et al. 2002: 44, pl. 1 fig. 11; Skelton and South 2002a: 162, fig. 23C, D; Littler and Littler 2003: 204; Abbott and Huisman 2004: 89, fig. 29B. Vouchers: UPF 1289, 1595 St. 4, 1218 St. 7, 1339 St. 8, 1319 St. 10, 1335 St. 9 Phyllodictyon anastomosans (Harvey) Kraft et Wynne, 1996: 139, figs 16–25. Leliaert et al. 1998: 186, figs 23–24; Payri et al. 2000: 74; Wynne 2001: 368, fig. 34; Littler and Littler 2003: 200, Abbott and Huisman 2004: 63, fig. 16A, B. Vouchers: UPF 1220, 1551, 1555 St. 1, 1476 St. 5; WSS 62 St. 31 Cladophora vagabunda (Linnaeus) van den Hoek, 1963: 144. Sartoni 1992: 304, fig. 6C–E; Littler and Littler 1997: 95, fig. 130; Kraft 2000: 562, fig. 21A–G; Leliaert and Coppejans 2003: 70, figs 14A–B, 15; Abbott and Huisman 2004: 79, fig. 24A–D. Vouchers: UPF 1221 St. 1, 1373 St. 8, 1562 St. 9 Siphonocladus tropicus (P. Crouan et H. Crouan) J. Agardh, 1887: 105. Sartoni 1992: 315, fig. 12A; Abbott and Huisman 2004: 90, fig. 30. Voucher: UPF 1633, St. 7 Siphonocladaceae Valoniaceae This family was previously listed under the Siphonocladales, but recent morphological and molecular evidence suggests that the latter order and the Cladophorales are monophyletic (Leliaert et al. 1998: 178, 2003). Valonia aegagropila C. Agardh, 1823: 429. N’Yeurt 1996: 372, fig. 15a, b; Leliaert et al. 1998: 192, figs 40, 41; Payri et al. 2000: 78; Littler and Littler 2003: 206; Abbott and Huisman 2004: 92, fig. 31A. Vouchers: UPF 1230 St. 7, 1351 St. 8 Boergesenia forbesii (Harvey) J. Feldmann, 1938: 1503, figs 3–5. Sartoni 1992: 306, fig. 7B; N’Yeurt 1996: 369, fig. 14; Leliaert et al. 1998: 184, fig. 13; Payri et al. 2002: 44, pl. 1 fig. 10; Littler and Littler 2003: 202. Voucher: UPF 1354 St. 8 Boodlea composita (Harvey) Brand, 1904: 187–190. Sartoni 1992: 306, fig. 7C; N’Yeurt 1996: 368, fig. 19 (as Boodlea coacta (Dickie) Murray); Leliart et al. 1998: 184, figs 16–19 (non figs 14–15, 20, F. Leliaert pers. com.); Kraft 2000: 569, fig. 24A–C; Payri et al. 2000: 70 (as Boodlea kaeneana Brand); Littler and Littler 2003: 200; Abbott and Huisman 2004: 85, fig. 26A–B. Vouchers: UPF 1219, 1546 St. 1, 1202 St. 2, 1189 St. 3, 1231 St. 7, 1308 St. 11, 1426, 1731 St. 23 Comments: this species forms spongy masses amidst spaces between dead coral. This genus (along with Phyllodictyon and several others) form a close cluster with Cladophoropsis in the family Cladophorophyceae based on recent phylogenetic studies (Kooistra et al. 1993; Wysor 2002; Leliaert et al. 2003) and a single, highly variable species may need to be recognised (F. Leliaert, pers. comm.). Dictyosphaeria cavernosa (Forsskål) Børgesen, 1932: 2. Sartoni 1992: 319, fig. 13A; N’Yeurt 1996: 371, fig. 12; Littler and Littler 1997: 87, fig. 117; Leliaert et al. 1998: Comments: see remarks above for Boodlea composita. Valonia fastigiata Harvey ex. J. Agardh, 1887: 101, pl. I fig. 5. Sartoni 1992: 321, fig. 14C; Coppejans et al. 1995: 98, fig. 39; Leliaert et al. 1998: 192, figs 42–44; Payri et al. 2000: 78; Payri et al. 2002: 44, pl. 1 fig. 7; Skelton and South 2002a: 162, fig. 24A; Littler and Littler 2003: 206. Vouchers: UPF 1217 St. 1, 1187, 1684 St. 3, 1281, 1596 St. 4, 1249, 1250 St. 7, 1340 St. 8, 1332 St. 9, 1309 St. 12 Comments: this species is quite common in Wallis, especially on the inner reef slopes where it forms abundant tightly adhering masses nested in the infractuosities of dead coral. It is also found in the seagrass beds, where it assumes an unusual, ball-like unattached form up to 10 cm in diameter. Valonia macrophysa Kützing, 1843: 307. Littler and Littler 1997: 91, fig. 121; Payri et al. 2000: 80; Littler and Littler 2003: 206. Vouchers: UPF 1631, 1632 St. 7, 1703 St. 20 Ventricaria ventricosa (J. Agardh) Olsen et West, 1988: 104. Sartoni 1992: 323, fig. 14E; N’Yeurt 1996: 372, fig. 23; Littler and Littler 1997: 89, fig. 119; Payri et al. 2000: 80; Payri et al. 2002: 44, pl. 1 fig. 12; Littler and Littler 2003: 204; Abbott and Huisman 2004: 94, fig. 32B. Vouchers: UPF 1226 St. 3, 1471 St. 5, 1329 St. 9, 1306 St. 12, 1377 St. 16 Marine plants from Wallis Bryopsidales Bryopsidaceae Bryopsis pennata Lamouroux var. secunda (Harvey) Collins et Hervey, 1917: 62. Mitchell et al. 1979: 109, pl. 1 figs 6–8; Littler and Littler 1997: 97, fig. 134; Payri et al. 2000: 82; Skelton and South 2002a: 163, fig. 24E; Littler and Littler 2003: 208; Abbott and Huisman 2004: 98, fig. 33C. Vouchers: UPF 1656; WSS 178 St. 1; UPF 1481 St. 2, 1566 St. 16 Australian Systematic Botany 373 and South and Skelton (2003: 540) reported C. biserrulata fronds to be thin and delicate, but Wallis plants were found to be quite coriaceous with thick assimilators, while still possessing conspicuous double rows of marginal teeth (cf. Payri et al. 2002, pl. 1 fig. 1). Joly and Semir (1973) described a further variety of C. brachypus from Brazil, var. nordestina, which has undulate margins and lacks marginal proliferations. The latter authors appropriately noted that it could represent a separate species, and it may be quite distinct from the Pacific material. Codium bulbopilum Setchell, 1924: 173, fig. 38; 1926: 84, pl. 11 fig. 1, pl. 12 fig. 2. Jones and Kraft 1984: 26, figs 4, 5B–F; N’Yeurt 1996: 384, figs 42, 45; Kraft 2000: 591, fig. 31F; Skelton and South 2002a: 165, fig. 25F; Littler and Littler 2003: 212. Voucher: UPF 1317, St. 10 Caulerpa cupressoides (Vahl) C. Agardh var. mamillosa (Montagne) Weber-van Bosse, 1898: 332, pl. 28, fig. 6. Sartoni 1978: 402, fig. 2b; Coppejans and Prud’homme van Reine 1992: 679, figs 3A, 8B; N’Yeurt 1996: 377, figs 25, 35; Kraft 2000: 597, fig. 32C, D; Payri et al. 2002: 44, pl. 1 fig. 6; Littler and Littler 2003: 216. Vouchers: UPF 1399, 1400 St. 19, 1456 St 26, 1242, 1243, 1245 St. 6 Comments: the Wallis material agrees with the hummock-like habit of Lord Howe plants in Kraft (2000). The relationship of Pacific records of C. geppiorum (e.g. Payri et al. 2000: 102) to C. bulbopilum remain unclear pending molecular studies (Skelton and South 2002). Caulerpa nummularia Harvey ex J. Agardh, 1873: 38. Littler and Littler 1997: 105, fig. 150; Littler and Littler 2003: 214; South and Skelton 2003: 542, fig. 3; Abbott and Huisman 2004: 121, fig. 44A. Vouchers: UPF 1363 St. 8, 1390 St. 17; WSS 234 St. 35 Codium mamillosum Harvey, 1855: 565. Womersley 1984: 230, figs 77A, 78A; Abbott 1986: 162, fig. 1; N’Yeurt 2001: 720, figs 48, 50, 69; Littler and Littler 2003: 212; Abbott and Huisman 2004: 109, fig. 39A–B. Voucher: UPF 1650, WSS 162 St. 1 Comments: this species is distinguished from superficially similar C. peltata by its crenulated disc-like assimilators which are tiered, as opposed to smooth, single assimilators in the latter species (Kraft 2000; Abbott and Huisman 2004). Codiaceae Comments: a common species at depths below 20 m on the outer reef slope, it can form large dominant covers at depths of up to 100 m in many parts of the South Pacific such as Fiji and McDonald Seamount (ADR N’Yeurt unpubl. obs). Caulerpa peltata Lamouroux, 1809b: 332. Weber-van Bosse 1898: 373, pl. 31 fig. 9; Kraft 2000: 601, fig. 33E; N’Yeurt 2001: 714; Skelton and South 2002a: 163, fig. 25B. Vouchers: UPF 1550 St. 1, 1195 St. 3, 1362 St. 8, 1708 St. 17, 1429 St. 23, 1513 St. 31, 1272 St. 5, 1533 St. 12 Caulerpaceae Comments: Caulerpa peltata is distinguished from C. racemosa var. peltata (Lamouroux) Eubank by having thin stolons with simple, never compound, disk-like assimilators which never bear clavate ramuli. The two entities are best kept separate until genetic and molecular studies provide a definitive answer on their status (Kraft 2000). Caulerpa biserrulata Sonder, 1871: 64, pl. 2 figs 10–12. Coppejans and al. 1995a: 74, figs 3, 4; Kraft 2000: 595, fig. 32A, B; Littler and Littler 2003: 214 (= Caulerpa brachypus Harvey forma parvifolia (Harvey) Cribb sensu Cribb 1958: 209, figs 1–7; Coppejans and Meinesz 1988: 184, figs 35–38; Coppejans and Prud’homme van Reine 1992: 673, figs 1A–C, 7A–B; N’Yeurt 2001: 711, fig. 40; Payri et al. 2002: 44, pl. 1 fig. 1). Vouchers: UPF 1360 St. 8, 1295 St. 13, 1402 St.14, 1453 St. 31 Comments: the Wallis plants have both simple, smooth and divided, toothed uprights, forming a continuum which circumscribes Caulerpa brachypus forma parvifolia (Cribb 1958b) and conforms with plants described from Papua New Guinea by Coppejans et al. (1995a). Caulerpa brachypus forma parvifolia was put in synonymy with C. biserrulata (Coppejans 1992: 388), and is distinguished from superficially similar C. brachypus by its double or triple rows of spines along the assimilator margins. Kraft (2000: 597) Caulerpa racemosa (Forsskål) J. Agardh, 1873: 35. Sartoni 1978: 406, fig. 4c; Coppejans and Meinesz 1988: 191, figs 22, 23; Coppejans 1992: 397, fig. 4C; Coppejans and Prud’homme van Reine 1992: 698, fig. 18A–B; Coppejans and al. 1995a: 78, fig. 7; N’Yeurt 1996: 378, figs 28, 36; Kraft 2000: 602, fig. 34A–D; Payri et al. 2000: 94; N’Yeurt 2001: 714, fig. 43; Payri et al. 2002: 44, pl. 1 fig. 2; Littler and Littler 2003: 226; Abbott and Huisman 2004: 122, fig. 44B, C. Vouchers: UPF 1251 St. 7, 1298 St. 13, 1403, 1404 St. 19, 1451 St. 31 Comments: includes C. racemosa vars peltata and uvifera; see Silva et al. (1996). 374 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Caulerpa serrulata (Forsskål) J. Agardh, 1837: 174. Sartoni 1978: 408, fig. 5a; Meñez and Calumpong 1982: 9, pl. 2E; Coppejans and Meinesz 1988: 192, figs 27, 28; Coppejans 1992: 403, fig. 7; Coppejans and al. 1992: 701, fig. 20B; 1995a: 78, figs 9, 10; N’Yeurt 1996: 382, figs 30, 40; Payri et al. 2000: 94; Leliaert et al. 2001: 452, fig. 1; N’Yeurt 2001: 715, fig. 44; Payri et al. 2002: 44, pl. 1 fig. 3; Skelton and South 2002a: 164, figs 24B–D; Littler and Littler 2003: 230; Abbott and Huisman 2004: 123, fig. 45A. Vouchers: UPF 1216, 1654 St. 1, 1232, 1233, 1234, 1235, 1236 St. 6, 1320 St. 10, 1296 St. 13, 1405 St. 20 Caulerpa sertularioides (S. G. Gmelin) Howe, 1905: 576. Sartoni 1978: 410, fig. 5b; Coppejans 1992: 404; Coppejans and Prud’homme van Reine 1992: 704, fig. 21A; Coppejans and al. 1995a: 80, fig. 8; Littler and Littler 1997: 107, fig. 154; Payri et al. 2000: 96; N’Yeurt 2001: 716, fig. 42; Payri et al. 2002: pl. 1 fig. 4; Littler and Littler 2003: 232; Abbott and Huisman 2004: 124, fig. 45B, C. Vouchers: UPF 1365 St. 8, 1237, 1238, 1239, 1240, 1241, 1244 St. 6, 1392 St. 19 Caulerpa taxifolia (Vahl) C. Agardh, 1817: 22. Sartoni 1978: 410, fig. 5c; Coppejans 1992: 406, fig. 8A–G; Coppejans and Prud’homme van Reine 1992: 706, figs 6B, 22B; South and N’Yeurt 1993: 122, fig. 17; Coppejans and al. 1995: 80, fig. 11; Littler and Littler 1997: 109, fig. 155; Kraft 2000: 604, figs 34E, 35A; Payri et al. 2000: 98; Leliaert et al. 2001: 452, fig. 2; Payri et al. 2002: 44, pl. 1 fig. 5; Littler and Littler 2003: 234; Abbott and Huisman 2004: 124, fig. 46A-B. Vouchers: UPF 1196, 1692 St. 3, 1710 St. 17, 1393, 1394, 1395, 1396, 1397, 1398 St. 19 Comments: Coppejans and Prud’homme van Reine (1992: 706) reported an ecad ‘mexicana’ of C. taxifolia from Indonesia, with partly compressed rachi, and basally constricted compressed pinnae. Later, Littler and Littler (2003: 222) reported a plant from Fiji with similar characteristics under Caulerpa mexicana Sonder ex Kützing. Caulerpa taxifolia and C. mexicana are distinct species (Olsen et al. 1998). Caulerpa mexicana from the Caribbean is characterised by a flattened main axis bearing non-constricted pinnae; the fronds also have a darker green colour with a lustrous sheen not found in C. taxifolia. (Meinesz et al. 1994; Littler and Littler 2000: 364, fig. on 365, upper). A main difference between the two species would be in the disposition of the rhizoids: sparse on the stolon and very thick and dense in C. taxifolia; close together and with few rhizoids per shoot in C. mexicana. Taylor (1977) cited C. mexicana to be present in the Hawaiian flora, but after re-examination of the herbarium sheets in question by Abbott and Huisman (2004: 124) they were found to be not distinct from C. taxifolia. True C. mexicana has yet to be confirmed from the South Pacific (A. Meinesz, pers. com.) and present records of this species from Fiji and other regional localities would best be ascribed to C. taxifolia. It would appear however to be present on the Andaman Sea coast, Thailand (Egerod 1975: 5, fig. 23). Caulerpa taxifolia (Vahl) C. Agardh ecad. tristichophylla Svedelius 1906: 112. fig. 5. Littler and Littler 2003: 222 (‘Caulerpa mexicana var. pluriseriata’) (= Caulerpa taxifolia var. asplenioides Harvey 1863: pl. 178). Voucher: UPF 1711 St. 17 Comments: this ecad is distinguished from the typical form of C. taxifolia by its three-dimentional, often radial arrangement of pinnulae on erect assimilators. Quite commonly, both two-dimentionally and three-dimentionally beset assimilators are found on the same plant. Also reported from Fiji (ADR N’Yeurt, unpubl. data), and Mangareva Island, French Polynesia (Weber-van Bosse 1910 (as C. taxifolia f. asplenioides); ADR N’Yeurt and CE Payri unpubl. data). This appears to be a shallow-water ecotype of C. taxifolia, rather than an infraspecific taxon (A Meinesz, pers. comm.). Plants from Fiji reported by Littler and Littler (2003) as Caulerpa mexicana var. pluriseriata and collections in SUVA-A examined by the first author conform to the Wallis material as C. taxifolia ecad. tristichophylla as understood here (see comments above on C. taxifolia). Weber-van Bosse (1898: 292; 1910: 2) reported this ecad under C. taxifolia f. asplenioides, based on C. asplenioides Greville (1853: figs 1, 2, pl. I) and C. taxifolia var. asplenioides (Greville) Harvey (1863: pl. 178). The latter forms and varieties were invalidated by Meinesz et al. (1984): 108) since the original description of C. asplenioides was found to represent C. mexicana. Halimedaceae The systematics of the Halimedaceae has recently been revised in molecular studies (Kooistra et al. 2002; Verbruggen and Kooistra 2004). Halimeda borneensis W.R. Taylor 1975: 81, figs 1, 2. (Misapplied name: Halimeda simulans Howe: Valet 1968: 48, pl. 9(4) fig. 3; Hillis 1980: 103, fig. 26; N’Yeurt 1996: 390, figs 76, 85; Littler and Littler 1997: 113, fig. 163; N’Yeurt 2001: 274); Hillis 1980: 105, fig. 27; Coppejans et al. 2001b: 411; Littler and Littler 2003: 240. Vouchers: UPF 1779 St. 3, 1781 St. 4, 1179, 1784, 1785, 1786 St. 5, 1782, 1783 St. 9, 1790 St. 14 Comments: previous Pacific records of Halimeda simulans are referable to H. borneensis (H. Verbruggen pers. com.) as H. simulans is strictly an Atlantic species (Kooistra et al. 2002: 134). Halimeda distorta (Yamada) Hillis 1968 : 33, figs 4, 6(2). (Misapplied name: Halimeda copiosa Goreau et Graham: Abbott and Huisman 2004: 131, fig. 49A). Hillis 1980: 120, fig. 34; 1985: 11; Verheij and Prud’homme van Reine 1993: 135, pl. 5 fig. 4; Payri et al. 2000: 108. Vouchers: UPF 2733 Marine plants from Wallis St. 1, 1172 St. 10, 1181, 1819 St. 11, 1823 St. 24, 1822 St. 31, 1818 St. 38 Comments: new molecular data suggests that previous Pacific records of H. copiosa are referable to H. distorta, as the former is exclusively an Atlantic Ocean species (H. Verbruggen, pers. comm.). H. distorta is characterised by having a single, basal holdfast and heavily calcified, contorted segments, which distinguishes it from superficially similar H. hederacea which has multiple rhizoidal holdfasts and keeled, uncontorted segments. The latest molecular evidence points to a merger of H. distorta and H. hederacea, but it is felt that further research might separate the two species again (H. Verbruggen, pers. comm.). Halimeda gracilis Harvey ex J. Agardh, 1887: 82. Coppejans et al. 1995: 86, fig. 24; Leliaert et al. 2001: 453, figs 3, 9–10; Littler and Littler 2003: 246; Abbott and Huisman 2004: 133, fig. 49F. Vouchers: UPF 1167, 1820, 1821 St. 30 Halimeda hederacea (Barton) Hillis 1968: 30, figs 3, 6:1, 6:4-8. (Misapplied name: Halimeda copiosa Goreau et Graham: Coppejans et al. 1995: 84, fig. 21). Payri et al. 2002: 46, pl. 2 fig. 1; Littler and Littler 2003: 242. Vouchers: UPF 1823 St. 29, 1822 St. 36 (= Halimeda opuntia var. hederacea (Barton) Hillis 1959: 360, pl. 2 fig. 7, pl. 5 fig. 4; Taylor 1950: 81, pl. 40 fig. 1; N’Yeurt 1996: 389, figs 70c, 84) Comments: This species is often listed as a variety or synonym of H. opuntia, but we prefer to maintain its specietal status in light of new molecular evidence (Kooistra et al. 2002; Verbruggen and Kooistra 2004). Additionally, it has a sprawling habit with smooth, shiny segments, as opposed to the clumped habit with dull, rough segments of H. opuntia. Halimeda incrassata (Ellis) Lamouroux 1816: 307; Hillis 1959: 365, pl. 4 figs 1, 2, pl. fig. 21, pl. 6 figs 21–24, pl. 12; Egerod 1974: 147, figs 62–64; Littler and Littler 1997: 111, fig. 159; Payri et al. 2000: 110; N’Yeurt 2001: 721, figs 26, 37; Payri et al. 2002: 46, pl. 2 fig. 3; Littler and Littler 2003: 246; Abbott and Huisman 2004: 133, fig. 50A. Vouchers: UPF 1779 St. 3, 1185, 1781 St. 6, 1179, 1784, 1785, 1786 St. 7, 1782, 1783 St 9 Halimeda macroloba Decaisne, 1841: 118. Hillis 1959: 375, pl. 3 fig. 3, pl. 5 figs 19, 20, pl. 6 fig. 17, pl. 12; 1980: 108, fig. 28; Egerod 1974: 148, figs 65–68; 1975: 61, fig. 33; Coppejans et al. 1995: 86, fig. 25; Payri et al. 2000: 112; N’Yeurt 2001: 722, fig. 38; Payri et al. 2002: 46, pl. 2 fig. 4; Littler and Littler 2003: 248; Abbott and Huisman 2004: 135, fig. 50B. Vouchers: UPF 1178, 1802 St. 6, 1803 St. 7, 1176, 1800, 1801 St. 8, 1799 St. 20 Comments: this species is the largest Halimeda in Wallis, with characteristic bulbous holdfast and segments to 2 cm in diameter. Australian Systematic Botany 375 Halimeda micronesica Yamada, 1941: 121, fig. 15. Hillis 1959: 364, pl. 3 fig. 1, pl. 5 figs 13, 14, pl. 6 fig. 2, pl. 9; Wynne 1993: 22, fig. 10; N’Yeurt 1996: 387, figs 69, 81–82; Payri et al. 2000: 112; Payri et al. 2002: 46, pl. 2 fig. 5; Littler and Littler 2003: 250. Vouchers: UPF 1832, 1833, 1834 St. 10, 1832 St. 35 Comments: the fan-shaped, fused basal segment of this species is distinctive in the field. Common in neighbouring Rotuma Island (N’Yeurt 1996), it was recently also found in Fiji (Littler and Littler 2003). Halimeda minima (W. R. Taylor) Hillis, 1968: 32. Hillis 1980: 113, fig. 30; Payri et al. 2000: 114; N’Yeurt 2001: 723, figs 25, 36; Littler and Littler 2003: 250. Vouchers: UPF 1169 St. 1, 1685 St. 3, 1177, 1792 St. 10, 1793 St. 9, 1794 St. 12, 1389 St. 17, 1797 St. 22, 1795 St. 23, 1173, 1787 St. 24, 1790 St. 25, 1796, 1798 St. 29, 1168 St. 30, 1788 St. 35, 1171 St. 36, 1791 St. 38 Halimeda opuntia (Linnaeus) Lamouroux, 1816: 308. Hillis 1959: 359, pl. 2 figs 7, 8, pl. 5 figs 3, 4, pl. 6 fig. 6, pl. 7 fig. 3, pl. 10; Egerod 1974: 147, figs 59–61; Coppejans et al. 1995: 86, fig. 27; N’Yeurt 1996: 388, figs 70a–c, 83; Littler and Littler 1997: 111; Payri et al. 2000: 114; N’Yeurt 2001: 723; Payri et al. 2002: 46, pl. 2 fig. 2; Skelton and South 2002a: 165, fig. 25C–D; Littler and Littler 2003: 252; Abbott and Huisman 2004: 135, fig. 50C. Vouchers: UPF 1170, 1805, 1827, St. 1, 1804 St. 3, 1807 St. 4, 1809 St. 5, 1811 St. 6, 1180, 1810 St. 7, 1812 St 6, 1184, 1808, 1829, 1830, 1831 St. 10, 1182 St. 11, 1813 St. 13, 1385 St. 16, 1814 St. 17, 1824 St. 23, 1174, 1815, 1825, 1826 St. 24, 1828 St. 35 Halimeda taenicola W. R. Taylor, 1950: 86, 207, pl. 46 fig. 1. Hillis 1959: 354, pl. 2 fig. 6, pl. 5 fig. 12, pl. 6 fig. 14, pls 11, 14; Hillis 1980: 139, fig. 42; N’Yeurt 1996: 390, figs 72, 74, 86; Payri et al. 2000: 116; Payri et al. 2002: 46, pl. 2 fig. 6; Littler and Littler 2003: 252. Vouchers: UPF 1817 St. 10, 1175, 1816 St. 9 Comments: this normally widely distributed tropical species was in Wallis restricted to the barrier reef on the eastern part of the island. Udoteaceae Chlorodesmis fastigiata (C. Agardh) Ducker, 1969: 17, fig. 1. Coppejans and Prud’homme van Reine 1989: 127, pl. 3 figs 1–4, 12; Payri et al. 2000: 120; Coppejans et al. 2001b: 421, figs 22–31; N’Yeurt 2001: 726, fig. 73; Payri et al. 2002: 44, pl. 1 fig. 8; Skelton and South 2002a: 165, fig. 26B; Littler and Littler 2003: 238. Vouchers: UPF 1299 St. 11, 1307 St. 12, 1457 St. 31 Chlorodesmis hildebrandtii A et E. S. Gepp, 1911: 16, pl. 8 figs 74–75. Egerod 1974: 143, figs 44–49; Coppejans and Prud’homme van Reine 1989: 129, pl. 3 figs 5–11; N’Yeurt 1996: 393, figs 54, 63; Coppejans et al. 2001b: p, 422, 376 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany figs 15–21; Littler and Littler 2003: 240. Vouchers: UPF 1267 St. 7; WSS 67 St. 10, 1545 St. 12 Comments: crystals were not seen in Wallis plants. Chlorodesmis hildebrandtii is distinguished from C. fastigiata by its truncated, equally constricted dichotomies; the latter species has unequal constrictions above dichotomies. Abbott and Huisman (2004: 138) follow the opinion of Ducker (1967) in ascribing previous Pacific records of C. hildebrandtii to C. caespitosa J. Agardh, but other authors (Coppejans et al. 2001; Littler and Littler 2003) retain the name for Pacific material. Rhipidosiphon javensis Montagne, 1842a: 15. Littler and Littler 1990: 35; Verheij and Prud’homme van Reine 1993: 140, pl. 7 fig. 6; Wynne 1993: 23, fig. 14; N’Yeurt 1996: 394, fig. 61; Payri et al. 2000: 120; Coppejans et al. 2001b: 422, figs 32–34; N’Yeurt 2001: 726, fig. 33; Littler and Littler 2003: 254; Abbott and Huisman 2004: 140, fig. 52C–D (= Udotea javensis (Montagne) A. et E. S. Gepp 1904: 363; Dawson 1954: 395, fig. 13b–c; Egerod 1975: 58, figs 28–29; Coppejans and Prud’homme van Reine 1989: 139, pl. 10 figs 3–9). Vouchers: UPF 1280, 1604 St. 4, 1275 St. 5; WSS 120 St. 8, 35 St. 11, 72 St. 12 Rhipilia tomentosa Kützing, 1858: 12, pl. 28I. Børgesen 1913: 92, fig. 76a–d; Bucher and Norris 1995: 2, figs 1–2; Millar and Kraft 2001: 25, figs 17, 18, 21–29 (= Rhipilia tenaculosa A et E. S. Gepp 1911: 56, figs 130–133; N’Yeurt 2001: 729, figs 51a–d, 70; Payri et al. 2002: 44, pl. 1 fig. 9). Vouchers: UPF 1200, 1491, WSS 31 St. 2; UPF 1530 St. 10, 1333, 1486 St. 9, 1376 St. 16, 1414, 1628 St. 22, 1442, 1506 St. 29, 1720 St. 30 Comments: after examining the relevant type material, Millar and Kraft (2001: 26) put R. tenaculosa in synonymy with R. tomentosa. This species is also found in nearby Fiji (N’Yeurt 2001). According to Millar and Kraft (2001), Fiji (178–180°E) appears to be the distributional limit eastward for the genus Rhipilia in the Pacific, but since Wallis Island (176° W) is slightly to the north-east of Fiji it would seem that the limit has been reset to the latter locality. However that distinction now belongs to the Phoenix Islands, (part of Kiribati), whose eastern limit is 170°30′ W. Indeed, South et al. (2001: 565) reported three species of Rhipilia from the Phoenix Islands (R. diaphana, R. geppii, and R. orientalis), but interestingly R. tomentosa is not listed from that locality, nor are any of the Phoenix species found in Wallis. Recent collections from Samoa (Skelton and South 2002) do not include any Rhipilia species; however other important nearby localities such as Tonga remain largely unknown phycologically. Rhipiliopsis howensis G. T. Kraft, 1986: 55, figs 22–30; 2000: 620, fig. 38F–G. N’Yeurt 2001: 729, fig. 77a–b. Voucher: WSS 50, St. 29 Comments: this is the second report of this species outside of its type locality of Lord Howe Island (Australia); it was also reported from the Suva Lagoon, Fiji (N’Yeurt 2001). Ochrophyta (= Heterokontophyta) The notion of a division of golden-brown algae has been the subject of much controversy since its inception by Luther (1899, ‘Heterokontae’, or having unequal flagella on gametes), as pointed out in de Reviers (2003: 148). van den Hoek et al. (1995) proposed to restrict the term ‘Heterokontophyta’ for brown algae with tripartite hairs on the longer flagella (Class Phaeophyceae), a definition contested by Cavalier-Smith (1995) who argued for the replacement term ‘Ochrophyta’ on the grounds that van den Hoek et al. (1995) did not apply the rules of the International Code of Botanical Nomenclature in their diagnosis. We use Ochrophyta here as adopted by Wynne (2001) and de Reviers (2003). Phaeophyceae Sphacelariales Sphacelariaceae Sphacelaria tribuloides Meneghini, 1840: 2. Littler and Littler 1997: 71, fig. 89; Draisma et al. 1998: 189, fig. 22; Abbott and Huisman 2004: 190, fig. 72D–E. Voucher: WSS 183, St. 40 Dictyotales Dictyotaceae Dictyopteris repens (Okamura) Børgesen, 1924: 265, fig. 13. Allender and Kraft 1983: 107, figs 19A–B; Coppejans et al. 1995b: 178, figs 3–4; N’Yeurt 1996: 399, figs 91–92; Payri et al. 2000: 128; Littler and Littler 2003: 166; Abbott and Huisman 2004: 197, fig. 75E–F. Vouchers: WSS 176 St. 1; UPF 1473 St. 5, 1625 St. 22, 1430, 1431 St. 23, 1526 St. 41 Dictyota bartayresiana Lamouroux, 1809a: 43. Allender and Kraft 1983: 112, figs 21E–F, 22D, 23A (‘D. bartayresii’); Ajisaka and Enomoto 1985: 38, fig. 1K–L; Coppejans et al. 1995b: 180, fig. 8; De Clerck and Coppejans 1997: 416, fig. 9; Payri et al. 2000: 132; De Clerck et al. 2001; Payri et al. 2002: 48, pl. 3 fig. 1; Abbott and Huisman 2004: 202, fig. 77A. Vouchers: UPF 1253, 1254, 1640, 1643 St. 7, 1345 St. 8, 1435 St. 25 Comments: this species forms ball-like hemispherical tufts, a distinctive feature of this species (Abbott and Huisman 2004). Dictyota divaricata Lamouroux, 1809a: 43. Allender and Kraft 1983: 112, figs 21C–D, 22B–C; Ajisaka and Enomoto 1985: 37, fig. 1H–I; Coppejans et al. 1995b: 182, fig. 19; Payri et al. 2000: 134. Vouchers: UPF 1212 St. 3, 1327, 1578 St. 9, 1614 St. 16 Marine plants from Wallis Dictyota friabilis Setchell, 1926: 91, pl. 13 figs 4–7. pl. 20 fig. 1; Dawson 1954: 401, fig. 16a–b; Ajisaka and Enomoto 1985: 37, fig. 1J; N’Yeurt 1996: 400, figs 93–94, 100–101; Payri et al. 2000: 132; N’Yeurt 2001: 742; Littler and Littler 2003: 168; Abbott and Huisman 2004: 205, fig. 77E. Vouchers: UPF 1548, 1660 St. 1, 1681, 1698 St. 3; WSS 159 St. 7 Lobophora variegata (Lamouroux) Womersley ex Oliveira, 1977: 217. Allender and Kraft 1983: 81, figs 4G–H, 5A–B; N’Yeurt 1996: 401, figs 105–106; Littler and Littler 1997: 79, fig. 99a–b; Payri et al. 2000: 136; Payri et al. 2002: 48, pl. 3 fig. 2; Skelton and South 2002a: 158, fig. 18C–D, G; Littler and Littler 2003: 172; Abbott and Huisman 2004: 209, fig. 80A–C. Vouchers: UPF 1549, 1651 St. 1, 1192, 1688 St. 3, 1597 St. 4, 1318 St. 10, 1318, 1528 St. 10, 1326 St. 9, 1447 St. 30, 1516 St. 41 Padina boryana Thivy in W. R. Taylor, 1966: 355, fig. 2. Payri et al. 2000: 138; Abbott and Huisman 2004: 213, fig. 81D–F (= Padina tenuis Bory 1827: 590; Egerod 1974: 150, fig. 84; Allender and Kraft 1983: 83, figs 5D–E, 6A; N’Yeurt 1996: 402, fig. 108f). Vouchers: UPF 1255, 1256, 1644, WSS 151 St. 7; UPF 1341, 1342, 1343, 1344 St. 8, 1409 St. 37, 1672 St. 43 Padina melemele Abbott et Magruder in Abbott, 1996: 143, figs 1–3. Coppejans et al. 1995b: 184, fig. 24 (‘Padina sp.’); Wynne 1998: 287; Coppejans et al. 2001: 29; Payri et al. 2002: 48, pl. 3 fig. 3; Littler and Littler 2003: 174; Abbott and Huisman 2004: 217, fig. 83A. Voucher: UPF 1384 St. 16 Comments: this deepwater species of Padina is characterised by a bright yellow to orange superior surface (surface towards which margin is curled; see Wynne 1998: 273) and a bright white calcified inferior surface. Originally described from Hawaii, it is now reported from Papua New Guinea (Coppejans et al. 2001), Fiji (N’Yeurt unpubl.; Littler and Littler 2003), New Caledonia (Payri unpubl.), Rapa Island, Austral Group, French Polynesia (ADR N’Yeurt and CE Payri, unpubl. data) and Wallis Island (this study). Padina melemele may be quite widespread in the tropical Pacific, but undercollected because of its inconspicuous subtidal habitat. Sargassum polycystum C. Agardh, 1824: 304. Chiang et al. 1992: 36, figs 1–12; N’Yeurt 1996: 403, figs 97, 108a–c; N’Yeurt 2001: 745; Payri et al. 2002: 48, pl. 3 fig. 4; Skelton and South 2002a: 160, fig. 21; Littler and Littler 2003: 184. Vouchers: UPF 1346, 1347 St. 8, 1460 St. 34 Comments: the characteristic knob-like proliferous outgrowths (elevated cryptostomata) on the distinct main axis, and large number of small vesicles on mature plants, are diagnostic for this species (Chiang et al. 1992). The only species of Sargassum and the largest algae found in Wallis, plants growing in the deep waters of Mata-Utu harbour attained up to 2 m in length. Its abundance at a port of entry Australian Systematic Botany 377 raises questions as to whether this represents an introduced species. From the literature, it is also present (around Wallis) in Fiji, New Caledonia, Samoa and Tonga. Fucales Sargassaceae Turbinaria conoides (J. Agardh) Kützing, 1860: 24, pl. 66 figs IIe, f. Barton 1891: 217, pl. LIV figs 1, 7–10, pl. LV figs 1–6, 8–10; Coppejans et al. 1995b: 192, fig. 36; Payri et al. 2002: 48, pl. 3 fig. 5; Littler and Littler 2003: 184. Voucher: UPF 1461, St. 34 Comments: this species is mainly distinguished from the usually less rare T. ornata by the absence of inward-directed set of teeth on the apical crown of branchlets. Plants are also somewhat less coarse, with thinner crown margins, than T. ornata. The Wallis material, however, has a few inward-directed set of teeth on some thalli, but otherwise conforms to the current species concept of T. conoides. This was also observed for material of T. conoides from New Caledonia (CE Payri, unpubl. data). On-going molecular analyses of the genera in the Fucales should reveal more information on the phylogeny of this group, and whether T. conoides is a valid species, or simply an ecomorph of T. ornata. Turbinaria ornata (Turner) J. Agardh, 1848: 266. Barton 1891: 219, pl. LIV figs 11–12, pl. LV fig. 7; Dawson 1954: 405, fig. 21; Coppejans et al. 1995b: 192, fig. 38; N’Yeurt 1996: 405; Payri et al. 2000: 148; N’Yeurt 2001: 745; Wynne 2001: 365, fig. 32; Payri et al. 2002: 48, pl. 3 fig. 6; Skelton and South 2002a: 160, fig. 19E; Littler and Littler 2003: 186; Abbott and Huisman 2004: 242, fig. 95. Vouchers: UPF 1222 St. 1, 1252 St. 7, 1316 St. 10 Comments: this species was not very common in the stations studied. The apical crown of branches has a distinct row of inward-directed teeth, and the branches are generally thicker than T. conoides (see comments above for latter species). Rhodophyta Porphyridiales Porphyridiaceae Chroodactylon ornatum (C. Agardh) Basson, 1979: 67, pl. IX fig. 52. Abbott 1999: 42, fig. 1A. Voucher: WSS 140 St. 4 Stylonema alsidii (Zanardini) K. Drew, 1956: 72. Abbott 1999: 44, fig. 1B–C. Voucher: WSS 08 St. 5 Erythropeltidales Erythrotrichiaceae Erythrotrichia carnea (Dillwyn) J. Agardh, 1883: 15, pl. I figs 8–10. Abbott 1999: 45, fig. 1E; N’Yeurt 2001: 746, fig. 106. Voucher: WSS 10 St. 5 378 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Acrochaetiales Acrochaetiaceae Acrochaetium barbadense (Vickers) Børgesen, 1915: 45. Abbott 1999: 52, fig. 3B. Voucher: WSS 99 St. 16 Acrochaetium kurogii (Lee et Lindstrom) Lee et Lee, 1988: 128. Lee and Yoshida 1997: 186, fig. 10A–E. Voucher: WSS 52 St. 29 Comments: this species is epiphytic on the brown alga Dictyopteris repens. This minute, elegant plant has opposite branching from every axial cell, with stellate chloroplasts and a central pyrenoid. The visible parts of the thallus grow parallel to the host surface, in a characteristic brittle-star pattern. The Wallis material is in good agreement with the description of the type species, and illustrations in Lee and Yoshida (1997). This appears to be the first report of the species outside of Japan. Pterocladiella caerulescens (Kützing) Santelices et Hommersand, 1997: 118. Abbott 1999: 197, fig. 51E; Payri et al. 2000: 168. Vouchers: UPF 1645, WSS 155 St. 7 Pterocladiella caloglossoides (Howe) Santelices, 1998: 244. Abbott 1999: 198, fig. 52A–B; Littler and Littler 2003: 64. Voucher: UPF 1664 St. 1; WSS 133 St. 8; UPF 1574, WSS 101 St. 16; UPF 1704 St. 20, 1624 St. 22; WSS 232 St. 36 Gelidiellaceae Gelidiella acerosa (Forsskål) Feldmann et Hamel, 1934: 533. Price and Scott 1992: 25, fig. 4A–E; Abbott 1999: 202, fig. 53A–C; Payri et al. 2000: 170; N’Yeurt 2001: 760; Littler and Littler 2003: 56. Vouchers: UPF 1225, 1655 St. 1, 1472 St. 5, 1263, 1264, 1635, 1636 St. 7, 1370 St. 8, 1499, WSS 36 St. 11; UPF 1539, 1544 St. 12, 1391 St. 17, 1705 St. 20, 1514 St. 31 Nemaliales Gracilariales Galaxauraceae Gracilariaceae Actinotrichia fragilis (Forsskål) Børgesen, 1932: 6. Abbott 1999: 64, fig. 7A–C; Payri et al. 2000: 160; N’Yeurt 2001: 749, figs 109, 117; Payri et al. 2002: 50, pl. 4 fig. 4; Littler and Littler 2003: 62. Vouchers: UPF 1283, 1598 St. 4, 1276 St. 5, 1257, 1634 St. 7, 1302 St. 12, 1379 St. 16, 1620 St. 22, 1433 St. 25 Gracilaria textorii (Suringar) De Toni, 1895: 27. Yamamoto 1978: 123, pls 12–14; pl. 42 figs 5–7; pl. 43 figs 1–4; N’Yeurt 2001: 772, figs 156, 168; Payri et al. 2002: 50, pl. 4 fig. 11. Vouchers: UPF 1357, 1361 St. 8 Galaxaura fasciculata Kjellman, 1900: 53, pl. figs 1–9; pl. 20 fig. 14. Abbott 1999: 66, fig. 7D–E; Payri et al. 2000: 162; N’Yeurt 2001: 749, fig. 116; Payri et al. 2002: 50, pl. 4 fig. 10. Vouchers: UPF 1215, 1224 St. 1, 1282, 1285 St. 4, 1303 St. 12, 1378 St. 16, 1408 St. 20, 1432, 1744 St. 25 Galaxaura filamentosa Chou in W.R. Taylor, 1945: 139. Abbott 1999: 67, fig. 7F; Payri et al. 2000: 164; Littler and Littler 2003: 64. Vouchers: UPF 1717 St. 1, 1600 St. 4; WSS 123 St. 8 Comments: this species was growing amidst seagrass beds, in a situation similar to that reported for this flattened species on the neighbouring island of Rotuma (N’Yeurt 1996: 415). Gracilaria sp. Vouchers: UPF 1696, St. 23 Comments: this unusual, flattened species of Gracilaria could represent a new species; its definite identification awaits the discovery of fertile spermatangial plants. Bonnemaisoniales Bonnemaisoniaceae Galaxaura marginata (Ellis et Solander) Lamouroux, 1816: 264. Abbott 1999: 67, fig. 7G–H; Payri et al. 2000: 164; N’Yeurt 2001: 752, figs 118–122; Littler and Littler 2003: 64. Vouchers: UPF 1284, 1592 St. 4 Asparagopsis taxiformis (Delile) Trevisan, 1845: 45. Abbott 1999: 174, fig. 43A–D; Payri et al. 2000: 160; N’Yeurt 2001: 775, figs 141, 142; Skelton and South 2002a: 141, fig. 6G; Littler and Littler 2003: 68. Voucher: UPF 1415 St. 22 Galaxaura subverticillata Kjellman, 1900: 48, pl. 3 figs 12; pl. 20 fig. 17. Abbott 1999: 69, fig. 8F–H. Vouchers: WSS 89 St. 1, UPF 1561 St. 2, 1593 St. 4, 1564 St. 16 Comments: the record represents the minute Falkenbergia stage in the life cycle of the species; larger thalli of the other stage presumably occur at other seasons in Wallis. Comments: we follow the opinion of Abbott (1999) in retaining the species distinct rather than being synonomyzed with Galaxaura rugosa, since the Wallisian plants also have abundant assimilatory filaments throughout the thallus. Gelidiales Gelidiaceae Gelidium pusillum (Stackhouse) Le Jolis, 1863: 139. Abbott 1999: 194, fig. 50A; Payri et al. 2000: 168; N’Yeurt 2001: 760. Vouchers: UPF 1552 St. 1, 1477, WSS 14 St. 5 Cryptonemiales Halymeniaceae Cryptonemia umbraticola Dawson, 1959: 43, figs 21F, 22A. Abbott 1999: 136, fig. 32B; Payri et al. 2000: 174. Vouchers: UPF 1475, WSS 13 St. 5, UPF 1323, 1325, 1488 St. 9, 1311, 1312, 1313, 1314, 1536, 1537, WSS 74, 75 St. 12; UPF 1387 St. 16, 1463 St. 36 Comments: this species forms loose rosettes at the base of coral heads, 12 m depth in the lagoon. Marine plants from Wallis Cryptonemia yendoi Weber-van Bosse, 1921: 249, fig. 77. Abbott 1999: 138, fig. 32C–E; Masuda et al. 1999: 449, figs 1–7. Vouchers: UPF 1579, WSS 108 St. 9; UPF 1535, 1538, WSS 73, 76 St. 12, 42 St. 23; UPF 1508 St. 29 Comments: this species can be distinguished from C. umbraticola by its erect habit, with a prominent stipe. Kallymeniaceae Kallymenia sp. Voucher: UPF 1207 St. 2 Comments: the lack of fertile material and fragmentary nature of the specimen precludes specietal assignment of this plant at this stage. Peyssonneliaceae Peyssonnelia bornetii Boudouresque et Denizot, 1973; 1975: 27, figs 24–47. Marcot et al. 1976: 237, fig. V; Payri et al. 2000: 172; N’Yeurt 2001: 777, figs 138, 146. Vouchers: UPF 1570, WSS 94, 98 St. 16 Peyssonnelia inamoena Pilger, 1911: 311, figs 24, 25. Boudouresque et Denizot 1975: 58, figs 107–115; Abbott 1999: 156, fig. 38B; Payri et al. 2000: 172; N’Yeurt 2001: 778, figs 139, 140a–e, 143–144; Littler and Littler 2003: 82. Vouchers: UPF 1204, WSS 221 St. 2; UPF 1304 St. 12, 1627, WSS 146 St 17; UPF 1502 St. 29 Peyssonnelia sp. Vouchers: WSS 180; UPF 1701, WSS 200 St. 3; UPF 1334 St. 9, 1669, 1729 St. 23, 1745 St. 25, 1721 St. 30 Comments: this species is closest to Peyssonnelia conchicola Piccone et Grunow sensu Marcot et al. 1988: 266; however, in the species from Madagascar the unicellular rhizoids are not laterally inserted, and the thallus is only 3–6 layers thick compared to 7–8 in the Wallis material. Both plants have terminal cystoliths. Peyssonnelia sp. also has been reported from the Cook Islands (ADR N’Yeurt unpubl. data). Gelidiella maschrisiana Dawson, 1957a: 17, fig. 4B. Abbott 1999: 204, fig. 53G–H; Payri et al. 2000: 170. Vouchers: WSS 127 St. 8; UPF 1605 St. 4, 1741, WSS 218 St. 10 Corallinales Corallinaceae Amphiroa foliacea Lamouroux in Quoy et Gaimard, 1824: 628, pl. 93 figs 2, 3. Abbott 1999: 178, figs 44C, 45B; Payri et al. 2002: 52, pl. 5 fig. 1 (as Amphiroa crassa); Skelton and South 2002a: 140, fig. 4D; Littler and Littler 2003: 26. Voucher: UPF 1201 St. 2, 1380 St. 16 Comments: Abbott (1999: 178) states that there exists a high degree of variation in intergenicula shape and size in Amphiroa foliacea, which may encompass the species circumsciption of A. tribulus, A. misakiensis and other taxa. Amphiroa tribulus would be the earlier name, but a critical study of its type is needed before making taxonomic Australian Systematic Botany 379 decisions. In the field however, A. foliacea can be readily distinguished by its flat, coarse habit with axes 3–5 mm wide, as opposed to the fragile, thinner branches of A. tribulus. Amphiroa fragilissima (Linnaeus) Lamouroux, 1816: 298. Price and Scott 1992: 45, fig. 11A–B; Payri et al. 2000: 182; Skelton and South 2002a: 140, fig. 4E; Littler and Littler 2003: 26. Vouchers: UPF 1661 St. 1, 1324, 1608 St. 9, 1445, 1512 St. 29 Amphiroa tribulus (Ellis et Solander) Lamouroux ,1816: 302. Payri et al. 2000: 182; N’Yeurt 2001: 761, figs 131, 134–135; Payri et al. 2002: 52, pl. 5 fig. 2; Littler and Littler 2003: 26. Voucher: UPF 1436 St. 25 Hydrolithon farinosum (Lamouroux) Penrose et Chamberlain, 1993: 295–303, figs 1–19. Gordon et al. 1976: 255, pl. III figs 1–4 (as Fosliella farinosa (Lamouroux) Howe); Penrose in Womersley 1996: 260, fig. 118A–D; Payri et al. 2000: 188; Ringeltaube and Harvey 2000: 437, figs 12–13; Littler and Littler 2003: 30. Voucher: WSS 161 St. 1 Hydrolithon murakoshii Iryu et Matsuda, 1996: 528. Payri et al. 2000: 190; Littler and Littler 2003: 32. Vouchers: WS 65 St. 1, 284 St. 17 Hydrolithon onkodes (Heydrich) Penrose et Woelkerling, 1992: 83. Gordon et al. 1976: 266, pl. IX figs 1–4; Ballesteros and Afonso-Carrillo 1995: 209, fig. 12; Womersley 1996: 261, fig. 119A–D; Payri et al. 2000: 192; Ringeltaube and Harvey 2000: 438, figs 14–16; Payri et al. 2002: 52, pl. 5 fig. 7. Vouchers: WS 75 St. 1, 1 St. 3, 120 St. 5, 283 St. 17, 399 St. 29, 403 St. 30 Hydrolithon reinboldii (Weber-van Bosse et Foslie in Foslie) Foslie, 1909: 55. Gordon et al. 1976: 255, pl. III figs 5–6, pl. IV figs 1–3; Payri et al. 2000: 192; Ringeltaube and Harvey 2000: 439, figs 17–20; Payri et al. 2002: 52, pl. 5 fig. 10; Littler and Littler 2003: 32. Vouchers: WS 67, 70 St. 1, 161 St. 11, 269 St. 16, 288 St. 17, 297 St. 18 Hydrolithon rupestris (Foslie) Penrose, 1996: 265, fig. 121. Penrose in Womersley 1996: 265, fig. 121A–C; Payri et al. 2000: 194. Voucher: WS 119 St. 5 Jania adhaerens Lamouroux, 1816: 270. Price and Scott 1992: 48, fig. 12A–C; Abbott 1999: 187, fig. 48A; N’Yeurt 2001: 765; Skelton and South 2002a: 141, fig. 6D–F; Littler and Littler 2003: 32. Vouchers: UPF 1547, 1727 St. 1, 1682 St. 3, 1716 St. 15, 1520 St. 41 Lithophyllum flavescens Keats, 1997: 357, figs 23–39. Payri et al. 2000: 204. Voucher: WS 333 St. 23 Lithophyllum insipidum Adey, Townsend et Boykins, 1982: 44, fig. 29A–F. Keats 1997: 352, figs 1–22; Payri et al. 2000: 206. Vouchers: WS 3 St. 3, 282 St. 17 Lithophyllum kotschyanum Unger, 1858: 22, pl V fig. 15. Gordon et al. 1976: 267, pl. IX figs 5, 6, pl. X figs 1, 2; Adey 380 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany et al. 1982: 37, fig. 24A–F; Ballesteros and Afonso-Carrillo 1995: 207, fig. 2; Payri et al. 2000: 206; Littler and Littler 2003: 36. Vouchers: WS 124 St. 5, 264 St. 16, 294 St. 18 Lithophyllum pygmaeum (Heydrich) Heydrich, 1897b: 412. Ballesteros and Afonso-Carrillo 1995: 207, fig. 3 (as Lithophyllum tamiense); N’Yeurt 1996: 414, fig. 115 (as Lithophyllum tamiense); Ringeltaube and Harvey 2000: 436, figs 7–11 (as Lithophyllum tamiense); Littler and Littler 2003: 38. Vouchers: WS 118 St. 5, 444 St. 37, 452 St. 38 Comments: forming unattached balls on the lagoon floor, or attached to coral boulders in the lagoon. According to Silva et al. (1996: 251), L. pygmaeum includes Lithophyllum tamiense (Heydrich) (Verheij 1994: 103). Lithothamnion proliferum Foslie in Weber-van Bosse and Foslie 1904: 18, fig. 8, pl. I figs 17–20. Keats et al. 1996: 188, figs 1–36 (‘L. prolifer’); Ringeltaube and Harvey 2000: 445, figs 32, 33 (‘L. prolifer’); Payri et al. 2002: 52, pl. 5 fig. 11; Skelton and South 2002a: 141, fig. 5B; Littler and Littler 2003: 38. Voucher: WS 262 St. 16 Mastophora pacifica (Heydrich) Foslie, 1903: 25. Woelkerling in Womersley 1996: 248, figs 111A–E, 112A–E, 113A–B; Payri et al. 2000: 196; Ringeltaube and Harvey 2000: 441, figs 21, 22; Payri et al. 2002: 52, pl. 5 fig. 4; Littler and Littler 2003: 40. Voucher: WS 136 St. 4, 361 St. 24, 396 St. 29 Mesophyllum erubescens (Foslie) Lemoine, 1928: 252. Gordon et al. 1976: 252, pl. I figs 5–8; Ballesteros and Afonso-Carrillo 1995: 209, fig. 11; Payri et al. 2000: 208; Ringeltaube and Harvey 2000: 445, figs 34–37; Payri et al. 2002: 52, pl. 5 fig. 8; Littler and Littler 2003: 42. Vouchers: WS 158 St. 11, 397 St. 29 Mesophyllum mesomorphum (Foslie) Adey, 1970: 25. Gordon et al. 1976: 252, pl. II figs 1–3. Voucher: WS 295 St. 18 Neogoniolithon brassica-florida (Harvey) Setchell et Mason, 1943: 91. Penrose in Womersley 1996: 281, fig. 129A–D; Ringeltaube and Harvey 2000: 441, figs 23–27; Payri et al. 2002: 52, pl. 5 fig. 5 (as Neogoniolithon frutescens). Vouchers: WS 67 St. 1, 4 St. 3, 291, St. 17, 360 St. 24, 376 St. 25, 450 St. 38 Pneophyllum conicum (Dawson) Keats, Chamberlain et Baba, 1997: 264. Gordon et al. 1976: 259, pl. IV figs 6–8, pl. VI fig. 1 (as Neogoniolithon conicum (Dawson) Gordon, Masaki et Akioka); Adey et al. 1982: 13, figs 6–7 (as Paragoniolithon conicum (Dawson) Adey, Townsend et Boykins); Payri et al. 2000: 202; Payri et al. 2002: 52, pl. 5 fig. 6; Littler and Littler 2003: 48. Voucher: WS 6 St. 3 Comments: this common Indo-Pacific species with a tortuous taxonomic history is reported to overgrow and kill live corals (Keats et al. 1997). Sporolithon ptychoides Heydrich, 1897: 67–69. Keats and Chamberlain 1993: 542, figs 1–13; Ballesteros and Afonso-Carrillo 1995: 209, fig. 13; Payri et al. 2000: 211; Littler and Littler 2003: 52. Vouchers: WS 66 St. 1106 St. 7, 247 St. 8 Gigartinales Caulacanthaceae Caulacanthus ustulatus (Turner) Kützing, 1843: 395. Wynne 1995: 277, figs 13–15; Abbott 1999: 104, fig. 20D–F; Masuda et al. 1999: 451, figs 8–11. Vouchers: UPF 1694, WSS 195 St. 1, 26, 27 St. 4, 67 St. 10; UPF 1715 St. 15 Comments: this species is superficially similar to Hypnea spinella, Caulacanthus ustulatus is distinguished by its more flattened axes, and its polygonal to hexagonal cortical cells in surface view. Corynocystaceae Corynocystis prostrata Kraft, in Kraft et al. 1999: 26, figs 6, 45–60. N’Yeurt 2001: 797, figs 107, 108, 217; Skelton and South 2002a: 142, fig. 7C; Littler and Littler 2003: 70. Voucher: UPF 1210, 1211, 1606 St. 2, 1322 St. 9, 1382 St. 16, 1623, WSS 144 St. 22; UPF 1446 St. 29 Comments: now found throughout the South Pacific (including French Polynesia, pers. obs. and In Herb. UPF), this characteristic alga is in most case associated with an ascidian which forms a thick white layer on the inferior surface. The genus, previously listed in the Acrotylaceae, has recently been transferred to a family of its own based on molecular studies (see Saunders et al. 2004). Dumontiaceae Gibsmithia hawaiiensis Doty, 1963: 458, figs 1–17. Abbott 1999: 108, fig. 22D–G; Payri et al. 2000: 212; Payri et al. 2002: 50, pl. 4 fig. 8. Vouchers: UPF 1205 St. 2; WSS 24 St. 4; UPF 1487 St. 9, 1416 St. 22, 1438, 1439 St. 29, 1462 St. 36 Hypneaceae Hypnea cervicornis J. Agardh, 1851: 451. Abbott 1999: 113, fig. 24A. Voucher: UPF 1587, St. 8 Comments: Haroun and Prud’homme van Reine (1993) reduced H. cervicornis in synonymy with H. spinella (C. Agardh) Kützing, but we prefer to follow the conservative opinion of Abbott (1999) to distinguish entities that look relatively different in the field. Hypnea charoides Lamouroux, 1813: 132, pl. 10 figs 1–3. Weber-van Bosse 1928: 449, figs 188, 189; Yamagishi and Masuda 1997: 136, figs 1–9; 2000: 31, figs 10–15; South 2004: 136, fig. 2. Vouchers: UPF 1261, 1262 St. 7, 1366 St. 8 Marine plants from Wallis Hypnea pannosa J. Agardh, 1847: 14. Price and Scott 1992: 38, figs 8A–D, 9A; Payri et al. 2000: 222; non N’Yeurt 2001: 780; Skelton and South 2002a: 143, fig. 7D; Littler and Littler 2003: 76. Vouchers: WSS 182 St. 1; UPF 1676 St. 3, 1279 St. 4, 1268, 1269 St. 5, 1259, 1260 St. 7, 1367, 1586 St. 8, 1424, 1425, 1428, 1671 St. 23, 1434 St. 25 Comments: records of Hypnea pannosa from the Suva Lagoon, Fiji in N’Yeurt (2001) were relegated to the status of an undescribed species close to the Japanese Hypnea charoides-valentiae complex by South (2004: 135). The Wallis material is in good agreement with H. pannosa from the Great Barrier Reef (Price and Scott 1992). Hypnea saidana Holmes, 1896: 256, pl. 11 figs 3a, b. Wynne 1995: 276, fig. 16; Payri et al. 2000: 224. Vouchers: UPF 1658 St. 1, 1193, 1194, 1646, 1699 St. 3, 1478, WSS 15, 16 St. 5; UPF 1543 St. 12, 1565 St. 16, 1523 St. 41 Hypnea spinella (C. Agardh) Kützing, 1847a: 23. Price and Scott 1992: 40, figs 9B, 10A–F; Abbott 1999: 117, fig. 25B–E; Payri et al. 2000: 224; Littler and Littler 2003: 76. Vouchers: UPF 1647, WSS 194 St. 3; UPF 1274, 1468, WSS 05, 06 St. 5; UPF 1368, 1588 St. 8, 1573, WSS 99 St. 16; UPF 1732, WSS 39 St. 23; UPF 1484 St. 26; WSS 53 St. 29; UPF 1521 St. 41 Comments: this is the most common minute Hypnea species in the collections, ubiquitous in most habitats. Based on culture and molecular studies, Yamagishi and Masuda (2000) relegated the entity previously known as Hypnea cervicornis/spinella from Japan to the rank of a new species, H. flexicaulis. The Wallis material agrees with the concept of H. spinella in Price and Scott (1992) and Abbott (1999). Hypnea valentiae (Turner) Montagne, 1841: 161. Weber-van Bosse 1928: 452, text fig. 190; Børgesen 1934: 17; Littler and Littler 2003: 76. Vouchers: UPF 1637, WSS 149, 152 St. 7; UPF 1673 St. 43 Comments: Yamagishi and Masuda (1997: 140; 2000) consider H. valentiae from Japan and Vietnam (Dawson 1954: 436, fig. 47) as belonging to the H. charoidesvalentiae complex. The Wallis material agrees with the descriptions of H. valentiae in Weber-van Bosse (1928) and Børgesen (1934), and the two names are retained separate as a conservative measure until future studies on their taxonomic status. Lomentariaceae Lomentaria corallicola Børgesen, 1939: 113, figs 30–32. Price and Scott 1992: 63, fig. 18A–E; Payri et al. 2000: 236; N’Yeurt 2001: 811, figs 223, 224; Littler and Littler 2003: 106. Vouchers: UPF 1663, WSS 171 St. 1 Nemastomataceae Platoma cyclocolpum (Montagne) Schmitz, 1894: 627. Masuda and Guiry 1994: 194–201, figs 1–34; Huisman Australian Systematic Botany 381 1999; N’Yeurt 2001: 781, figs 169–173; Payri et al. 2002: 50, pl. 4 fig. 6. Vouchers: UPF 1440, WSS 45, 46, 47 St. 29 Comments: the largest and most spectacular fleshy red alga collected from Wallis, the material agrees well with the description of material from the Canary Islands by by Masuda and Guiry (1994) and from Australia by Huisman (1999). Predaea laciniosa Kraft, 1984: 11, figs 25–35. Payri et al. 2000: 214; Payri et al. 2002: 50, pl. 4 fig. 5; Littler and Littler 2003: 78. Vouchers: WSS 19 St. 5; UPF 1626, WSS 145 St. 22; UPF 1419, 1482 St. 23 Predaea weldii Kraft et Abbott, 1971: 194, figs 1–15. Kraft 1984: 15, figs 36–42; Abbott 1999: 153, fig. 37D–F; N’Yeurt 2001: 783, figs 183, 228; Littler and Littler 2003: 78. Vouchers: UPF 1417, 1418, WSS 198 St. 22 Solieriaceae Meristotheca procumbens Gabrielson et Kraft, 1984: 241, fig. 14A–D. N’Yeurt 1995: 248, figs 3–10; 1996: 416, figs 137, 142–147, 203–205; Payri et al. 2000: 220; N’Yeurt 2001: 793, figs 208, 209, 231; Littler and Littler 2003: 92. Vouchers: UPF 1223, 1719, WSS 204 St. 1, 102 St. 2; UPF 1203 St. 3; WSS 136 St. 4; UPF 1575 St. 9, 1613 St. 16 Comments: this edible species is relatively infrequent and cryptic in Wallis, presumably the reason why it is not consumed traditionally as in neighbouring islands such as Rotuma (N’Yeurt 1995, 1996) where it is locally abundant in intertidal habitats. Sarconema filiforme (Sonder) Kylin, 1932: 22. Børgesen 1932: 11, fig. 7; Papenfuss and Edelstein 1974; 32, figs 1–3, 13, 20–25; Payri et al. 2000: 220. Vouchers: UPF 1348, 1349, 1350, 1591, WSS 117 St. 8 Comments: this species was recorded growing amidst seagrass beds, with other algae. This distinctive species can be recognised by its lubricous, terete thalli with bifurcate tips. This is the fourth report for this predominantly Indian Ocean genus in the South Pacific. Papenfuss and Edelstein (1974: 41) mention having examined herbarium material from New South Wales, Australia, and Upolu, Western Samoa (the latter as Dicranema setaceum var. upolensis Grunow). It is also found seasonally in Tahiti, French Polynesia (Payri et al. 2000). Halymeniales Halymeniaceae Halymenia actinophysa Howe, 1911: 509, pl. 34. Abbott 1999: 143, fig. 34A. Vouchers: WSS 134 St. 8; UPF 1611 St. 16 Comments: the Wallis material is in good agreement with the description of the Type of the species by Howe (1911), especially the macelike subcortical cells with several blunt arms. 382 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Prionitis sp. Voucher: UPF 1413, St. 22. Comments: The material being sterile, indentification is not possible at this stage. specific Sebdeniaceae Sebdenia sp. Vouchers: WSS 177, 179 St. 1 (tetrasporic); UPF 1501, 1670 St. 23 Comments: the fragmentary nature of the material precludes the definite identification of this taxon at the present time, but it could represent a new species. It is close to the flattened Sebdenia sp. mentioned from Fiji in N’Yeurt (2001: 806). Rhodymeniales Champiaceae Champia compressa Harvey, 1838: 402. Millar 1990: 371, figs 30A–D; Masuda et al. 2001: 84, figs 14–17; Skelton and South 2002a: 143, fig. 7E, F; Littler and Littler 2003: 102. Vouchers: WSS 84, 181 St. 1, UPF 1691 St. 3; WSS 153 St. 7, 121, 130 St. 8; UPF 1531 St. 10, 1577, WSS 105 St. 9, 222, 239 St. 16; UPF 1723 St. 30 Champia parvula (C. Agardh) Harvey, 1853: 76. Millar 1990: 371, figs 29G, H; Price and Scott 1992: 55, fig. 14A–E; Abbott 1999: 218, fig. 60A–C; Payri et al. 2000: 238; N’Yeurt 2001: 808, fig. 238; Skelton and South 2002a: 143, fig. 8A–C. Vouchers: WSS 170 St. 1, 156 St. 7 Rhodymeniaceae Chamaebotrys boergesenii (Weber-van Bosse) Huisman, 1996: 106, figs 35–38, 40–42. Abbott 1999: 228, fig. 63E–G; Payri et al. 2000: 230; Masuda et al. 2001: 81, figs 1–6; N’Yeurt 2001: 812, figs 218, 235, 236; Littler and Littler 2003: 110. Voucher: UPF 1695, WSS 196 St. 3; UPF 1465, WSS 04 St. 5; UPF 1702 St. 9, 1496 St. 11, 1737, 1500, WSS 37 St. 23 Chrysymenia okamurae Yamada et Segawa, 1953: 110, fig. 3. Abbott 1999: 230, fig. 64C–D. Vouchers: UPF 1572, WSS 96 St. 16 Coelothrix irregularis (Harvey) Børgesen, 1920: 389. Price and Scott 1992: 60, fig. 17A–D; Abbott 1999: 233, fig. 65A–D; Payri et al. 2000: 230. Vouchers: UPF 1638, 1639, WSS 150 St. 7 Gelidiopsis intricata (C. Agardh) Vickers, 1905: 61. Price and Scott 1992: 51, fig. 13A–F; Abbott 1999: 221, fig. 61A; Payri et al. 2000: 232; Skelton and South 2002a: 144, fig. 8D–G. Vouchers: UPF 1493 St. 2, 1641, 1642, WSS 154 St. 7, UPF 1328, 1331, 1580 St. 9, 1310 St. 12, 1374, 1375, 1714 St. 15, 1388 St. 16, 1494 St. 40; WSS 34 St. 37 Gelidiopsis scoparia (Montagne et Millardet) De Toni, 1900: 410. Abbott 1999: 221, fig. 61B; Littler and Littler 2003: 106. Vouchers: UPF 1206 St. 2, 1690 St. 3, 1470 St. 5, 1497 St. 11, 1563, 1618 St. 16, 1739 St. 36, 1495 St. 40 Halichrysis coalescens (Farlow) R.E. Norris et Millar in R. Norris, 1991: 583, 585, 587, figs 7–13. Abbott 1999: 235, fig. 66A–C; N’Yeurt 2001: 812, figs 220, 221, 234; Littler and Littler 2003: 116. Vouchers: UPF 1469, 1480, WSS 18 St. 5; UPF 1738, WSS 211 St. 23 Leptofauchea anastomosans (Weber-van Bosse) R. Norris et Aken, 1985: 58, figs 10–12. Dawson 1956: 52, fig 49 (‘Rhodymenia anastomosans’). Vouchers: UPF 1571, WSS 95 St. 16; WSH 277, WSS 43, 212 St. 23; UPF 1443, 1509 St. 29 Comments: growing intricately associated with articulated coralline algae (Amphiroa spp.) to which the blades form numerous secondary attachments, this species also occurs in the Cook islands in similar habitats (ADR N’Yeurt, unpubl. data). Ceramiales Ceramiaceae Anotrichum tenue (C. Agardh) Nägeli, 1862: 399. Abbott 1999: 247, fig. 68D; Masuda et al. 1999: 453, figs 15–23; Payri et al. 2000: 240; Skelton and South 2002a: 145, fig. 9A–F. Voucher: WSS 67 St. 10 Comments: Womersley (1998: 344) discusses the range of characteristics in specimens of the A. tenue complex, which support the concept of a single species with several varieties sometimes described as separate species. Antithamnion decipiens (J. Agardh) Athanasiadis, 1996: 151. Abbott 1999: 250, fig. 69C–D. Voucher: WSS 40, St. 23 Balliella repens Huisman et Kraft, 1984: 77, figs 16–26. Price and Scott 1992: 72, figs 20D, 21A–B; Abbott 1999: 257, fig. 71H. Vouchers: WSS 140 St. 4, 69 St. 12, 49 St. 29 Centroceras minutum Yamada, 1944: 42. Dawson 1956: 54, fig. 54; Ardré 1987: 285, figs 30–37; Wynne 1993: 12, fig. 7; 1995: 290, fig. 27; Abbott 1999: 262, fig. 73H–I; N’Yeurt 2001: 817, figs 249, 250. Vouchers: WSS 228 St. 3, 68 St. 10 Comments: reasons to maintain Centroceras minutum as a separate species from C. clavulatum are given in Wynne (1993: 12). N’Yeurt (2001: 817) examined slide-mounted type material of C. minutum in SAP and found it also to differ from C. clavulatum. Ceramium codii (Richards) Mazoyer, 1938: 324. Millar 1990: 393, fig. 41D–F; Abbott 1999: 270, fig. 75D–F; Payri et al. 2000: 246; South and Skelton 2000: 56, figs 11–14. Vouchers: WSS 08 St. 5, 126 St. 8, 218 St. 10, 81, 82, St. 12 Ceramium flaccidum (Kützing) Ardissone, 1871: 40. Womersley 1978: 234, figs 4A–D, 14E–H; Price and Scott 1992: 89, fig. 27A–E; Womersley 1998: 410, figs 188E, H, 190A–D; Abbott 1999: 274, fig. 76D–H; Payri et al. 2000: 248; South and Skelton 2000: 65, figs 32–39, 41–44; Marine plants from Wallis Australian Systematic Botany 383 N’Yeurt 2001: 817, fig. 253. Vouchers: UPF 1683, 1697, WSS 188, 199, 228 St. 3; UPF 1585, 1713, WSS 114, 202 St. 8; UPF 1742, WSS 67, 217 St. 10, 48 St. 29 Diplothamnion jolyi van den Hoek 1978: 47; Huisman 1991; Stegenga and Vroman 1987: 411, figs 35, 36; Abbott 1999: 297, fig. 83C. Voucher: WSS 25, St. 4 Comments: the broad rows of basipetal cortical cells and forcipate apices are characterisctic for this species. This is the largest and most common Ceramium species in Wallis, and is locally abundant especially forming reddish, silky tufts on branches of dead coral. Euptilota articulata (J. Agardh) Schmitz, 1896: 7. Womersley 1998: 355, pl. 2 fig. 2, figs 141E, 164, 165; N’Yeurt 2001: 820, figs 254–263; Payri et al. 2002: 50, pl. 4 fig. 3; Littler and Littler 2003: 122. Vouchers: UPF 1214 St. 3, WSS 139 St. 4, 129 St. 8; UPF 1421, 1422, 1423, 1609, 1621 St. 22, 1449, 1722 St. 30 Ceramium kramerii South et Skelton, 2000: 69, figs 45–51. Voucher: WSS 126, St. 8 Comments: a minute, distinctive species, epiphytic on larger algae growing on coral debris. Ceramium macilentum J. Agardh, 1894: 15. Womersley 1978: 232, figs 3E, 14A–D; Price and Scott 1992: 100, fig. 31; South and Skelton 2000: 71, figs 52–62. Vouchers: WSS 194 St. 3, 08 St. 5, 208 St. 23, 48 St. 29 Ceramium sp. aff. C. marshallense Dawson, 1957b: 120, figs 27a, b. Price and Scott 1992: 102, fig. 32; Wynne 1993: 14, figs 8–9; 1995: 294, figs 38, 39 South and Skelton 2000: 75, figs 63–71, 73. Vouchers: WSS 25, 139 St. 4 Ceramium subdichotomum Weber-van Bosse, 1923: 333, fig. 125. Wynne 1995: 296, figs 44, 45; 1999: 195, figs 8, 9; Payri et al. 2000: 250 (‘subdichotum’); South and Skelton 2000: 78, figs 74–79. Voucher: WSS 208, St. 23 Ceramium upolense South et Skelton, 2000: 81, figs 80–88. Vouchers: WSS 07, 08 St. 5, 203 St. 9, 99 St. 16, 60, 64 St. 31 Corallophila apiculata (Yamada) R. Norris, 1993: 395. N’Yeurt 1996: 420, figs 148–151, 163 (‘Centroceras apiculatum’); Abbott 1999: 288, fig. 81A–C; Payri et al. 2000: 242. Vouchers: WSS 08 St. 5, 67 St. 10 Corallophila huysmansii (Weber-van Bosse) R. Norris, 1993: 396. Abbott 1999: 290, fig. 81D–E; South and Skelton 2002: 148, fig. 11A. Voucher: WSS 127 St. 8 Crouania minutissima Yamada, 1944: 40. Dawson 1956: 55, fig. 56; Abbott 1999: 294, fig. 82E–G. Vouchers: WSS 33 St. 2, 195 St. 3; UPF 1529, WSS 65 St. 10, 104 St. 9 Comments: the Wallis material has straight to slightly upcurved whorl branches with spherical to slightly elongate terminal cells. Cribb (1983) suggested that previous records of C. minutissima with three whorl-branchlets actually represent C. capricornica Saenger et Wollaston, as discussed in Skelton and South (2002: 149). On the other hand, Abbot (1999: 294) reported having examined authentic C. minutissima from Ant atoll in SAP, as well as recent material from that locality, and found three whorl branchlets in both cases. There hence would seem to be an error in Yamada’s original diagnosis, and C. minutissima is retained here as a conservative measure. Comments: this elegant, plumose material agrees well with that described by Itono (1977: 139) from Southern Japan. Haloplegma duperreyi Montagne, 1842b: 258, pl. 7 fig. 1. Price and Scott 1992: 127, fig. 43A–D; Womersley 1998: 282, fig. 133A–E; Abbott 1999: 305, fig. 86E; Littler and Littler 2003: 122. Vouchers: UPF 1492, WSS 32 St. 2; UPF 1213 St. 3, WSS 141 St. 4; UPF 1315 St. 10, 1412 St. 22, 1448, 1724 St. 30 Ptilothamnion sp. Voucher: WSS 128 St. 8. Comments: The material is sterile and specific identification is not possible. Spyridia filamentosa (Wulfen) Harvey in W. Hooker, 1833: 337. Womersley and Cartledge 1975: 222, figs 1, 3A–B; Price and Scott 1992: 131, fig. 45A–E; Womersley 1998: 372, figs 171, 173A–B; Abbott 1999: 313, fig. 88A–B; Payri et al. 2000: 256; Littler and Littler 2003: 124. Vouchers: UPF 1358, 1359, 1584, WSS 113 St. 8 Comments: frowing as unattached clumps within seagrass beds. Wrangelia argus (Montagne) Montagne, 1856: 444. Price and Scott 1992: 134, fig. 46A–E; N’Yeurt 1996: 424, figs 167, 169–172. Vouchers: WSS 86 St. 1, 192 St. 3 Wrangelia dumontii (Dawson) Abbott, 1979: 222, figs 21–26, 29–30; 1999: 316, fig. 89A–B. Littler and Littler 2003: 126. Voucher: WSS 139 St. 4 Comments: Wrangelia dumontii is readily distinguished from W. argus by its obtuse branch tips, contrasted to the needle-like tips of the former species. Dasyaceae Dasya anastomosans (Weber-van Bosse) M.J. Wynne, 2002: 539. Millar 1990: 433, fig. 59D–G (‘Dasya pilosa’); Abbott 1999: 325, fig. 93A–E (‘Dasya pilosa’); Payri et al. 2000: 264 (‘Dasya pilosa’); Littler and Littler 2003: 128 (‘Dasya pilosa’). Vouchers: UPF 1665, WSS 173 St. 1 Comments: the tortuous taxonomy of this species is discussed in Wynne (2002). The Wallis plants have the typical furry habit with long assimilatory filaments. Weber-van Bosse (1921: 309, pl. 7 fig. 10) only gave a Latin diagnosis for her new species, with a fuller description given in Weber-van Bosse (1923: 376). 384 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Dasya baillouviana (S. Gmelin) Montagne, 1841: 165. Littler and Littler 1997: 55, fig. 62; 2000: 170, fig. middle 171; Payri et al. 2000: 260. Vouchers: UPF 1534, WSS 71 St. 12 Eupogodon geppii (Weber-van Bosse) P. Silva in P. Silva, Meñez et Moe, 1987: 130 (misapplied name: Exophyllum wentii Weber-van Bosse (1928: 478, pl. VI figs 5, 6): Hollenberg 1968c: 81, figs 6, 7; Abbott 1999: 363, fig. 105D-H; Payri et al. 2000: 278; Skelton and South 2002a: 155, fig. 16C). Vouchers: UPF 1652, WSS 163, 164, 165 St. 1 Comments: Dasyopsis geppii was first described from the Chagos Archipelago and Seychelles by Weber-van Bosse (1913: 130, pl. 13 figs 18–20; pl. 14 fig. 33), together with Dasyopsis palmatifida Weber-van Bosse (1913: 130, pl. 13 fig. 21). Later, Weber-van Bosse (1923: 379) was of the opinion that both species could be conspecific, a view echoed by Dawson (1957b: 124) and confirmed by de De Jong et al. (1997: 445). Both entities were earlier transferred to the genus Eupogodon Kützing (1845: 312) by Silva et al. (1987: 130). More recently, Millar (1996: 156) and De Jong et al. (1997: 445) alluded to the possibility that Eupogodon geppi (Weber-van Bosse) P. C. Silva and E. palmatifidus (Weber-van Bosse) PC Silva formed a genus of their own. The first author has examined type material of Dasyopsis palmatifida (L 938.7–422) and it matches well the Wallis plants, but its affinities to the genus Eupogodon is not immediately apparent. Some Pacific records were misidentified in the past as Exophyllum wentii Weber-van Bosse; the first author has examined the dried Lectotype (L 0061135) and the liquid-preserved Syntype (L 0109993) and Isotype (L 0109994) of the latter species, and they do not match at all the Wallis, Hawaiian, Fijian or Samoan plants in vegetative or reproductive anatomy. Ongoing molecular analyses (N’Yeurt and PW Gabrielson, unpubl. data) should shed more light on the taxonomy and generic affinities of this unusual plant in the near future. Heterosiphonia crispella (C. Agardh) Wynne var. laxa (Børgesen) Wynne, 1985: 87. Price and Scott 1992: 161, fig. 56A–D; Lluch and Garreta 1993: 468, fig. 4; N’Yeurt 1996: 425, figs 175, 181; Payri et al. 2000: 266; Littler and Littler 2003: 130; Yamagishi et al. 2003: 540, figs 14–23. Vouchers: WSS 176 St. 1, 214 St. 23 Delesseriaceae Hypoglossum simulans Wynne, I. Price et Ballantine, 1989: 31, figs 12–26. Wynne 1989: 515, figs 1H, 2A–E; Abbott 1999: 340, fig. 98A–B; Payri et al. 2000: 258; N’Yeurt 2001: 832, figs 281, 282, 286; Skelton and South 2002a: 152, fig. 12C–D; Littler and Littler 2003: 134. Voucher: WSS 233 St. 35 Martensia fragilis Harvey, 1854: 145. Millar 1990: 418, figs 53C–E; Yoshida and Mikami 1996: 101, figs 1, 4–18; Abbott 1999: 344, fig. 99C–E; Payri et al. 2002: 50, pl. 4 fig. 12; Skelton and South 2002a: 153, fig. 14A–D; Littler and Littler 2003: 36. Vouchers: WSS 177 St 1; UPF 1209 St. 3; WSS 11, 12 St. 4; UPF 1383, 1569, 1607, WSS 93 St. 16; UPF 1725 St. 23, 1441, 1505 St. 29; WSS 63 St. 31; UPF 1525 St. 41 Comments: Martensia fragilis can be distinguished from superficially similar M. elegans by the alternate production of membranous and reticulate portions of the thallus, whereas in M. elegans reticulate portions never produce further membranous parts (Millar 1990). This elegant alga is quite common in subtidal habitats surveyed in Wallis. Nitophyllum adhaerens Wynne, 1997: 215, figs 1–14. Abbott 1999: 347, fig. 100E. Vouchers: WSS 177 St. 1, 191 St. 3; UPF 1576, WSS 103 St. 9; UPF 1668 St 18; WSS 51 St. 29 Rhodomelaceae Acanthophora pacifica (Setchell) G.T. Kraft, 1979: 128, figs 1–6, 18–24. Abbott 1999: 353, fig. 102A–C; De Jong et al. 1999: 229, fig. 43; Payri et al. 2000: 268; Payri et al. 2002: 50, pl. 4 fig. 1; N’Yeurt 2001: 33, figs 293–299; Littler and Littler 2003: 142. Vouchers: UPF 1420, 1622 St. 22, 1503 St. 29 Acanthophora spicifera (Vahl) Børgesen, 1910: 201, figs 18A–C; 19A-E. Abbott 1999: 355, fig. 102D–E; De Jong et al. 1999: 31, figs 3, 33–39, 46; Payri et al. 2000: 270; Skelton and South 2002a: 153, figs 16A, 17A; Littler and Littler 2003: 142. Vouchers: UPF 1352, 1353 St. 8 Comments: an often overlooked edible species, growing on the reef crest and seagrass beds. Bostrychia tenella (Lamouroux) J. Agardh, 1863 (1851– 1863): 869. Falkenberg 1901: 515, pl. 12 figs 10–13; King and Puttock 1989: 34, fig. 15A–D; N’Yeurt 1996: 429, figs 129, 186; Littler and Littler 1997: 59, fig. 68; Payri et al. 2002: 50, pl. 4 fig. 2. Voucher: UPF 1410 St. 38 Comments: found as spongy, sediment-infiltrated mats at the base of mangrove roots and on beach rock at a marina on the south coast of the island. This species grows together with Laurencia brachyclados. It is also found growing on mangrove roots on nearby Rotuma Island (N’Yeurt 1996). Chondria arcuata Hollenberg, 1945: 447, figs 2–4. Gordon-Mills and Womersley 1987: 551, fig 24D–E, 27; Abbott 1999: 57, fig. 103A; Womersley 2003: 427, fig. 186A–E. Voucher: WSS 236, St. 35 Chondria armata (Kützing) Okamura, 1907: 69, pl. XVI figs 9–19. Millar 1990: 459, figs 71A–B; Ballesteros 1994: 538, fig. 1a–c; Payri et al. 2002: 50, pl. 4 fig. 7; Littler and Marine plants from Wallis Littler 2003: 144. Vouchers: UPF 1265, 1266, 1301 St. 7, 1542, WSS 79, 80, 81 St. 12 Comments: this distinctive species of Chondria is also found in neighbouring Fiji (Garbary et al. 1991). Chondria dangeardii Dawson, 1954: 60, fig. 62f–g. Price and Scott 1992: 165, fig. 57A–D; Ballesteros 1994: 538 fig. 2a–c; Abbott 1999: 359, fig. 103B–C; Payri et al. 2000: 273; Skelton and South 2002a: 155, fig. 17B. Vouchers: UPF 1718, 1659, WSS 167, 168, 169 St. 1 Chondria dasyphylla (Woodward) C. Agardh, 1817: XVIII. Gordon-Mills and Womersley 1987: 246, figs 30–57; N’Yeurt 1996: 430, figs 134, 188; Payri et al. 2000: 74. Vouchers: UPF 1364, 1589 St. 8 Chondria leptacremon (Melvill) De Toni, 1903: 848. Littler and Littler 2000: 202. Vouchers: UPF 1188, 1677, 1693, 1700, WSS 184, 185, 193 St. 3; UPF 1649, WSS 160, 161 St. 7; UPF 1406, WSS 225 St. 19 Chondria simpliciuscula Weber-van Bosse, 1913: 125, pl. 12 figs 9, 10. Price and Scott 1992: 169, fig. 59A–E; N’Yeurt 1996: 431, figs 130, 133, 190; Abbott 1999: 361 fig. 104A–F; Payri et al. 2000: 276. Vouchers: WSS 06, 10 St. 5, 60 St. 31 Herposiphonia delicatula Hollenberg, 1968b: 540, figs 1A,B, 2H, 3. Wynne 1995: 303, fig. 56; Abbott 1999: 371, fig. 107F-G. Voucher: WSS 170 St. 1 Herposiphonia obscura Hollenberg, 1968b: 549, fig. 25. Abbott 1999: 73, fig. 108B. Voucher: WSS 67 St. 10 Australian Systematic Botany 385 842, figs 309–312, 318); Littler and Littler 2003: 148. Vouchers: UPF 1662, WSS 172 St. 1, 135 St. 8; UPF 1612 St. 16 Comments: this species can be readily distinguished by its markedly projecting outer cortical cells. Chondrophycus succica (Cribb) Nam, 1999: 463. (= Laurencia succica Cribb 1958a: 163, pl. 1 figs 1–3; Price and Scott 1992: 192, fig. 69A–D; N’Yeurt 2001: 845, figs 314, 316, 317; Payri et al. 2000: 280). Vouchers: WSS 175 St. 1; UPF 1678 St. 3, 1594, WSS 118 St. 4; UPF 1467, WSS 03 St. 5, 119 St. 8; UPF 1386, WSS 239 St. 16; UPF 1667, 1733, 1734, WSS 41 St. 23; UPF 1444, 1507 St. 29 Laurencia brachyclados Pilger, 1920: 6, figs 9, 10. Dawson 1954: 458, fig. 61; McDermid 1988: 232, figs 1, 2; Abbott 1999: 382, fig. 111A–C Vouchers: UPF 1411, WSS 207 St. 38 Comments: growing mixed with Bostrychia tenella at the base of mangrove roots. Laurencia cervicornis Harvey, 1853: 73, pl. 18C. Littler and Littler 2000: 212. Vouchers: UPF 1735, WSS 209 St. 23 Laurencia flexilis Setchell, 1926: 101, pl. 19. Masuda et al. 1999: 455, figs 29–34; Payri et al. 2000: 280; Littler and Littler 2003: 52. Vouchers: UPF 1629, 1630, WSS 148 St. 7; UPF 1483 St. 26 Chondrophycus cartilaginea (Yamada) Nam, 2004: 267. (= Laurencia cartilaginea Yamada 1931: 230, pl. 19 figs a, o; McDermid 1988: 232, fig. 3; Nam and Saito 1990: 08, figs 1–7; Abbott 1999: 384, fig. 111D–E). Vouchers: WSS 174, St. 1; UPF 1675, WSS 186 St. 3; UPF 1355, 1356, 1590, WSS 115, 116 St. 8; UPF 1666, 1736, WSS 210 St. 23 Comments: originally described from Tahiti (Setchell 1926), this species has since been widely reported in the Pacific and Indian Ocean (Silva et al. 1996). It could be confused with Laurencia venusta Yamada, but the latter has both longitudinally oriented secondary pit connection between superficial cortical cells and lenticular thickenings in medullary cells, which are not found in L. flexilis (Masuda et al. 1999). Comments: Nam and Saito (1999) and Nam (2004) discuss the Laurencia complex and its relationship to Chondrophycus. Laurencia implicata J. Agardh, 1852: 745. Price and Scott 1992: 185, figs 65A–E, 66A–C. Vouchers: UPF 1712, WSS 201 St. 17; UPF 1458, 1515, WSS 56 St. 31 Chondrophycus dotyi (Saito) Nam, 1999: 463. (= Laurencia dotyi Saito 1969: 154, figs 9, 10; McDermid 1988: 233, figs 7, 8; Abbott 1999: 385, fig. 112A–B; Payri et al. 2000: 284; Payri et al. 2002: 50, pl. 4 fig. 9). Vouchers: UPF 1190, 1208 St. 3; WSS 143 St. 4; UPF 1270, 1271, 1466, WSS 02 St. 5, 131, 132 St. 8; UPF 1381, 1610, 1619 St. 16, 1427 St. 23, 1504 St. 29, 1524 St. 41 Laurencia mariannensis Yamada, 1931: 200, pl. 5 fig. b, text figs f, g. Abbott 1999: 388, fig. 113A–B. Vouchers: UPF 1485, WSS 23 St. 4 Comments: Garbary and Harper (1998) delineated the characteristics that separate the genus Chondrophycus from Laurencia and Osmundea. Further new combinations were given by Nam (1999: 463). Chondrophycus parvipapillata (C. K. Tseng) Garbary et Harper, 1998: 195. (= Laurencia parvipapillata Tseng 1943: 204, pl. IV; Abbott 1999: 391, fig. 114A–B; N’Yeurt 2001: Laurencia cf. L. sp. in Price and Scott, 1992: 194, fig. 70A–C. Vouchers: UPF 1567, WSS 91 St. 16 Comments: this species is similar to the minute Laurencia reported from the Great Barrier Reef by Price and Scott (1992), and could well represent a new species. Further studies are required to elucidate its affinities. Polysiphonia apiculata Hollenberg, 1968a: 61, figs 1D, 8, 9. Abbott 1999: 411, fig. 120C. Vouchers: WSS 22 St. 2; UPF 1474, WSS 09 St. 5, 127, 128 St. 8, UPF 1581, WSS 109 St. 9 386 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany Comments: this species is readily distinguished by its abruptly tapering apices with a prominent apical cell. Polysiphonia delicatula Hollenberg, 1968a: 62, fig. 1F. Abbott 1999: 412, fig. 120E. Vouchers: UPF 1687, WSS 190 St. 3, 25 St. 4, 07, 08 St. 5, 60, 64 St. 31 Polysiphonia homoia Setchell et Gardner, 1930: 162. Abbott 1999: 417, fig. 122F–G. Voucher: WSS 67 St. 10 Polysiphonia poko Hollenberg, 1968a: 70, figs 3A, 15. Price and Scott 1992: 209, fig. 76A–C; Abbott 1999: 421, fig. 123D-G. Voucher: WSS 233, 237 St. 35 Polysiphonia scopulorum Harvey, 1855: 540. Stegenga and Vroman 1988: 307, figs 21–26, 29–31; Millar 1990: 445, fig. 65E–G; Price and Scott 1992: 210, fig. 77A–D; Payri et al. 2000: 286; N’Yeurt 2001: 37, figs 304, 308; Payri et al. 2000: 286; Womersley 2003: 175, fig. 78A–E. Voucher: WSS 205 St. 30 Polysiphonia sphaerocarpa Børgesen, 1918: 271, figs 267–271. Stegenga and Vroman 1988: 307, figs 27, 28; Millar 1990: 446, fig. 66A–E; Price and Scott 1992: 212, fig. 78A–D; Schneider and Searles 1997: 204, fig. 19; Abbott 1999: 28, fig. 127A–G. Voucher: WSS 219, 220 St. 10 Comments: this species is easily recognised by its small, erect tufted habit and large, adaxially curved globular cystocarps regularly placed on the main axes. Kim and Lee (1999: 280) transferred P. sphaerocarpa to their new genus Neosiphonia based on reproductive and vegetative differences with Polysiphonia, but we concur with Skelton and South (2002: 156) in temporarily maintaining the South Pacific material under Polysiphonia pending a better understanding and verification of relevant type material of regional species. Tolypiocladia glomerulata (C. Agardh) Schmitz in Schmitz and Falkenberg, 1897: 441. Falkenberg 1901: 177, pl. 21 figs 27–29; Dawson 1954: 452, fig. 59b–c; Price and Scott 1992: 219, fig. 81A–D; Wynne 1995: 321, fig. 80; Abbott 1999: 442, fig. 132F–H; N’Yeurt 2001: 837, figs 302, 303; Littler and Littler 2003: 158. Vouchers: WSS 157, St. 7; UPF 1730, WSS 208 St. 23 Incertae sedis Wurdemannia miniata (Sprengel) Feldmann et Hamel, 1934: 544. Hatta and Prud’homme van Reine 1991: 375, fig. 14a–f; Price and Scott 1992: 31, fig. 6A–E; Abbott 1999: 238. Vouchers: UPF 1556 St. 1, 1599 St. 4; WSS 122 St. 8, UPF 1540, WSS 77 St. 12; UPF 1706 St. 20; WSS 44 St. 23; UPF 1510 St. 29 Magnoliophyta (seagrasses) Hydrocharitales Hydrocharitaceae Halophila ovalis (R. Brown) J.D. Hooker, 1858: 45. Dawson 1954: 376, fig. 1d–f; Meñez et al. 1983: 30, figs 21A–C, 22; Lanyon 1986: 27, fig. 8, pl. 1 fig. f, pl. 3 fig. f; Payri et al. 2000: 96; Payri et al. 2002: 53, pl. 6 fig. 2; Littler and Littler 2003: 186. Vouchers: UPF 1197, 1198, 1199 St. 3, 1227 St. 6, 1246 St. 7, 1369 St. 8, 1459 St. 34 Comments: the smallest seagrass characteristic translucent oval leaves. in Wallis, with Potamogetonales Cymodoceaceae Halodule pinifolia (Miki) den Hartog, 1964: 309. Meñez et al. 1983: 13, figs 7A–B, 8; Lanyon 1986: 20, pl. 1 fig. d, pl. 3 fig. d; Payri et al. 2002: 53, pl. 6 fig. 3; Littler and Littler 2003: 288. Voucher: UPF 1371, WSS 245 St. 8 Comments: the Wallis material has irregularly serrate margins and furcate midribs at the apices of leaves, which are diagnostic features of the species. The blades are grass-like, flattened. Syringodium isoetifolium (Ascherson) Dandy in Dandy and Tandy, 1939: 116. Meñez et al. 1983: 18, figs 11A–D, 12; Lanyon 1986: 35, fig. 12, pl. 2 fig. c, pl. 4 fig. c; Payri et al. 2002: 53, pl. 6 fig. 4; Littler and Littler 2003: 288. Vouchers: UPF 1228, 1229, 1247, 1248 St. 7, 1372 St. 8 Comments: the fleshy, terete blades of this species easily distinguishes it in the field from Halodule pinifolia. Discussion Taxonomy and algal distribution patterns This study has yielded a total of 194 species of macrophytes, including 14 Cyanobacteria, 11 Heterokontophyta, 41 Chlorophyta, 128 Rhodophyta and three species of seagrasses. The non-geniculate Corallinaceae are not fully treated, and numerous species remain to be added in future. This study represents the first published records of seagrasses and marine algae for the Wallis islands, increasing our knowledge of the tropical marine flora of the western Pacific. Even though the outer part of the reef has not been sampled in the eastern part of the island, most of the habitats have been examined in the other areas from the surface to a depth of 25 m, both inside and outside of the lagoon. With respect to the distribution of species among the 42 different stations (Table 2), the highest species richness (130 species) is in the area of the fringing reef, owing to the presence of the large seagrass beds in the sheltered, shallow and soft-bottomed areas which provide microhabitats for various algae. The canopy of the seagrass beds comprises the large Syringodium isoetifolium, and the dominant algae are common green species such as Halimeda macroloba, H. incrassata, H. opuntia, Caulerpa spp. as well as the diminutive seagrass Halophila ovalis. A third species of seagrass Halodule pinifolia was less common and found at only one site. The fringing shallow reef flats were dominated by brown algae such as Padina boryana, Dictyota Marine plants from Wallis Australian Systematic Botany 387 Table 3. Taxa comparison between Wallis and several tropical Pacific areas (excluding Cyanobacteria and coralline algae) Fiji and Rotuma are treated here as a single entity, since all species from Rotuma in common with Wallis also occur in Fiji Locality Wallis (this study) Fiji and Rotuma (N’Yeurt et al. 1996b; Littler and Littler 2003; South and Skelton 2003) French Polynesia (Payri and N’Yeurt 1997; Payri et al. 2000; Littler and Littler 2003) Solomon Islands (Womersley and Bailey 1970; Littler and Littler 2003) Rarotonga, Cook Islands (Chapman 1977; ADR N’Yeurt unpubl. data) Samoa (including American Samoa) (Skelton and South 1999, 2002a, 2002b; Littler and Littler 2003) bartayresiana, Turbinaria conoides and Sargassum polycystum. Moroever, 30% of the species sampled in the fringing reefs were only found in this particular area (e.g. Acanthophora spicifera, Coelothrix irregularis, Gracilaria textorii, Hypnea cervicornis, Sarconema filiforme, Pterocladia caerulescens, Spyridia filamentosa, Hypnea charoides, Anadyomene tropicus, Siphonocladus tropicus, Borgesenia forbesii, Chaetomorpha linum, Valonia aegagropila, Sargassum polycystum and Turbinaria conoides). The least species rich of all sites are the pinnacles with only 24 species, none of which is restricted to this zone. Seventy-one per cent of the pinnacle flora also occur on the fringing reef, which is undoubtedly due to the very similar geomorphological features of the reef flats between the two areas, and the influence of the eroding land mass (terrigenous inputs leading to sedimentation, water turbidity). Unlike the barrier reefs, most of the pinnacles are not under direct oceanic water influence. The flora of the barrier reef and lagoon reef patches is rich and similar in terms of species number (115 v. 106 sp.), and most of the species in these areas have a wide distribution pattern (18 and 14%, respectively, are restricted to these areas). However, barrier reefs and lagoon components cannot be considered as one single area, because the species composition differs from one area to the next, and only five species were found in both areas, (e.g. Chlorodesmis fastigiata, Hydrolithon murakoshii and Sebdenia sp.). The barrier reef is dominated by non-geniculate coralline algae, especially mastophoroids (Hydrolithon spp. and Neogoniolithon spp.), and Lithophylloids (Lithophyllum insipidium, L. flavescens and the branching L. kotschyanum) which form large crusts on hard substratum and play an important role in consolidating the rubbles and large pieces of corals resulting from destructive dynamite fishing practices. The shallow and soft-bottom areas of the lagoon were locally dominated by patches of green algae such as Halimeda incrassata and Caulerpa spp. (C. brachypus, C. taxifolia and C. sertularioides), and Cyanobacteria assemblages, which appear to be seasonal based on Chlorophyta Ochrophyta Rhodophyta Total 041 136 11 46 111 226 163 408 097 42 146 285 076 30 116 222 044 23 099 166 075 43 155 273 examination of satellite imagery data over several years. The flora of the coral reef patches in the lagoon area is dominated by abundant clumps of the common green algae, Halimeda opuntia and H. minima, as well as turf communities of algae, mostly red (e.g. Polysiphonia spp., Ceramium spp.). Few species (five) were restricted to this area: Codium mamillosum and Dasya spp. Finally, the outer reef slopes which are less rich than the adjacent barrier reef (90 species), contain 23% of the species that are not found in other areas; they are mostly Rhodophyta, such as Acanthophora pacifica, Chrysymenia okamurae, Platoma cyclocolpum, Predaea weldii, Prionitis sp., and a few Chlorophyta such as Rhipiliopsis howensis, Halimeda gracilis and H. hederacea. Few species are found in both the outer slope and barrier reef areas; they are mostly Halimeda spp. (H. distorta, H. taenicola and H. micronesica) and coralline algae. Overall, the most represented genera are Laurencia (10 species), followed by Halimeda (nine species), Ceramium (seven species) and Hydrolithon (five species). The general pattern observed from the distribution of the Wallis marine flora is characterised by very few species (4%: e.g. Halimeda opuntia, H. minima, Hydrolithon onkodes, Valonia fastigiata) shared between the different sites, while most of them are area-restricted. This agrees with what is described in French Polynesia by Payri et al. (2000) for the algae, and also by Galzin and Legendre (1988) for fishes as a result of biotic (inter specific competition, and grazing) and abiotic factors (microhabitat, nutrient levels and water movement, Littler and Littler 1994). Biogeographic comparisons Establishing biogeographic comparisons of an area’s flora requires accurate species taxonomy and also a similar sampling effort from both the study site and the other regions selected for the comparison. Thus, in respect of this assumption, the comparison has been restricted to the neighbouring regions and those located at the same latitude range (Table 3). Coralline algae and Cyanobacteria have not 388 A. D. R. N’Yeurt and C. E. Payri Australian Systematic Botany been included as they are generally understudied; forms and varieties of species have been counted as distinct taxa. The number of taxa (crustose corallines and Cyanobacteria excluded) of Wallis at first sight appear to be generally low compared with neighbouring islands such as Fiji and Samoa. However, this may be a reflection of limited sampling rather than a true biogeographical difference, as this study was based on a single 16-day survey in which some habitats (such as the turbulent eastern region of Wallis) were excluded. Wallis has no endemic or notable records; neighbouring Rotuma, on the other hand, has one endemic taxon (Avrainvillea rotumensis; N’Yeurt et al. 1996a) but this species has since been reported north of Rotuma, in Tuvalu (J. Orempuller, pers. comm.). It is interesting that several very common tropical genera, such as Hydroclathrus, Rosenvingea, Colpomenia, Chnoospora, Asteronema, Neomeris and Cheilosporum, are absent and to note that (excepted for Asteronema, Chnoospora, Neomeris and Cheilosporum) they do not occur either north of Fiji such as in Rotuma (N’Yeurt 1996), the Solomon Islands (Womesley and Bailey 1970), Nauru (South and Yen 1992) or Kiribati (South et al. 2001). It could be a seasonal factor, as these species are known to be mainly present during the warm season (Payri 1987) and collecting was done in Wallis at the onset of the cooler season. In the Solomon Islands, Womersley and Bailey (1970: 294) report another seasonal brown alga, Colpomenia sp., as rare during the cooler month of August. On the other hand, in isolated oceanic islands close to equatorial regions such as Rotuma, Wallis, Nauru, Kiribati there is no marked variation in air and sea temperatures throughout the year, and these conditions could exclude the presence of seasonal species whose distribution range does not reach these higher latitudes. In the case of habitat-specific species such as the widespread brown algae Chnoospora minima and usually associated Asteronema breviarticulatum, their absence from Wallis (but presence in Rotuma) could be explained by the lack of exposed, rocky habitas in the former island. However, seasonality cannot explain the puzzling absence of the green alga Halimeda discoidea, which is usually ubiquitous in the tropical Pacific throughout the year, or Cheilosporum spp. which is usually common in overhangs on the reef crest (both of these species, incidentally, do occur in neighbouring Rotuma). An instance of apparently disjunct north–south biogeographical distribution is the presence of the distinctive Japanese species Acrochaetium kurogii in Wallis, and also in the Cook Islands (ADR N’Yeurt, unpubl. data). However, hasty conclusions should not be made as such minute epiphytic species are easily overlooked in floristic surveys, and sampling effort is far from normalised in the region, with many areas remaining unexplored. It is becoming clear that an increasing number of species previously considered endemic or restricted to certain areas are in fact much more widespread or pantropical, as shown in the recent discovery of the deepwater red algae Predaea weldii and Gibsmithia hawaiiensis (type localities in the central Pacific) in South Africa (De Clerck et al. 2002) and Amansia rhodantha (type locality in the Indian Ocean) in the South Pacific (N’Yeurt 2002). Doty (1954) discussed the mutual exclusiveness of the genera Rhipilia and Sargassum in equatorial localities, such as Kiribati and Bikini atoll. Wallis island (13°18' S) could be considered close to the equator, and the presence of Rhipilia tenaculosa would, following this logic, prevent the occurrence of Sargassum. Interestingly, a single species of Sargassum, S. polycystum, occurs at Wallis, albeit only at one site which is a port of entry. It could be hypothesised that Sargassum does not occur in the natural state in Wallis, and its presence is the result of introduction via shipping. Whatever the case might be, other species are surely to be discovered in Wallis, especially in the turbulent eastern region, which is yet to be explored. The present study provides our first knowledge of the benthic marine macrophytes associated with the ecosystem diversity of Wallis. It is the first, but important, step towards the conservation and management of the habitats of this island’s marine environment. Acknowledgments The authors acknowledge financial support for this study from the French Coral Reef Reasearch Institute, IFRECOR. Mr Heroen Verbruggen (Ghent University, Belgium) is thanked for identifying Halimeda species, as well as for helping us with collections and sorting of material in the field, and for being a valuable dive buddy. We thank the following individuals: Mr Paino Vanai, Head of the Environmental Service and all his devoted staff, for logistical support in the field and on land, without which this mission would have been impossible; the Prefect and Chief Administrator of Wallis and Futuna; Lavelua, King of Wallis; the members of the Traditional Council of Chiefs of Wallis, and Professor C. Chauvet, Université de la Nouvelle-Calédonie. 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Memoirs of the Faculty of Fisheries, Hokkaido University 25, 97–152. Yoshida T, Mikami H (1996) Observations on Japanese species of the genus Martensia (Delesseriaceae, Rhodophyta), with the description of Neomartensia gen. nov. Phycological Research 44, 101–106. Manuscript received 17 September 2003, accepted 24 June 2004 Taxonomic index Acanthophora pacifica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthophora spicifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrochaetium barbadense. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrochaetium kurogii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actinotrichia fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiroa foliacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiroa fragilissima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiroa tribulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anotrichium tenue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antithamnion decipiens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asparagopsis taxiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balliella repens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boergesenia forbesii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boodlea composita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bostrychia tenella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bryopsis pennata var. secunda . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calothrix confervicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulacanthus ustulatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa biserrulata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa cupressoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa nummularia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa peltata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa racemosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa serrulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa sertularioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa taxifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa taxifolia ecad. tristichophylla . . . . . . . . . . . . . . . . . . . . . Centroceras minutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium codii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium flaccidum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium kramerii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium macilentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium sp. aff. C. marshallense . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium subdichotomum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceramium upolense. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 384 378 378 378 379 379 379 382 382 378 382 372 372 384 373 370 380 373 373 373 373 373 374 374 374 374 382 382 382 383 383 383 383 383 Chaetomorpha crassa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetomorpha linum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamaebotrys boergesenii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Champia compressa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Champia parvula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlorodesmis fastigiata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlorodesmis hildebrandtii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria arcuata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria armata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria dangeardii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria dasyphylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria leptacremon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondria simpliciuscula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chroodactylon ornatum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chronodrophycus cartilaginea . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrondrophycus dotyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrondrophycus parvipapillata . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrondrophycus succica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysymenia okamurae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cladophora vagabunda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Codium bulbopilum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Codium mamillosum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coelothrix irregularis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corallophila apiculata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corallophila huysmansii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corynocystis prostrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crouania minutissima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptonemia umbraticola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptonemia yendoi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasya anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasya baillouviana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyopteris repens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyosphaeria cavernosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyosphaeria versluysii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyota bartayresiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 372 382 382 382 375 375 384 384 385 385 385 385 377 385 385 385 385 382 372 373 373 382 383 383 380 383 378 379 383 384 376 372 372 376 Marine plants from Wallis Australian Systematic Botany Dictyota divaricata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Dictyota friabilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Diplothamnion jolyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Erythrotrichia carnea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Eupogodon geppii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Euptilota articulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Galaxaura fasciculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Galaxaura filamentosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Galaxaura marginata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Galaxaura subverticillata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Gelidiella acerosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Gelidiella maschrisiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Gelidiopsis intricata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Gelidiopsis scoparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Gelidium pusillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Gibsmithia hawaiiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Gracilaria textorii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Gracilaria sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Halichrysis coalescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Halimeda borneensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374 Halimeda distorta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374 Halimeda gracilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda hederacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda incrassata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda macroloba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda micronesica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda minima. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda opuntia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halimeda taenicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375 Halodule pinifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Halophila ovalis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Haloplegma duperreyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Halymenia actinophysa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Herposiphonia delicatula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Herposiphonia obscura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Heterosiphonia crispella var. laxa . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Hydrolithon farinosum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Hydrolithon murakoshii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Hydrolithon onkodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Hydrolithon reinboldii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Hydrolithon rupestris. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Hypnea cervicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Hypnea charoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Hypnea pannosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Hypnea saidana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Hypnea spinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Hypnea valentiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Hypoglossum simulans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Jania adhaerens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Kallymenia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Laurencia brachyclados . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Laurencia cervicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Laurencia flexilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Laurencia implicata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Laurencia mariannensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Laurencia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Leptofauchea anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Lithophyllum flavescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Lithophyllum insipidum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Lithophyllum kotschyanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Lithothamnion proliferum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Lithophyllum pygmaeum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Lobophora variegata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Lomentaria corallicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 397 Lyngbya majuscula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370 Lyngbya polychroa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Martensia fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Mastophora pacifica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Mesophyllum erubescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Mesophyllum mesomorphum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Meristotheca procumbens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Neogoniolithon brassica-florida. . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Nitophyllum adhaerens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Nostoc calcicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370 Oscillatoria bonnemaisonii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Oscillatoria margaritifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Padina boryana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Padina melemele. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Peyssonnelia bornetii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Peyssonnelia inamoena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Peyssonnelia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 Phormidium nigroviride . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Phormidium papyraceum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Phyllodictyon anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Platoma cyclocolpum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Pneophyllum conicum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Polysiphonia apiculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 Polysiphonia delicatula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Polysiphonia homoia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Polysiphonia poko . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Polysiphonia scopulorum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Polysiphonia sphaerocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Predaea laciniosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Predaea weldii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Prionitis sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Pterocladiella caerulescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Pterocladiella caloglossoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 Ptilothamnion sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Rhipidosiphon javensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Rhipilia tormentosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Rhipiliopsis howensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Sarconema filiforme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381 Sargassum polycystum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Scytonema polycystum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370 Sebdenia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Siphonocladus tropicus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Sphacelaria tribuloides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Sphaenosiphon olivaceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369 Spirulina subsalsa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Sporolithon ptychoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 Spyridia filamentosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Stylonema alsidii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Symploca atlantica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Symploca hydnoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Symploca laeteviridis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Syringodium isoetifolium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Tolypiocladia glomerulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 Turbinaria conoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Turbinaria ornata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Ulva clathrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Ulva flexuosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Valonia aegagropila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Valonia fastigiata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Valonia macrophysa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Ventricaria ventricosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372 Wrangelia argus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Wrangelia dumontii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Wurdemannia miniata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 http://www.publish.csiro.au/journals/asb

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