Marine algal flora of french polynesia. II. Chlorophyceae (green algae)

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.

Cryptogamie, Algol., 2007, 28 (1): 3-88 © 2007 Adac. Tous droits réservés Marine Algal Flora of French Polynesia II. Chlorophyceae (green algae) Antoine D. R. N’YEURT a* & Claude E. PAYRI a, b a Laboratoire Terre-Océan, Université de la Polynésie française, B.P. 6570 Faa’a 98702, Tahiti, French Polynesia b UMR 7138, Systématique, Adaptation, Evolution, Equipe Biodiversité Marine Tropicale, IRD-Nouméa BPA5, 98848 Nouméa cedex, New Caledonia (Received 9 December 2005, accepted 8 May 2006) Abstract – This second paper in a monographic series on the marine algae of French Polynesia gives a detailed coverage of the species of Chlorophyceae occurring in these islands. A total of 81 taxa are presented; of these, 23 represent new records for the local flora, 1 (Halimeda heteromorpha) is a new species and 2 (Microdictyon sp. and Struveopsis sp.) are newly discovered taxa. The benthic marine green algal flora has most affinities with the neighbouring Cook Islands (Sørensen Index of 0.43) but only with 44% of shared species with that archipelago, while 69 and 64% of species are shared with the Fijian and Micronesian islands, respectively. About 75% of the flora is shared with the Indian and Pacific Ocean in general; 3 species (or about 4% of the total flora) seem restricted to French Polynesia. algae / biogeography / distribution / floristics / French Polynesia / Chlorophyceae / new records / taxonomy Résumé – Flore des algues marines de la Polynésie française. II. Chlorophyta (algues vertes). Cette deuxième publication dans une série monographique sur la flore marine de la Polynésie française donne une distribution détaillée des espèces de Chlorophycées qui sont présentes dans ces îles. Un total de 81 taxons sont présentés, dont 23 sont nouveaux pour la Polynésie française, 1 (Halimeda heteromorpha) représente une nouvelle espèce et 2 (Microdictyon sp. et Struveopsis sp.) représentent de nouveaux taxons. La flore a le plus d’affinités avec l’archipel voisin des îles Cook (Index de Sørensen de 0.43), mais seulement avec 44 % des espèces en commun avec ce groupe d’îles, tandis que 69 et 64 % des espèces sont en commun avec les îles Fidji et la Micronésie, respectivement. Environ 75 % de la flore est en commun avec l’Océan Indien et l’Océan Pacifique en général ; 3 espèces (ou environ 4 % de la flore) semblent restreintes à la Polynésie française. algues / biogéographie / distribution / floristique / Polynésie française / Chlorophyceae / nouveaux reports / taxonomie * Correspondence and reprints: nyeurt@yahoo.com Communicating editor: Frederik Leliaert 4 A. D. R. N’Yeurt & C. E. Payri INTRODUCTION This study gives a detailed overview of the species of Chlorophyceae occurring in French Polynesia and supplements and revises the taxa previously reported in Setchell (1926), Meinesz et al. (1981), Payri & Meinesz (1985b), Payri & N’Yeurt (1997), Payri et al. (2000), and Conte & Payri (2002, 2006). The history of early phycological collecting in French Polynesia was treated in N’Yeurt & Payri (2006). The earlier works covered some 96 taxa of Chlorophyta, but this number could not be confirmed. The present paper documents the distribution of Chlorophyta within French Polynesia and compares the similarity of the benthic marine green algal flora of French Polynesia to other Pacific areas and discusses the biogeographical patterns encountered. MATERIALS AND METHODS Material was collected using SCUBA, snorkelling or reef-walking. Herbarium specimens were pressed using standard techniques, and representative parts of thalli and turf algae stored in 4% buffered formalin in sealed plastic bags packed in a light-proof container for shipment and later anatomical examination in the laboratory. Herbarium specimens were photographed using a benchmounted digital camera (NIKON Coolpix 995). Photomicrographs were obtained using an OLYMPUS C-4040 digital camera fitted on an OLYMPUS BH2 microscope. Voucher specimens are housed at the Phycological Herbarium of the Université de Polynésie française in Tahiti (UPF), with “S” referring to slide collections. These collections are accessible online at the following address: http:/ /biodiv.upf.pf/base/. “IFR” refers to holdings in the herbarium of the French Institute of Research on Coral Reef Environment (IFRECOR) in Bora Bora, French Polynesia (curated by Mr. Denis Schneider). Where necessary, available French Polynesian herbarium records in UPF have been re-verified in the light of new taxonomic information, and some of Setchell & Parks and other early collections in BM (British Museum, London), PC (Museum National d’Histoire Naturelle, Paris), SAP (Hokkaido University, Sapporo) and UC (University of California, Berkeley) examined either on site or on loan. Unfortunately, some voucher specimens of species from Moorea listed in Payri (1987), which contain some taxa not held at UPF, could not be located and are presumed lost. Consequently, only those records which could be confirmed on the basis of existing collections have been included in this study. The taxonomy generally follows that of Silva et al. (1987; 1996), De Reviers (2003), and is updated with other sources where necessary as stated in the text. For each taxon, basionym and type locality information is provided, followed by relevant synonyms and selected bibliographic references. Asterisks (*) indicate new published records or species for French Polynesia. French Polynesian green algae 5 RESULTS AND OBSERVATIONS Key to genera of French Polynesian Chlorophyta 1a. Thallus calcified or partly calcified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Thallus totally uncalcified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2a. Thallus cylindrical (club-shaped) and unsegmented, or a flabellate blade 3 2b. Thallus flattened or circular, segmented, straight or slightly arcuate . . . . . . 4 3a. Thallus club-shaped, unbranched, slightly arcuate. . . . . . . . . . . . . . Neomeris 3b. Thallus fan-shaped, dichotomously branched, monostromatic Rhipidosiphon 4a. Segments clavate, fused into a circular stipitate disc . . . . . . Parvocaulis 4b. Segments flattened, in branched chains, not fused into a disk Halimeda 5a. Thallus as a solitary ovoid vesicle, to 5 cm high . . . . . . . . . . . . . . Ventricaria 5b. Thallus not ovoid, usually gregarious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Thallus blade-like, compressed or tubular, or of branches forming a fused network . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b. Thallus vesicular, filamentous or plumose, not forming a network . . . . . . 11 7a. Thallus three-dimentional, forming spongy crispy masses . . . . . . . . Boodlea 7b. Thallus in a single plane, not a spongy mass . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Thallus thick and gelatinous, composed of a mass of non-organized microscopic cells less than 10 µm in diameter, with sickle-shaped chloroplasts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verdigellas 8b. Thallus thin, not gelatinous, cells more than 10 µm in diameter, chloroplasts not sickle-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9a. Thallus monostromatic, mesh-like and composed of fused branches, not membranous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Thallus distromatic, hollow tubular, compressed or membranous, not meshlike or composed of fused components . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulva 10a. Thallus broadly lanceolate, with a long stipe; tenacular cells predominantly consisting of small, modified, distally crenellated (type-3) cells on terminal segments. . . . . . . . . . . . . . . . . . . . . . . . . . Phyllodictyon 10b. Thallus forming rosettes or blunt paddle-shaped, attached via short rhizoids, tenacular cells (type-1) consisting solely of unmodified segments with a distal crenellated or smooth thickened pad . . . . . . Microdictyon 11a. Thallus filamentous or tuft-like. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11b. Thallus not filamentous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 12a. Filaments regularly separated by cross-walls (septate) or lacking crosswalls at the base of at least some lateral branches . . . . . . . . . . . . . . . 13 12b. Filaments coenocytic, or with few partitions if any. . . . . . . . . . . . . . . 14 13a. Thallus branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 13b. Thallus unbranched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 14a. Axes sparsely branched, constrictions absent or rare . . . . . . . . Derbesia 14b. Axes profusely branched, constrictions abundant . . . . . . . . . . . . . . . . 15 15a. Thallus with radial, siphonous vesicles about an erect axis; cell divison segregative (presence of globular lateral branch precursors in axes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphonocladus 15b. Thallus with dichotomous, pinnate or secund branching; cell division not segregative . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 16a. Thallus lacking cross-walls at the base of at least some lateral branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 16b. Thallus with cross walls at the base of all lateral branches . . Cladophora 6 A. D. R. N’Yeurt & C. E. Payri 17a. Thallus forming fleecy masses; filament diameter less than 70 µm; basal cell absent or unmodified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhizoclonium 17b. Thallus erect or in clumps; filament diameter more than 70 µm; modified basal cell often present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetomorpha 18a. Thallus plumose, with pinnate or secund branching . . . . . . . . Bryopsis 18b. Thallus tuft-like, filaments dichotomous . . . . . . . . . . . . . . . . . . . . . . . . 20 19a. Thallus flaccid or in clumps, lacking a percurrent main axis Cladophoropsis 19b. Thallus tufted, with a stratified percurrent main axis . . . . . . . . . Struveopsis 20a. Thallus tufted, filaments not adhering except at the base Chlorodesmis 20b. Thallus blade-like, filaments interwoven throughout . . . . Avrainvillea 21a. Thallus bubble-like, or of loosely-adhering, clavate or cylindrical vesicles 22 21b. Thallus otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 22a. Thallus bubble-like, composed of small closely-adhering hexagonal cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyosphaeria 22b. Thallus consisting of large clavate or cylindrical vesicles . . . . . . . . . . 23 23a. Vesicles irregular or clavate; thallus loosely aggregated, not prostrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valonia 23b. Vesicles cylindrical, sometimes arcuate; thallus prostrate or forming balls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valoniopsis 24b. Thallus with a creeping stolon bearing uprights of various shapes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa 24a. Thallus without a creeping stolon, spongiose; habit globular, upright or convoluted, internal filaments with terminal inflated utricles . . Codium Division CHLOROPHYTA T. Cavalier-Smith Order Tetrasporales Lemmermann Family Palmellopsidaceae Korshikov Genus Verdigellas Ballantine et J. Norris *Verdigellas peltata Ballantine et J. Norris, 1994: 369, figs 5-7 (type locality: Mona Island, Puerto Rico). Brazil: Bravin et al., 1999: 124, figs 6-8; Fiji: Littler & Littler, 2003: 194 (as Verdigellas sp.). (Figs 1-2) Misapplied name: Palmogloea protuberans (Smith et Sowerby) Kützing, 1843: 176. French Polynesia: Payri et al., 2000: 62. Material examined: Hiti, 10 Nov. 1996, leg. J. Orempuller, UPF 129; Marokau, 22 Nov. 1996, leg. J. Orempuller, UPF 127, 128; Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 698. Thallus to 10 cm high and up to 2 mm thick, dark green and turgid, attached to the substratum via small, discrete basal holdfasts. Habit irregularly lobed, erect (Fig. 1). The surface of the thallus is smooth and gelatinous, with usually ruffled, thickened margins. Internally, structure consists of a mass of nonorganized, microscopic spherical cells 6-9 µm in diameter, with characteristic sickle-shaped chloroplasts (Fig. 2). Remarks: Growing as isolated thalli, Verdigellas peltata has been recorded from 30 to 60 m deep, in atolls of the Tuamotu archipelago. This jelly-like alga of grassgreen colour appears to be relatively common in deepwater and low-light habitats in tropical regions, from the Caribbean to the South Atlantic (Bravin et al., 1999) and the South Pacific (Littler & Littler, 2003; this study). The nearest related genus to Verdigellas is Palmophyllum Kützing (1846), but species in that genus have a gelatinous stalk, are leathery in texture (as opposed to soft and jelly-like in Verdigellas) and the species closest to V. peltata, Palmophyllum crassum (Naccari) Rabenhorst (1868) differs in having concentrically zoned lobes (Ballantine & French Polynesian green algae 7 Norris, 1994: 371; Feldmann, 1937: 176). Palmophyllum umbracola (Nelson & Ryan, 1986) from New Zealand could be mistaken for V. peltata, but the former has a leathery texture unlike the soft, slippery consistency of V. peltata (Ballantine & Norris, 1994: 371). The French Polynesian material is broadly peltate, forming gelatinous unbranched cushions 1-2 mm thick attached to the substratum by several discrete stipe-like holdfasts. It hence differs from V. fimbriata (Ballantine & J. Norris, 1994: 369) and V. nektongammea (Ballantine & Aponte, 1996) which are erect with marginal branches, or lobed with cavities in transverse sections, respectively. In French Polynesia it has been previously misidentified as Palmogloea protuberans (Smith et Sowerby) Kützing, a superficially similar freshwater species (Payri et al., 2000). Order Ulvales Blackman et Tansley Family Ulvaceae J.V. Lamouroux Genus Ulva Linnaeus Recent molecular studies (Hayden et al., 2003; Shimada et al., 2003) have confirmed previous vegetative and reproductive indications (Bonneau, 1977; Provasoli & Pintner, 1980; Kraft, 2000) that Ulva and Enteromorpha are not distinct genera. The artificial separation of Ulva into distromatic (“Ulva”) and tubular (“Enteromorpha”) forms (Link, 1820) had been done for purposes of convenience, and retained since. French Polynesian records of Enteromorpha are hence transferred to Ulva based on the new combinations proposed by Hayden et al. (2003: 288, table 4). Five species of the genus are represented in collections from French Polynesia in UPF. Since most of the South Pacific species of Ulva have been identified in the past using European names, this taxonomy may not be applicable to tropical populations (G.T. Kraft, pers. com.). Key to the French Polynesian species of Ulva 1a. Thallus usually flabellate, sheet-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Thallus usually tubular to compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Cells in cross-section rectilinear, arranged in distinct palisade layers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. rigida 2b. Cells in cross-section rectangular, not in distinct palisade layers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. lactuca 3a. Thallus cylindrical, sparsely branched with uniseriate laterals . . . U. flexuosa 3b. Thallus compressed, uniseriate laterals absent. . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Laterals radially arranged, at a narrow angle. . . . . . . . . . . . . U. clathrata 4b. Laterals not radially arranged, at wide angles . . . . . . . . . . . U. compressa *Ulva clathrata (Roth) C. Agardh, 1811: 23 (Fig. 3) Basionym: Conferva clathrata Roth, 1806: 175-178 (type locality: Fehmarn, southwestern Baltic, according to Berger et al., 2003: 288). Homotypic synonym (given by Hayden et al., 2003): Enteromorpha clathrata (Roth) Greville, 1830: lxvi, 181. Viêt Nam: Dawson, (1954: 384, fig. 6d-e); Thailand: Egerod, 1974: 134, fig. 4; Indian Ocean: Silva et al., 1996: 729; Lord Howe Island: Kraft, 2000: 525, fig. 8; Fiji: N’Yeurt, 2001: 693, figs 2, 3a-b; Samoa: Skelton & South, 2002a: 160, fig. 22A-B; 2002b: 8, pl. 5 figs 29-33, pl. 6 fig. 34; Hawaiian Islands: Abbott & Huisman, 2004: 46, fig. 5A-C. Material examined: Tarakoi islet, Rapa Island, Australs, 5 Nov. 2002, leg. C. E. Payri, UPF 2018, 2019. 8 A. D. R. N’Yeurt & C. E. Payri Thallus irregularly branched and proliferous, forming entangled masses. Filaments of main axis 280-300 µm in diameter, lateral branchlets 40-100 µm in diameter, issued distichously to radially at a narrow angle. Ultimate branchlets and distal portions of axes multiseriate, 20-40 µm in diameter, with characteristic bulbous expansions. Cells angular and mostly regularly arranged in surface view, 11-17 µm in diameter. 1-4 pyrenoids per cell (usually 2-3). Remarks: Ulva clathrata is readily distinguished by multiseriate laterals ending in bulbous expansions. Ulva compressa Linnaeus, 1753: pl. 2: 1163 (type locality: Baltic Sea) (Figs 4-5) Homotypic synonym (given by Hayden et al., 2003): Enteromorpha compressa (Linnaeus) Greville, 1830: 180. Marshall Islands: Dawson, 1956: 27, fig. 1; Europe: Bliding, 1963: 132, fig. 82a-f, 83a-d, 84a-f; Indian Ocean: Silva et al., 1996: 729; Blomster et al., 1998: 325, figs 4-6, 35-41; Lord Howe Island: Kraft, 2000: 521, fig. 5A-F; Fiji: N’Yeurt, 2001: 695. Material examined: Opunohu, Moorea, 1 Jul. 1997, leg. A. D. R. N’Yeurt, UPF 81; Faa’a, Tahiti, leg. A. D. R. N’Yeurt, 13 Apr. 1997, UPF 83-84, 7 May 1997, UPF 82; Taharaa, Tahiti, 22 Apr. 1998, leg. C. E. Payri, UPF 714. Thallus simple to repeatedly branched, 16-25 mm high, composed of compressed, upwardly broadened axes 2-5 mm in diameter. Cells polygonal to rounded, about 14 × 23 µm, densely arranged and appearing compressed in surface view. Remarks: Ulva compressa is distinct from U. intestinalis, and Enteromorpha usneoides Bonnemaison ex. J. Agardh (1883: 159) appears to be an ecotype of U. compressa (Blomster et al., 1998, Hayden et al., 2003). Ulva compressa is morphologically differentiated from U. intestinalis by the lack of branching in the latter. It is a cosmopolitan biofouling species, with extreme morphological plasticity varying from low tubular fronds to flattened blades to 120 mm across (Kraft, 2000: 521). The latter observation of morphological plasticity is in support of the accumulating molecular evidence for merging of Enteromorpha into Ulva. Ulva flexuosa Wulfen, 1803: 1 (Fig. 6) Basionym: Conferva flexuosa Roth, 1800: 188, nom. illeg. (type locality: Duino, near Trieste, Adriatic Sea). Homotypic synonym (given by Hayden et al., 2003): Enteromorpha flexuosa (Wulfen) J. Agardh, 1883: 126 (incl. subsp. flexuosa). French Polynesia: Payri & N’Yeurt, 1997: 879, Payri et al., 2000: 62, Conte & Payri, 2002: 166, fig. 1; Europe: Bliding, 1963: 73, figs 38-40; Indian Ocean: Silva et al., 1996: 733; Lord Howe Island: Kraft, 2000: 523, fig. 7 A-E; Namibia: Lluch, 2002: 179, figs 199, 200A-D. Heterotypic synonym (given by Hayden et al., 2003): Enteromorpha tubulosa (Kützing) Kützing, 1856: 11. French Polynesia: Setchell, 1926: 68. Material Examined: Ua Huka, Marquesas, Aug. 1999, leg. E. Conte & C. E. Payri, UPF 801, 819; Opunohu, Moorea, 1 Jul. 1997, leg. A. D. R. N’Yeurt, UPF 81; Faa’a, Tahiti, 13 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 83, 84, 7 May 1997, UPF 82; Taharaa, Tahiti, 22 Apr. 1998, leg. C. E. Payri, UPF 714. Plants light-green and fleecy, tubular in cross-section, 150-155 µm in diameter and up to 20 cm long; sparsely to profusely branched with a flexuous basal portion. Lateral branches often uniseriate, 25-30 µm in diameter. In surface view cells subrectangular, up to 25 by 35 µm, arranged in distinct brick-like longitudinal rows; 3-6 (usually 3) pyrenoids per cell. French Polynesian green algae 9 Figs 1-9. 1. Verdigellas peltata. Habit of dried plant (UPF 698). Scale = 10 mm. 2. Verdigellas peltata. Cellular matrix, with sickle-shaped chloroplast (arrow) (UPF 698). Scale= 20 µm. 3. Ulva clathrata (UPF 2018); habit. Scale = 200 µm. 4. Ulva compressa (UPF 81); habit. Scale = 10 mm. 5. Ulva compressa (UPF 82). View of surface cells, showing pyrenoids (arrowhead). Scale = 20 µm. 6. Ulva flexuosa (UPF WSS228). Habit showing uniseriate lateral branchlet (arrowhead). Scale = 100 µm. 7. Ulva lactuca (UPF 136). Habit. Scale = 10 mm. 8. Ulva lactuca (UPF 2715). Transverse section of thallus, showing squarish double row of cells. Scale = 20 µm. 9. Ulva rigida (UPF RPS 184). Transverse section of thallus, showing rows of rectilinear cells. Scale = 30 µm. 10 A. D. R. N’Yeurt & C. E. Payri Remarks: This species, which is one of the commonest of the genus in the tropical Pacific, includes at least one subspecies, subsp. paradoxa (Dillwyn) Bliding, (1963: 79), characterized by abundant uniseriate laterals and less regular arrangement of cell rows. Ulva lactuca Linnaeus, 1753: 1163 (type locality: “in Oceano”). French Polynesia: Setchell, 1926: 69, pl. 7 figs 1-4 (as f. lacinulata), Payri et al., 2000: 64, Conte & Payri, 2002: 167, fig. 5; South Australia: Womersley, 1984: 141, figs 44 A, 45 A-C; Indian Ocean: Silva et al., 1996: 745; Lord Howe Island: Kraft, 2000: 531, fig. 10. (Figs 7-8) Material examined: Trou du souffleur, Tahiti, 18 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 137; Hotel Beachcomber, Faa’a, Tahiti, 13 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 135, 139; 7 May 1997, UPF 136; Taharaa, Tahiti, 14 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 138; Arue, Tahiti, 6 Apr. 1998, leg. C. E. Payri, UPF 699; Ua Huka, Marquesas, Aug. 1999, leg. E. Conte & C. E. Payri, UPF 818; Motu Togegie, Gambier, 22 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 810, 811; Rapa Iti, 4 Nov. 2002, leg. C. E. Payri, UPF 1941, 2689; Pt. Komire, Rapa, 13 Nov. 2002, leg. J. L. Menou, UPF 2198. Thallus to 10 cm broad, flat and simple, membranous and irregularly lacerate with small holes, attached by a small rhizoidal holdfast (Fig. 7). Blade distromatic; cells squarish, 20-23 µm long and 20-21 µm tall, subrectangular and adherent (Fig. 8), with a single outer chloroplast. Pyrenoids 1-2 per cell. Remarks: The commonest Ulva species reported in the literature, U. lactuca is a poorly defined and documented species. Some authors (e.g. Abbott & Huisman, 2004) do not venture using the name for tropical species. We follow the description of Kraft (2000) for the French Polynesian specimens, which have squarish cells and are distinct from U. rigida (described below) which has rectilinear cells. Setchell (1926: 69) reports this edible species as being consumed by Tahitians at the time, but recently in French Polynesia it is only being used as food in the Marquesas (Conte & Payri, 2002, 2006). Ulva rigida C. Agardh, 1823: 410 (type locality: Cádiz, Spain). French Polynesia: Taylor, 1973: 38 (Makatea); Netherlands: Koeman & van den Hoek, 1980: 37, figs 78-107; Indian Ocean: Silva et al., 1996: 750; Belize: Littler & Littler, 1997: 87, fig. 114; Lord Howe Island: Kraft, 2000: 532, fig. 11; Namibia: Lluch, 2002: 185, figs 210, 211A-B. (Fig. 9) Material examined: Tarakoi Islet, Rapa Island, Australs, 5 Nov. 2002, leg. C. E. Payri, UPF 2687, RPS 182, 183, 184. Thallus 25-30 mm broad and 88-100 µm thick, sparsely perforate, with a coarser basal portion. Cell rectilinear, 45-50 µm long and 13-18 µm broad, arranged in distinct double palisade layers. Remarks: This species is distinguished by its rectilinear cells arranged in palisade layers, and the coarseness of the lower portions of the thallus (Kraft, 2000). So far in French Polynesia, only reported from Makatea (Tuamotu) and in the cooler waters of the southern Australs (Rapa Island). Order Cladophorales Haeckel Family Anadyomenaceae Kützing Genus Microdictyon Decaisne With some 20 tropical to subtropical species, the most comprehensive monographic treatment of Microdictyon to date remains the work of Setchell (1929). In French Polynesia, Microdictyon occurs in atolls of the Tuamotu French Polynesian green algae 11 Archipelago, the Gambier, Society and Rapa in the Australs. Covers of Microdictyon in typical rosettes among coral debris are typical of the algal flora of the lagoons in atolls of the Tuamotu Archipelago. Payri & N’Yeurt (1997) report four species of the genus in French Polynesia, but only three have been found, some growing together in the same habitats. Reports of M. agardhianum and M. setchellianum need verification as both these names are taxonomically questionable (see Silva et al., 1996: 758, 934). Key to the French Polynesian species of Microdictyon 1a. Thallus firm, leaf-like, with clear basal umbilicate rhizoidal holdfast. . M. sp. 1b. Thallus flaccid, forming expanded blades without clear basal holdfast . . . . 2 2a. Main filaments below 250 µm in diameter, attachment pads smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. umbilicatum 2b. Main filaments above 350 µm in diameter, attachment pads crenellated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. okamurae Microdictyon okamurae Setchell, 1925: 107; 1929: 553, figs 76-84 (type locality: Ryukyuretto, Japan). French Polynesia: Taylor, 1973: 38 (Tikehau); Payri & N’Yeurt, 1997: 880; Payri et al., 2000: 66; Indian Ocean: Silva et al., 1996: 757. (Figs 10-12) Material examined: Reitoru, 25 Sep. 1992, leg. R. Galzin, UPF 120; Marokau, 22 Sep. 1996, leg. J. Orempuller, UPF 113; Tekokota, 26 Sep. 1996, leg. J. Orempuller, UPF 122; Hiti, 12 Nov. 1996, leg. J. Orempuller, UPF 123; Haraiki, 15 Nov. 1996, leg. J. Orempuller, UPF 112, 117; Hikueru, 16 Nov. 1996, leg. J. Orempuller, UPF 116; Hiti, 12 Nov. 1996, leg. J. Orempuller, UPF 118; Marokau, 22 Nov. 1996, leg. J. Orempuller, UPF 119. Thallus light yellow-green (remaining so when dried), consisting of a monostromatic, delicate reticulate blade up to 5 by 12 cm, forming a meshwork with veins (Fig. 10). Main filaments 400-500 µm in diameter, terminal cells 180200 µm in diameter, slightly tapered. Branching mostly opposite. Branch anastomosis frequent, taking place by means of crenulated attachments pads formed on the tip of unmodified terminal cells (type-1 tenacular cells in Leliaert, 2004: 104; Figs 11-12). Remarks: A common and often dominant component of the atoll marine flora, forming extensive rosettes on coralline formations and other hard substrata in the lagoons. This coarser species in French Polynesia appears to be restricted to atoll habitats, unlike M. umbilicatum which occurs as far south as the high islands of Rapa in the Australs, and east to the Gambier. Microdictyon umbilicatum (Velley) Zanardini, 1862: 461. French Polynesia: Payri & N’Yeurt, 1997: 880, Payri et al., 2000: 66; Setchell, 1929: 503, figs 21-27; South Australia: Womersley, 1984: 217, fig. 72; Indian Ocean: Silva et al., 1996: 758; Lord Howe Island: Kraft, 2000: 570, fig. 24 D-F; Hawaiian Islands: Egerod, 1952: 363, pl. 32b figs 5, 6b; Abbott & Huisman, 2004: 62, fig. 15B. (Figs 13-14) Basionym: Conferva umbilicata Velley, 1800: 169, pl. 7 (type locality: New South Wales, Australia). Heterotypic synonym (according to Abbott & Huisman, 2004): Microdictyon japonicum Setchell, 1925: 107. Fiji: N’Yeurt, 2001: 699, fig. 27; Samoa: Skelton & South, 2002a: 161, fig. 22C. Material examined: Reitoru, 25 Sep. 1992, leg. R. Galzin, UPF 121; Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 614; Papeari, Tahiti, 24 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 133; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 894; Marotiri, Australs, 6 Nov. 2002, leg. IRD, UPF 2109; Rapa Island, Australs: Karapoo, 1 Nov. 2002, leg. J. L. Menou, UPF 1873; Ha’urei, 11 Nov. 2002, leg. J. L. Menou, UPF 2141, 13 Nov. 2002, UPF 2172, 2182; 12 A. D. R. N’Yeurt & C. E. Payri Figs 10-15. 10. Microdictyon okamurae (UPF 2697). Habit of mesh-like thallus. Scale = 300 µm. 11. Microdictyon okamurae (UPF 123). Detail of crenulate attachment pad making contact with another part of the thallus (arrowhead). Scale = 40 µm. 12. Microdictyon okamurae (UPF 2697). Branchlet with crenulate attachment pad (arrowhead). Scale = 40 µm. 13. Microdictyon umbilicatum (UPF RPS 217). Habit of mesh-like thallus. Scale = 400 µm. 14. Microdictyon umbilicatum (UPF 2698). Detail of smooth, circular attachment pad (arrowhead). Scale = 40 µm. 15. Microdictyon sp. (UPF RPS 33). Detail of stipe, showing a basal rhizoid (arrowhead). Scale = 200 µm. Rarapai, 12 Nov. 2002, leg. J. L. Menou, UPF 2153, 30 Nov. 2002, UPF 2442; Komire Pt., 13 Nov. 2002, leg. J. L. Menou, UPF 2200; east oceanic slope, 23 Nov. 2002, leg. J. L. Menou, UPF 2329; Tarakoi, 6 Nov. 2002, leg. C. E. Payri, UPF 2038; Anakaturinako Bay, 8 Nov. 2002, leg. C. E. Payri, UPF 2074; Tauna islet, 27 Nov. 2002, leg. C. E. Payri, UPF 2369, 2370. French Polynesian green algae 13 Thallus dark green and rigid (turning blackish when dry), consisting of a monostromatic, delicate reticulate blade up to 5 by 12 cm, forming a mesh with prominent veins (Fig. 13). Attachment to the substratum via an umbilicate stipe. Main filaments (120) – 200-220 µm in diameter, terminal cells 40-52 µm in diameter, gradually tapered with a blunt tip. Branching irregularly alternate or opposite to stellate, with up to 4 branches per mother cells. Branch anastomosis frequent, taking place by means of smooth, rounded attachments pads formed on the tip of unmodified terminal cells (Fig. 14). Remarks: By far the commonest Microdictyon species encountered in French Polynesia, to date reported from all archipelagos except the Marquesas, it can be distinguished from M. okamurae by its finer mesh texture and generally darker green colour, usually turning black when dried. Plants from Tahiti, Gambier and Marotiri (UPF 133, 894, 2109) have smaller main filament diameters (120-130 µm) and do not turn black on drying, and would have previously been placed under M. japonicum Setchell. Kraft (2000) and Abbott & Huisman (2004), after comparing respective type specimens, have placed M. japonicum in synonymy with M. umbilicatum. *Microdictyon sp. (Fig. 15) Material examined: Marotiri, Australs, 6 Nov. 2002, leg. IRD, UPF 2110; Northern plateau, Rapa Island, 25 Nov. 2002, leg. J. L. Menou, UPF RPS 33. Thallus grass-green, rigid and orbicular, rugose, 5-10 mm high and 4 to 20 mm broad, with an umbilicate stipe 1-2 mm long, attached via long blunt-tipped rhizoids issued from basal segments. Branching acropetal and fan-shaped, becoming stellate, with secondary branchlets attaching perpendicularly to other primary segments via smooth attachment pads. Cells of mid-thallus 80-110 µm in diameter, broader above than below; ultimate segments blunt-ended, 40-30 µm in diameter. Edge of thallus irregularly open, without lateral cohesion between ultimate cells. Remarks: Growing at 40 to 52 m depth, in French Polynesia so far only reported from the southern Australs (Rapa, Marotiri). This diminutive, stiped Microdictyon species bears some resemblance to M. calodictyon (Montagne) Kützing, from the Canary Islands, and to an undescribed species from New Caledonia. As pointed out by Børgesen (1925: 35), M. calodictyon is similar, especially as regards the basal portion, to Rhipidiphyllon reticulatum (Askenasy) Heydrich, reported from Easter Island by Børgesen (1924: 251, figs 3-4), and from the Marshall Islands by Dawson (1956: 32, fig. 10). Rhipidiphyllon, however, differs from the French Polynesian plants by its more lax branching, smaller stature (no more than 5 mm broad) and does not form a network as in even young plants of M. calodictyon. It is not ruled out, however, that both species could be related or represented different ecomorphs or growth stages of the same entity. Studies in progress (Payri et al., in prep) should elucidate the affinities of the South Pacific plants in the near future. Family Cladophoraceae Wille Genus Chaetomorpha Kützing Of the two species of Chaetomorpha accredited to Tahiti by Setchell (1926: 72), none have been recollected since, although a further two species described below are reported in collections. 14 A. D. R. N’Yeurt & C. E. Payri Key to the French Polynesian species of Chaetomorpha 1a. Mature filaments straight and basally attached, 200-250 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. antennina 1b. Mature filaments bent or coiled, free or creeping, entangled . . . . . . . . . . . . 2 2a. Filaments 80-200 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . C. fibrosa 2b. Filaments 300-600 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Basal cells absent, cells of uniform diameter throughout, rhizoids absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. linum 3b. Basal cells curved, cells deflexed and creeping with abundant rhizoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. basiretrorsa Chaetomorpha antennina (Bory de Saint-Vincent) Kützing, 1847: 166. French Polynesia: Payri et al., 2000: 68; Indian Ocean: Silva et al., 1996: 759; Lord Howe Island: Kraft, 2000: 538, fig. 13 A-C. (Fig. 16) Basionym: Conferva antennina Bory de Saint-Vincent, 1804: 381 (type locality: La Réunion). Material examined: Taharaa, Tahiti, 18 May 1996, leg C. E. Payri, UPF 687; 14 Apr. 1997, leg C. E. Payri & A. D. R. N’Yeurt, UPF 57; 30 Mar. 1998, leg. A. D. R. N’Yeurt, UPF 680; Karapoo, Rapa, 1 Nov. 2002, leg. C. E. Payri, UPF 1857; Akatamiro Bay, Rapa, 2 Dec. 2002, leg. J. L. Menou, UPF 2451. Thallus to 30 mm high and 15 mm broad, light yellowish-green, consisting of tufts of stiff, unbranched, segmented filaments to 250 µm in diameter. Basal cells straight, erect and permanent, 1.5-3 mm in length (Fig. 16b). Individual segments are barrel-shaped, with a thick cell wall (Fig. 16a). Remarks: Forming tufts on the reefs rim and in areas exposed to surf. The Tahitian and Rapa plants grow attached in dense tufts and have straight basal cells 1.5-3 mm in length, and thus conform to C. antennina and not to C. basiretrorsa Setchell (1926: 72, pl. 8 figs 4-6), a new species described from Punaruu, Tahiti. This leaves the question as to why the presently common C. antennina (a species familiar to Setchell, who had found it earlier in American Samoa) was absent from the reefs when Setchell visited Tahiti in 1922, and especially from the Tahara’a site from where he had made an extensive collection of algae. Chaetomorpha basiretrorsa Setchell, 1926: 72, pl. 8 figs 4-6 (type locality: Punaruu Pass, Tahiti). (Fig. 17) Heterotypic synonyms: Chaetomorpha prostrata P. Anand, 1940: 5, figs 5A-B, 6 (type locality: Manora, Karachi, Pakistan). Pakistan: Nizamuddin & Begum, 1973: 16, figs 67-68. — Rhizoclonium grande Børgesen, 1935: 14, figs 5-6 (type locality: Bombay, India). Pakistan: Nizamuddin & Begum, 1973: 16, fig. 69; Thailand: Egerod, 1975: 43, figs. 1-4; Tanzania: Jaasund, 1976: 5, fig. 12; Somalia: Sartoni, 1986: 361, fig. 3D; Seychelles: Wynne, 1995: 328, fig. 87; Rotuma: N’Yeurt, 1996: 368, figs 9, 20; Hawaiian Islands: Gilbert, 1965: 486, fig. 4E; Abbott & Huisman, 2004: 82, fig. 25B. — Rhizoclonium robustum Setchell et Gardner, 1937: 73, pl. 5 fig. 11 (type locality: North Seymour Island, Galapagos). Misapplied name: Cladophoropsis javanica (Kützing) P. Silva in Silva et al., 1996: 792 (Indian Ocean). Material examined: Punaruu Pass, Tahiti, 12 Jul. 1922, leg. W. A. Setchell & H. E. Parks, UC 261301 (type). Thallus caespitose, creeping, 400-600 µm in diameter, with thick stratified cell walls. Rhizoids abundant, up to 100 mm thick at the base, arising from nearly French Polynesian green algae 15 every cell. Individual cells 2-3 diameters long, deflexed. Basal cell up to 2 mm long, distinctly bent. Remarks: Growing in deep pools under Sargassum. This species has not been recollected in Tahiti since Setchell’s report. The conspecificity of C. basiretrorsa and Rhizoclonium grande was first suggested by H. Ohba, in an unpublished annotation on the holotype herbarium sheet dated 11 February 1998. This observation was confirmed by our own examination of Setchell’s material and comparison with R. grande from other Pacific Islands (e.g. N’Yeurt 1996, Rotuma). According to Silva et al. (1996: 793) and based on the authority of Dixit (1968: 13), R. grande and Chaetomorpha prostrata are synonyms of Cladophoropsis javanica (Kützing) P. Silva; however Hanyuda et al. (2003: 568, fig. 1) have shown from analysis of 18S rRNA sequences that R. grande is closely allied to Chaetomorpha. This is also confirmed by 28S rDNA sequences (unpublished data, F. Leliaert, pers. com.) where “R. grande” from the East African coast is closely related to Chaetomorpha crassa and C. spiralis Okamura. Leliaert (2004) also provides arguments for the distinction between Rhizoclonium and Cladophoropsis, on morphological grounds. As Setchell’s name is the earlier one, it takes precedence over Børgesen’s name for this entity (Article 11, Section 3 of the International Code of Botanical Nomenclature; Greuter et al., 2000) and R. grande is treated as a later synonym. The synonymy of Rhizoclonium profundum and R. grande was suggested by Dawson (1959: 7), and an examination of Setchell & Gardner’s illustrations and description of the Galapagos material would seem to confirm this. Chaetomorpha fibrosa Kützing, 1849: 377 (type locality: Baltic Sea). French Polynesia: Setchell, 1926: 72, pl. 8 figs 7-8. Material examined: Tongatapu, Tonga, Jul. 1874, leg. H. Moseley, BM 841401; Maraa, Tahiti, 28 Sep. 1928, leg. C. Crossland, BM 841379. Thallus forming crisp entangled masses over larger fixed algae. Segments 80-110 µm in diameter and 70-240 µm long. Cell walls stratified, about 12 µm thick. Remarks: This alga is little-reported in the literature, and of uncertain taxonomic status (Furnari et al., 1999). It mainly differs from C. linum by its smaller filament diameter. The French Polynesian material agrees with plants from Tonga in BM. *Chaetomorpha linum (O. F. Müller) Kützing, 1845: 204. India: Børgesen, 1935: 12, fig. 3; Pakistan: Nizamuddin & Begum, 1973: 15, fig. 62; Thailand: Egerod, 1974: 135, fig. 9; South Australia: Womersley, 1984: 176, pl. 13, fig. 2, figs 54D, 57A; Indian Ocean: Silva et al., 1996: 765; Belize: Littler & Littler, 1997: 93, fig. 126; Papua New Guinea: Littler & Littler, 2003: 198. (Figs 18-19) Basionym: Conferva linum O. F. Müller, 1778: 7, pl. 771(2) (type locality: Denmark). Misapplied name: Chaetomorpha basiretrorsa Setchell. French Polynesia: Setchell, 1926: 72. Material examined: Tahiti, Oct. 1909, leg. J. E. Tilden South Pacific Algae Nr. 21, BM 841428 (as Chaetomorpha antennina (Bory) Kützing). Thallus forming entangled, coiled unattached masses to 40 mm across (Fig. 18). Filaments 320-400 µm in diameter, of uniform thickness throughout, composed of cells 430-950 µm long, slightly incised at cross-walls (Fig. 19). Cell walls thin, 10 µm thick. Basal cells absent. Remarks: This species was not recollected since, and possibly interpreted by Setchell (1926) as a free, entangled form of C. basiretrorsa. However, the Tahitian 16 A. D. R. N’Yeurt & C. E. Payri material in BM agrees well with C. linum from other regional localities such as the Mariana Islands (Guam, BM 841419) or Tonga (BM 841384). Usually associated with high nutrient or polluted areas (Littler & Littler, 2003), a shift in habitat composition might explain this alga’s current disappearance. It mainly differs from the widespread Chaetomorpha crassa (C. Agardh) Kützing by its thinner cells walls, and rectangular cells never exceeding 500 µm in diameter in French Polynesian specimens. Chaetomorpha crassa usually attains diameters of up to 1000 µm, and has almost quadrate barrel-shaped cells (Littler & Littler, 2000: 314). Genus Rhizoclonium Kützing It has been proposed (Nienhuis, 1975; Kraft, 2000: 543) that there is only a single, continuously variable and widely distributed species, R. riparium (Roth) Harvey (1849). Furthermore, the only distinguishing character at present between Chaetomorpha and Rhizoclonium is the occasional production of morphologically distinct short laterals in the latter genus, and both genera could be congeneric, with Rhizoclonium being the earlier name. Moroever, Hanyuda et al. (2003: 570) and recent studies (F. Leliaert, pers. com.) have shown Rhizoclonium to be polyphyletic, forming a clade with Chaetomorpha and Cladophora species, and suggesting plural independent evolution of the genus morphology within the Cladophorales. For the time being, traditional taxonomic characters have been used to identify the two French Polynesian species of Rhizoclonium. Key to the French Polynesian species of Rhizoclonium 1a. Filament diameter 25-30 µm, often curled, apical cell swollen, found in estuaries. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. riparium 1b. Filament diameter 40-60 µm, not curled, apical cell not swollen, found in uppermost littoral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. africanum Rhizoclonium africanum Kützing, 1853: 21, pl. 67 fig. 2 (type locality: Senegambia, Africa). French Polynesia: Payri & N’Yeurt, 1997: 880, Payri et al., 2000: 70; Indian Ocean: Silva et al., 1996: 784; Lord Howe Island: Kraft, 2000: 542; Samoa: Skelton & South, 2002b: 10, pl. 7 figs 44-46. Material examined: Maraa, Tahiti, Sep. 1928, leg. C. Crossland, BM 840828; Taharaa, Tahiti, on cliff walls, 30 Mar. 1998, leg. A. D. R. N’Yeurt, UPF 703; Atituiti, Mangareva, Gambier, base of cliffs, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 920; Rikitea, Mangareva, Gambier, on breakwater, 20 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 892. Thallus light green and filamentous, to 10 cm long, forming entangled fleecy masses composed of straight, coarse filaments 45-60 µm in diameter. Individual cells rectangular, with thick cell walls. Apical cells of same diameter as main filaments. Multicellular rhizoids absent; some unicellular outgrowths occur very rarely. Remarks: Growing in the upper littoral, on moist cliff walls or surfaces periodically exposed to surf. Normally occurring above the high tide mark, this species can withstand considerable desiccation, often forming semi-dry skeins which can be confused with bryophytes. At times it can form “blooms” such as what occurred on beaches of Losin Island, Thailand, in June 2001 (K. Passfield, pers. com.). The French Polynesian species fall within the range of filament diameters of R. africanum, as described for Lord Howe Island material by Kraft (2000). French Polynesian green algae 17 Figs 16-22. 16. Chaetomorpha antennina (UPF RPS 202). Apical cell (a) and characteristically long basal cell (b). Scale = 100 µm. 17. Chaetomorpha basiretrorsa (UC 261301). Habit, showing deflexed cell and ventral rhizoid. Scale = 300 µm. 18. Chaetomorpha linum (BM 841428). Habit of pressed specimen collected by J. E. Tilden in Tahiti. Scale = 5 mm. 19. Chaetomorpha linum (BM 841428). Detail of rehydrated filament, showing slight incision at cross-wall (arrowhead). Scale = 20 µm. 20. Rhizoclonium riparium (UPF 2694). Habit, showing coiled filaments. Scale = 50 µm. 21. Rhizoclonium riparium (UPF 2694). Detail of typically swollen filament end. Scale = 25 µm. 22. Cladophora aokii (UPF RPS 211). Habit of thallus. Scale = 450 µm. 18 A. D. R. N’Yeurt & C. E. Payri *Rhizoclonium riparium (Roth) Harvey, 1849: pl. CCXXXVIII. Peru: Dawson et al., 1964: 11, pl. 8 figs B-E; Trinidad: Richardson, 1975: 85, pl. 18 fig. 2; Indian Ocean: Silva et al., 1996: 786; Belize: Littler & Littler, 1997: 95, fig. 131; 2000: 324, fig. on p. 325. (Figs 20-21) Basionym: Conferva riparia Roth, 1806: 216 (type locality: Norderney, East Frisian Islands, Germany; type lost according to Dawson et al., 1964). Material examined: Hakaui, Marquesas, 6 May 2002, leg. S. Sidolle, UPF 2694. Thallus filamentous, composed of unbranched, mostly curled filaments, 25-30 µm in diameter, forming yellow-green tangled masses. Short rhizoidal holdfast present. Basal cells 31-40 µm in diameter and 131-155 µm long. Cells of mid-thallus cylindrical, 63-123 µm long. Apical cells characteristically swollen. Remarks: Growing in brackish water at river mouths. Rhizoclonium riparium can be distinguished from R. africanum by its finer, darker coloured and more lubricous, somewhat curled filaments with swollen apical cells. It would represent the lower extreme of the range of width variations encountered in the genus, if we follow the opinion of Nienhuis, 1975. In the Marquesas, this species is used as the primary ingredient in a traditional remedy against joint pains (Conte & Payri, 2006). Genus Cladophora Kützing Seven species of this large polymorphic genus have been reported for French Polynesia by Payri & N’Yeurt (1997), but eight species (including five new records not listed in Payri & N’Yeurt, 1997) have been encountered and confirmed in collections. Setchell (1926) described two new species from Tahiti (C. leucobryoides and C. rhadina) but they have not been encountered since, although they have been reported from Hawaiian Islands by Abbott & Huisman (2004) who, after examining type material, put them in synonymy with C. socialis Kützing and C. seriacea (Hudson) Kützing, respectively. Similarly, C. inserta f. typica and f. ungulata Setchell were put in synonymy with C. vagabunda (Linnaeus) Hoek. Two other Cladophora species reported by Setchell [C. beneckii Moebius and C. insignis (Agardh) Kützing] were freshwater species not treated here. Because of the taxonomic difficulties associated with morphological plasticity of the species under various environmental conditions and geographic isolation, it is possible that several of the species reported earlier represent the same entities. In particular, the distinction between C. socialis Kützing (1849: 416) and C. patentiramea (Montagne) Kützing (both originally described from Tahiti) is based on the diameter of apical cells and the length of segments, which were found to be very variable and encompass the circumscription of both species in question (Kraft, 2000: 568). Members of this genus have recently been revised by van den Hoek (1963, 1982), van den Hoek & Chihara (2000), van den Hoek & Womersley (1984) and Leliaert (2004). Key to the French Polynesian species of Cladophora 1a. Thallus to 50 cm long, forming fleecy masses. . . . . . . . . . . . . C. patentiramea 1b. Thallus never exceeding 5 cm in length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Thallus erect, rigid or wiry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2b. Thallus forming short tufts or mats, not rigid or wiry. . . . . . . . . . . . . . . 3 3a. Filaments of mid-thallus 100 µm or less in diameter. . . . . . . . . . . . . . . . . . . . . 7 3b. Filaments of mid-thallus 150 µm or more in diameter . . . . . . . . . . . . . . . . . . . 4 French Polynesian green algae 5a. 5b. 7a. 7b. 19 4a. Filaments of mid-thallus 400-450 µm in diameter, branching irregular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. catenata 4b. Filaments of mid-thallus 150-250 µm in diameter, branching lateral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. herpestica Holdfast spreading and lacerate . . . . . . . . . . . . . . . . . . . . . . . . . . C. ohkuboana Holdfast rhizoidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Unbranched basal portion about half of thallus length, median segments 50-100 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. feredayoides 6b. Basal portion much shorter than thallus length, median segments 170350 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aokii Filaments delicate, main axes not reaching over 65 µm in diameter, branching irregular with non-percurrent axes, forming cushions . . . . . . . . . . . C. socialis Filaments about 100 µm in diameter, slippery, branching pseudodichotomous with percurrent axes, forming matted masses . . . . . . . . . . . . . . . . . C. seriacea *Cladophora aokii Yamada, 1925: 85-86, fig. III (type locality: Mao-pi T’ou, Taiwan). Taiwan: Sakai, 1964: 54; Japan & Russian Far East: van den Hoek & Chihara, 2000: 64, fig. 27A-C. (Fig. 22) Heterotypic synonym (given by van den Hoek & Chihara, 2000): Cladophora ryukyuensis Sakai et Yoshida in Yoshida, 1977: 71 (type locality: Loo Choo Islands, Ryukyus, Japan). Lord Howe Island: Kraft, 2000: 550, fig. 17A-C. Material examined: Rapa Island, Australs: Karapoo, Sargassum beds –28 m, 1 Nov. 2002, leg. J. L. Menou, UPF 1860; Tematapi Pt., 1. Nov. 2002, leg. J. L. Menou, UPF 1878; Komire Pt., 13 Nov. 2002, leg. J. L. Menou, UPF 2199; Ha’urei, 13 / 26 Nov. 2002, leg. J. L. Menou, UPF 2176, 2337; east oceanic slope, -68 m, 23 Nov. 2002, leg. J. L. Menou, UPF 2315, 2328; Tarakoi Islet, 5 / 6 Nov. 2002, leg. C. E. Payri, UPF 2011, 2034; Anarua Bay, 7 Nov. 2002, leg. C. E. Payri, UPF 2057; Aruroa Pt., 22 Nov. 2002, leg. A. D. R. N’Yeurt, UPF 2239, 2240; Tauna Islet, 27 Nov. 2002, leg. C. E. Payri, UPF 2351, 2352. Thallus olive green, 15-35 mm high; coarse and wiry, with secund or alternate branching at acute angles. Attached to the substratum via basal cells with stout, subparallel primary rhizoids, and secondary rhizoids from lower axial cells. Apical cells rounded, segments of mid-thallus 170-350 µm in diameter and 250-4000 µm long. Nodes with 2-4 axes occur near ultimate branches. Remarks: Found growing subtidally from -10 to as deep as –68 metres, this species is characterised by its coarse habit and thick branches. It ressembles C. dotyana Gilbert (1965) from Hawaii, but the latter is much coarser and has mature cells of the order of 500-700 µm in diameter, while C. aokii never exceeds 450 µm even in apical portions. According to van den Hoek & Chihara (2000: 65), Sakai (1964: 52) incorrectly interpreted the type of Cladophora fastigiata Harvey, on which C. ryukyuensis was based, but his description of the latter species conforms to that of the type of C. aokii Yamada, which has nomenclatural priority. *Cladophora catenata (Linnaeus) Kützing, 1843: 271. Papua New Guinea: Leliaert & Coppejans, 2003: 53, fig. 4. Basionym: Conferva catenata Linnaeus, 1753: 1166 (type locality: Bahamas according to van den Hoek, 1963: 12, 123). Misapplied name (given by Gilbert, 1962: 138): Cladophoropsis membranacea (C. Agardh) Børgesen. Hawaiian Islands: Egerod, 1952: 356, fig. 3. Heterotypic synonyms (given by Leliaert, 2004): Cladophoropsis luxurians Gilbert, 1962: 136, fig. 3A-B (type locality: shore of Molokai opposite Mokuhooniki Island, Hawaii). French Polynesia: Payri et al., 2000: 72. — Cladophora luxurians (Gilbert) Abbott & Huisman, 2003: 282; 2004: 77, fig. 22D (Hawaii). 20 A. D. R. N’Yeurt & C. E. Payri Material examined: Taiaro, 23 Sep. 1992, leg. R. Galzin, UPF 67; Haraiki, 14 Nov. 1996, leg. J. Orempuller, UPF 68, 554; Marokau, 22 Nov. 1996, leg. J. Orempuller, UPF 70; Nuku Hiva, Marquesas, 1997, leg. J. Orempuller, UPF 587. Thallus dark-green, up to 3 cm tall and forming small, tough mats on the reef top. Branching sparse to profuse and irregular, filaments 400-450 µm in diameter, turgid, with lateral branchlets secundly or irregularly issued from distal or proximal ends of mother cells, often curving downwards with frequent formation of secondary holdfasts from the crenulate tips of filaments. Formation of cross-walls at the base of the lateral branches markedly delayed. Remarks: Found on coral rubble, in the lagoons and passes of atolls of the Tuamotu archipelago and in the Marquesas. Abbott & Huisman (2003) transferred the Hawaiian species Cladophoropsis luxurians to Cladophora luxurians based on the presence of delayed cross-wall formations at the base of lateral branchlets, a feature inconsistent with the criteria for Cladophoropsis. Concurrently, Leliaert (2004: 122, 235), after examining relevant material, transferred Hawaiian records of Cladophoropsis membranacea and C. luxurians to Cladophora catenata as he found them to be indistinguishable from the latter species. *Cladophora feredayoides Kraft et Millar in Kraft, 2000: 556, fig. 19 (type locality: Erskotts Hole, Lord Howe Island, Australia). (Fig. 23) Material examined: Rapa Island, Australs: Rukuaga, base of cliffs –10 m, 30 Nov. 2002, leg. J. L. Menou, UPF 2429, 2690, 2691, RPS 75; Tupua’i Bay, 2 Dec. 2002, leg. J. L. Menou, UPF RPS 71. Thallus stiff and wiry, 30-40 mm high, growing solitarily from entangled basal rhizoids. Lower portion of plant unbranched for 10-20 mm. Segments of mid-thallus cylindrical, 50-100 µm in diameter and 630-2200 µm long; branching secund or alternate. Remarks: Growing from 8-10 m depth, this small stiff, wiry species is characterised by an unbranched lower portion of the thallus reaching at least half the length of the plant (Fig. 23b). The Rapa plants are in good agreement with the Lord Howe Island material described in Kraft (2000), and this represents the first published record for this species outside of the type locality. *Cladophora herpestica (Montagne) Kützing, 1849: 415. (Fig. 24) Basionym: Conferva herpestica Montagne, 1842: 15 (type locality: New Zealand). Homotypic synonym (given by Leliaert, 2004): Cladophoropsis herpestica (Montagne) Howe, 1914: 31. French Polynesia: Setchell, 1926: 77, pl. 8 figs 1-3, Payri & N’Yeurt, 1997: 881; South Australia: Womersley, 1984: 184, figs 58B, 59C; Indian Ocean: Silva et al., 1996: 791; Lord Howe Island: Kraft, 2000: 575, fig. 25E-F. Heterotypic synonym (given by Leliaert, 2004): Cladophoropsis adhaerens Gilbert, 1962: 136 (type locality: Hawaii). Material examined: Punaruu Pass, Tahiti, 11 Jul. 1922, leg. W. A. Setchell & H. E. Parks, BM 841145; Akananue Bay, Rapa Island, Australs, 2 Nov. 2002, leg. C. E. Payri, UPF 2686. Thallus forming compact, sediment-infiltrated dark green cushions to 15 mm thick and 200 mm wide, composed of stiff, subparallel often curved filaments 150-250 µm in diameter. Cross walls always present between two cells, but generally absent at the base of lateral branches, with a distal, single nonseptate rhizoid being issued from the basal poles of most cells. Branching lateral, distal to intercalary cells. Remarks: Leliaert (2004: 240) and Leliaert & Coppejans (2006) excluded Cladophoropsis herpestica (Montagne) Howe from Cladophoropsis, and placed it, French Polynesian green algae 21 along with Cladophoropsis zollingeri (Kützing) Reinbold, in synonymy with Cladophora herpestica. Abbott & Huisman (2004: 87) considered Cladophoropsis adhaerens Gilbert (1962: 136) from the Hawaiian Islands a synonym of C. membranacea, but Leliaert (2004: 244) found it to be indistinguishable from Cladophora herpestica (Montagne) Kützing. The species has recently not been found other than in the Austral islands in French Polynesia, although it occurs in neighbouring Cook Islands (N’Yeurt & Payri, in prep). Setchell’s (1926) record of this species from Tahiti (verified by us in BM) has not been recollected since from that locality. *Cladophora ohkuboana Holmes, 1896: 249, pl. 10 fig. 1 (type locality: Japan). Japan: Sakai, 1964: 15; Lord Howe Island: Kraft, 2000: 554, fig. 18 D-E; Japan & Russian Far-East: van den Hoek & Chihara, 2000: 84, figs 35, 36A-D. (Fig. 25) Material examined: Tauna Islet, Rapa Island, -20 m on coral rubble, 27 Nov. 2002, leg. C. E. Payri, UPF 2692, RPS 85. Thallus olive-green, rigid, up to 70 mm high. Holdfast spreading and lacerate, composed of a fibrous disc with radiating furcate, unseptated flattened rhizoids to 2 mm long. Branching alternate, opposite or secund, with laterals issued from main axis at narrow angles. Segments thick, cylindrical, 400-500 µm in diameter and 900-4000 µm long. Remarks: Superficially similar to C. dotyana Gilbert, this coarsest of Cladophora species is characterised by its thick cell dimensions and unique, spreading lacerate holdfast. Cladophora patentiramea (Montagne) Kützing, 1849: 416 pl. 60, fig. I. French Polynesia: Setchell, 1926: 74, pl. 9, Payri & N’Yeurt, 1997: 880, Payri et al., 2000: 68, Conte & Payri, 2002: 167, fig. 4; Viêt Nam: Dawson, 1954: 388, fig. 7e; Indian Ocean: Silva et al., 1996: 778; Lord Howe Island: Kraft, 2000: 566, fig. 23. Basionym: Conferva patentiramea Montagne, 1842: 15 (type locality: Tahiti). Material examined: Botanical Gardens Papeari, Tahiti, 1 Oct. 1995, 10 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 62, 60; Hotel Beachcomber, Faa’a, Tahiti, 13 Apr. 1997, 7 May 1997, leg. A. D. R. N’Yeurt, UPF 61, 64; Opunohu, Moorea, 1 Jul. 1997, leg. A. D. R. N’Yeurt, UPF 63; Tarakoi islet, Rapa, 5 Nov. 2002, leg. C. E. Payri, UPF 2693. Thallus to 50 cm long, light yellowish green in colour, forming fine fleecy tufts attached to the substratum via a small inconspicuous holdfast. Main axis to 250 µm in diameter, sparsely branched, bearing profuse, plumose, basally septate lateral branchlets in an irregularly opposite manner. Remarks: A common species growing in estuaries (Tahiti), or on the reef face 15-20 m depth (Rapa). Cladophora sericea (Hudson) Kützing, 1843: 264. Indian Ocean: Silva et al., 1996: 780; Hawaiian Islands: Abbott & Huisman, 2004: 77, fig. 22E. (Figs 26-27) Basionym: Conferva sericea Hudson, 1762: 485 (lectotype locality: Isle of Sheppey, Kent, England). Heterotypic synonym (given by Abbott & Huisman, 2004): Cladophora rhadina Setchell, 1926: 76, pl. 10 fig. 1 (type locality: Afaahiti District, Tahiti). Material examined: Port Phaeton, Afaahiti, Tahiti, 24 Jun. 1922, leg. W. A. Setchell & H. E. Parks, !UC 261235. Thallus 30-40 cm high, with matted holdfasts, forming pale green silky, slippery floating masses. Branching dense and pseudodichotomous, with both long and short laterals of mixed ages. Main filaments about 100 µm in diameter, 22 A. D. R. N’Yeurt & C. E. Payri Figs 23-30. 23. Cladophora feredayoides (UPF RPS 71). (a) Habit of stiff, erect thallus. (b) Lower holdfast of extensive, unbranched basal portion. Scale = 80 µm. 24. Cladophora herpestica (UPF RPS 42). Detail showing production of basal, non-septate rhizoid (arrowhead) from filament cell. Scale = 200 µm. 25. Cladophora ohkuboana (UPF RPS 85). Habit of distal portion of thallus. Scale = 500 µm. 26. Cladophora sericea (UC 261235). Habit of dried Holotype collected by Setchell, and Parks from Afaahiti, Tahiti. Note annotation by I. A. Abbott on leftern side. Scale = 20 mm. 27. Cladophora sericea (UC 261235). Detail of rehydrated portion of thallus showing pseudodichotomous branching. Scale = 50 µm. 28. Cladophora socialis (UC 233473). Habit of dried, felt-like material collected from Tahiti in 1909 by Josephine Tilden. Scale = 15 mm. 29. Cladophora socialis (UC 233473). Detail of rehydrated portion of thallus, showing abundant epiphytic diatoms (arrowhead) imparting greyish colour to plant. Scale = 50 µm. 30. Cladophora socialis (UPF 3003). Detail of portion of mid-thallus, showing irregular branching. Scale = 50 µm. French Polynesian green algae 23 sharply distinct from secondary branches, with segments up to 400 µm long. Apical cells 12-16 µm in diameter and up to 280 µm long; apices rounded. Secondary branches often in 3s or 4s in lower nodes. Remarks: Floating near causeway. The synonymy of Cladophora rhadina and C. sericea was proposed by I. A. Abbott (October 1990, annotation on herbarium sheet of type material, Fig. 26) and reiterated in Abbott & Huisman (2004: 77). The Tahitian material from Afaahiti District in UC also contains fragments of the red alga Sarconema filiforme (Sonder) Kylin, which in recent time has been reported from neighbouring Papeari District (Payri et al., 2000). Cladophora socialis Kützing, 1849: 416 (type locality: Tahiti). French Polynesia: Setchell, 1926: 74; Easter Island: Børgesen, 1924: 249, fig. 2a-c; Marshall Islands: Dawson, 1957: 103, fig. 4B; Pakistan: Nizamuddin & Begum, 1973: 9, figs 44-49; Thailand: Egerod, 1975: 137, figs 17-21; Somalia: Sartoni, 1986: 361, fig. 4A; 1992: 302, fig. 6A-B; South Africa: Leliaert & Coppejans, 2003: 51, fig. 3; Hawaiian Islands: Abbott & Huisman, 2004: 78, fig. 23A-C. (Figs 28-30) Heterotypic synonym (given by Abbott & Huisman, 2004): Cladophora leucobryoides Setchell, 1926: 74 (type locality: Tahiti). Material examined: Tahiti, October 1909, leg. J. E. Tilden, !UC 233473; Punaauia, 3 Apr. 2005, leg. A. D. R. N’Yeurt & K. Pouira, UPF 3003. Thallus forming pale to dark green dense felty cushions about 10 mm in diameter and 15-20 mm thick (Fig. 28). Branching irregular. Basal filaments to 52 µm in diameter with segments up to 180 µm long (Fig. 30). Apical cells to 20 µm in diameter, obtuse. Secundly branched one-cell laterals present near apices. Axes slender and intertwined, with cylindrical cells. Uniseriate rhizoids common from basal poles of cells (even in distal parts of the thallus), ending in disclike attachment pads. Distinctly felt-like when dry. Remarks: Growing as dark green moss-like masses, in small inlets (Tautira) and on semi-exposed rocks on the foreshore (Punaauia). The synonymy of Cladophora leucobryoides and C. socialis was proposed by I. A. Abbott (October 1990, annotation on herbarium sheet of type material) and reiterated in Abbott & Huisman (2004: 78). The plants often appear grey due to the presence of abundant epiphytic diatoms (Fig. 29). Family Siphonocladaceae Schmitz Genus Boodlea G. Murray et De Toni in Murray Boodlea, a genus of some six species, differs from closely related Phyllodictyon by its spongy three dimensional network, lacking a stalk and growing directly on the substratum (Kraft & Wynne, 1996). Species demarcations within the genus are poorly defined, and there exists wide variations in single populations depending on environmental conditions. Three species are reported from French Polynesia in Payri & N’Yeurt (1997), with only a single species confirmed in collections. Boodlea composita (Harvey) Brand, 1904: 187, pl. 6 figs 28-35. French Polynesia: Taylor, 1973: 38 (Bora Bora, Huahine); Hawaiian Islands: Egerod, 1952: 362, pl. 32a, fig. 6a, Abbott & Huisman, 2004: 85, figs 26A-B; Viêt Nam: Dawson, 1954: 390, fig. 9c-d; New Caledonia: Valet, 1968: 38, pl. 12 fig. 2; Thailand: Egerod, 1975: 50, fig. 19; Somalia: Sartoni, 1986: 306, fig. 7C; Indian Ocean: Silva et al., 1996: 789; Papua New Guinea: Leliaert et al., 1998: 184, figs 14-20; Lord Howe Island: Kraft, 24 A. D. R. N’Yeurt & C. E. Payri 2000: 569, fig. 24 A-C; Fiji: N’Yeurt, 2001: 703, Littler & Littler, 2003: 200; Samoa: Skelton & South, 2002b: 10, pl. 7 figs 44-46; Rodrigues Island: Coppejans et al., 2004: 2970. (Figs 31-34) Basionym: Conferva composita Harvey, 1834: 157 (type locality: Mauritius). Misapplied name: Boodlea kaeneana Brand, 1904: 190, pl. 6 figs 36-39; French Polynesia: Setchell, 1926: 77, pl. 7 fig. 5, Payri & N’Yeurt, 1997: 880, Payri et al., 2000: 70. Heterotypic synonyms (given by Leliaert, 2004): Boodlea coacta (Dickie) Murray et De Toni in Murray, 1889: 245. Rotuma: N’Yeurt, 1996: 368, fig. 19. — Boodlea montagnei (Harvey ex J.E. Gray) Egerod, 1952: 332. Cook Islands: Dickie, 1877: 33; Somalia: Sartoni, 1986: 307, fig. 7D; Hawaiian Islands: Abbott & Huisman, 2004: 86, fig. 27. Material examined: Tahiti, 3 Apr. 1911, leg. J. E. Tilden, BM 701624; Maraa, Tahiti, Sep. 1928, leg. C. Crossland, BM 841661; Taharaa, Tahiti, 14 Apr. 1997, leg. C. E. Payri & A. D. R. N’Yeurt, UPF 9; Taapuna, Tahiti, 7 May. 1997, leg. A. D. R. N’Yeurt, UPF 8; Tiahura, Moorea, 1 Jul. 1997, leg. A. D. R. N’Yeurt, UPF 383; Atituiti, Mangareva, Gambier, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 917; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2522; Tauna Islet, Rapa, 27 Nov. 2002, leg. C. E. Payri, UPF 2368; Rapa Iti, 28 Nov. 2002, leg. J. L. Menou, UPF 2413; Punaauia, Tahiti, 15 Apr. 2005, leg. A. D. R. N’Yeurt, and V. Pouira, UPF L15-04-05. Thallus shiny light green, spongy and crisp, forming unattached masses to 13 cm in diameter. Main filaments about 240 µm in diameter, composed of cells 500-560 mm long. Thallus highly polymorphic; arrangement of older branches are mostly three-dimensional and random (Fig. 31), while younger portions of thalli range from semi-plumose (Fig. 32) to regularly organised, Phyllodictyon-like twodimensional blades (Fig. 33). Branching irregularly lateral, the terminal branchlets 35-65 mm in diameter, with frequent secondary attachments between branches via terminal, crenellated sucker-like pads on unmodified segments (type-1 tenacular cells in Leliaert, 2004: 204; Fig. 34). Remarks: Setchell (1926: 77) distinguished Tahitian and Hawaiian Boodlea kaeneana Brand by its single or unilateral second-order branchlets, contrasting to the regularly opposite branching of the Mauritius species B. composita (Børgesen, 1940: 21, fig. 6). The French Polynesian plants were found to exhibit variable regular, irregular or unilateral branching. Thus it is reasonable to assume Tahitian B. kaeneana records a misapplied name for B. composita, a species previously reported from the Society Islands by Taylor (1973). Furthermore, the Hawaiian type material of B. kaeneana was found conspecific with Cladophoropsis membranacea (Leliaert, 2004: 120). The genus Boodlea (along with Phyllodictyon and several others) form a close cluster with Cladophoropsis in the Cladophorophyceae based on recent phylogenetic studies (Kooistra et al., 1993; Wysor, 2002; Leliaert et al., 2003) and a single, highly variable species (Boodlea composita) may need to be recognized (Leliaert, 2004). We have ourselves examined the type specimens of Conferva composita, Cladophora coacta, Microdictyon montagnei, Boodlea paradoxa and Boodlea siamensis in BM, and have to agree with Leliaert’s encompassing view of this highly polymorphic genus. Genus Cladophoropsis Børgesen amend. Papenfuss The taxonomy of this polyphyletic genus has most recently been revised by Leliaert (2004) and Leliaert & Coppejans (2006). French Polynesian green algae 25 Cladophoropsis sundanensis Reinbold, 1905: 147 (lectotype locality: Kangean, Indonesia according to Leliaert, 2004: 127). French Polynesia: Setchell, 1926: 77; W.R. Taylor, 1973: 38 (Makatea); Payri & N’Yeurt, 1997: 881, Payri et al., 2000: 72; Indian Ocean: Silva et al., 1996: 793; Rotuma: N’Yeurt, 1996: 370, fig. 11; Lord Howe Island: Kraft, 2000: 573, fig. 25A-D; South Africa: Leliaert et al., 2001: 452, figs 6-8; Fiji: N’Yeurt, 2001: 706, fig. 16; Leliaert, 2004: 127, figs 3L-M, 13, 14. Heterotypic synonym (given by Leliaert, 2004): Cladophoropsis carolinensis Trono, 1971: 48 (type locality: Truk, Micronesia). Material examined: Tikehau, lagoon, 7 Nov. 1995, leg. A. D. R. N’Yeurt, UPF S154; Motu Taraururoa, Gambier, fringing reef on ocean side, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 851; Motu Totegegie, Gambier, fringing reef, 22 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 796; north plateau, -40 m, Rapa Island, Australs, 25 Nov. 2002, leg. J. L. Menou, UPF 2332. Plants tufted, greenish-brown, forming sediment-infiltrated flaccid cushions up to 15 mm high with filaments 60-176 µm in diameter; loosely branched with branchlets at 300-500 µm intervals along the main axis. Branches non-septate and entangled at the base, secund or irregular and projecting from the distal end of the primary axial cells, with a rhizoid arising from the base of most cells immediately above a lateral in open connection with the cell immediately below. Remarks: Leliaert (2004: 127), after examining relevant type material, found no basis to distinguish Cladophoropsis carolinensis from C. sundanensis. Recent molecular phylogeographic studies on Cladophoropsis membranacea (Kooistra, 1992; van der Strate et al., 2002) revealed the presence of cryptic species, notably separate clades for Atlantic, Caribbean and Pacific (Japanese, Hawaiian) material. In the same studies, C. sundanensis was found to be a sister species to the C. membranacea clade. Genus Dictyosphaeria Decaisne ex Endlicher Key to the French Polynesian species of Dictyosphaeria 1a. Thallus hollow, intracellular spines absent . . . . . . . . . . . . . . . . . . D. cavernosa 1b. Thallus solid, intracellular spines present . . . . . . . . . . . . . . . . . . . . D. versluysii Dictyosphaeria cavernosa (Forsskål) Børgesen, 1932: 2, pl. 1 fig. 1. French Polynesia: Taylor, 1973: 38 (Bora Bora); Payri & N’Yeurt, 1997: 881, Payri et al., 2000: 76; Hawaiian Islands: Egerod, 1952: 350, figs 1b-f, 2f-g; Abbott & Huisman, 2004: 89, fig. 29A; Viêt Nam: Dawson, 1954: 388, fig. 8i; Indian Ocean: Silva et al., 1996: 794; Rotuma: N’Yeurt, 1996: 371, fig. 12; Papua New Guinea: Leliaert et al., 1998: 188, figs 30-33; Lord Howe Island: Kraft, 2000: 578, fig. 27 A-B; Fiji: N’Yeurt, 2001: 707; Littler & Littler, 2003: 202; South Africa: De Clerck et al., 2005: 54, fig. 23. (Fig. 35) Basionym: Ulva cavernosa Forsskål, 1775: 187 (syntype localities: “Gomfodae” (Al-Qunfidha), Saudi Arabia; Mokha, Yemen). Heterotypic synonym (given by Silva et al., 1996): Dictyosphaeria favulosa (C. Agardh) Decaisne ex Endlicher, 1843: 18. French Polynesia: Setchell, 1926: 78. Material examined: Reka Reka, 9 Nov. 1996, leg. J. Orempuller, UPF 80; Haraiki, 15 Nov. 1996, leg. J. Orempuller, UPF 79, 707; Hotel Intercontinental, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2590; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2601; Tarakoi islet, Rapa, shelves with Sargassum, 6 Nov. 2002, leg. C. E. Payri, UPF 2036; Akaeke Pt., Rapa, dead patch reef, 7 Nov. 2002, leg. C. E. Payri, UPF 2055; Tiahura, Moorea, 1 Oct. 2004, leg. A. D. R. N’Yeurt, UPF 2955. 26 A. D. R. N’Yeurt & C. E. Payri Thallus 8-13 cm in diameter; shiny light green, sessile, sometimes spherical and often irregularly lobed. Internal structure hollow, the walls one-cell thick, with angular or polygonal cells clearly seen with the naked eye. Intracellular spines absent. Thallus lightly attached to substratum via small rhizoids. Remarks: Commonly found growing on coral and other hard substrata in the lagoon of atolls, where it can reach large sizes. Dictyosphaeria versluysii Weber-van Bosse, 1905: 144 (“versluysi”) (syntype localities: “Plusieurs récifs dans l’Archipel Malaisien”). French Polynesia: Payri et al., 2000: 76; Hawaiian Islands: Egerod, 1952: 351, 354-355, figs 1a, 2h-k; Abbott & Huisman, 2004: 89, fig. 29B; New Caledonia: Valet, 1966a: 256, figs 1-2; Indian Ocean: Silva et al., 1996: 796; Papua New Guinea: Leliaert et al., 1998: 190, figs 37-39; Fiji: N’Yeurt, 2001: 707, Littler & Littler, 2003: 204; South Africa: De Clerck et al., 2005: 56, fig. 24. (Fig. 36) Heterotypic synonym (given by Valet, 1966a): Dictyosphaeria australis Setchell, 1926: 79, pl. 8 figs 9, 10 (type locality: Arue, Tahiti). French Polynesia: Payri & N’Yeurt, 1997: 881. Material examined: Maraa, Tahiti, 24 Sep. 1928, leg. C. Crossland, BM 841316; Taharaa, Tahiti 14 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 77; Taapuna, Tahiti, 7 May 1997, leg. A. D. R. N’Yeurt, UPF 78; Punaauia, Tahiti, 10 May 1997, leg. A. D. R. N’Yeurt, UPF 76; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2601; Atituiti, Mangareva, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 916. Thallus to 4 cm in diameter, shiny dark green, sessile and cartilaginous, with angular or polygonal cells clearly seen with the naked eye. Internal structure solid throughout, with sparse intracellular spines 60-88 µm long. Attached to the substratum via sparse basal rhizoids. Remarks: Forming tightly adhering, solid button-like crusts on coral debris. Valet (1966a) found that characters previously used to separate solid species of Dictyosphaeria of the versluysii group were overlapping and unreliable; D. australis Setchell fell within that category. Genus Phyllodictyon J.E. Gray Phyllodictyon anastomosans (Harvey) Kraft et Wynne, 1996: 139, figs 16-25. French Polynesia: Payri et al., 2000: 74; Papua New Guinea: Leliaert et al., 1998: 186, figs 23-24; Oman: Wynne, 2001: 368, fig. 34; Fiji: Littler & Littler, 2003: 200; Hawaiian Islands: Abbott & Huisman, 2004: 63, fig. 16A-B. Basionym: Cladophora? anastomosans Harvey, 1859: pl. CI (type locality: Fremantle, Western Australia). Heterotypic synonyms (given by Kraft & Wynne, 1996): Struvea anastomosans (Harvey) Piccone et Grunow ex Piccone, 1884: 20. West Indies: Børgesen, 1913: 54, fig. 39; Mauritius: Børgesen, 1952: 7, fig. 3; Hawaiian Islands: Egerod, 1952: 359, pl. 31 fig. 4; Viêt Nam: Dawson, 1954: 390, fig. 8g; Thailand: Egerod, 1975: 50, fig. 15a-b; Somalia: Sartoni, 1992: 317, fig. 12B-C; Papua New Guinea: Coppejans et al., 1995: 96, fig. 38; Maldives: Wynne, 1993: 20; 1995: 332, fig. 86; Indian Ocean: Silva et al., 1996: 798; Rotuma: N’Yeurt, 1996: 369, figs 21-22. — Struvea delicatula Kützing, 1866: 1, pl. 2 figs e-g (type locality: New Caledonia). Japan: Okamura, 1908 (1907-1909): 203, pl. 40 figs 9-12; Segawa, 1938: 135, fig. 3; Ryukyu Islands: Yamada, 1934: 46, fig. 10; New Caledonia: Valet, 1968: 37. Material examined: Punaauia PK 18, Tahiti, 28 Feb. 1998, leg. A. D. R. N’Yeurt, UPF 613. French Polynesian green algae 27 Figs 31-38. 31. Boodlea composita (UPF L15-04-05). Habit of randomly organised, older part of thallus. Scale = 250 µm. 32. Boodlea composita (UPF L15-04-05). Habit of younger part of thallus. Scale = 100 µm. 33. Boodlea composita (UPF L15-04-05). Habit of very young blade-like part of thallus. Scale = 100 µm. 34. Boodlea composita (UPF L15-04-05). Detail of crenellated attachment pad (arrowhead) between adjacent branchlets. Scale = 100 µm. 35. Dictyosphaeria cavernosa (UPF 2036). Habit of flattened-out thallus. Scale = 10 mm. 36. Dictyosphaeria versluysii (UPF 2601). Habit of solid thallus. Scale = 5 mm. 37. Siphonocladus tropicus (UPF 2091). Habit of freshly-collected plant. Scale = 2 mm. 38. Struveopsis sp. (UPF RPS 20). Habit of thallus, showing irregular branching and portions with segregative cell division (arrowheads). Also shown is the lack of cross wall between the main axis and side branchlets (small arrow). Scale = 500 µm. Thallus to 30 mm high, dark to light green, composed of a segmented, hollow stalk to 800 µm wide bearing an irregularly shaped flat blade to 10 mm wide and 15 mm long. The blade is composed of a network of segmented, anastomosing filaments 60-100 µm in diameter repeatedly branched in a mostly opposite manner along upper parts of the main axis. Tenacular cells 28 A. D. R. N’Yeurt & C. E. Payri predominantly composed of small modified cells produced distally on terminal segments (type-3 cells in Leliaert, 2004: 104). Remarks: Growing usually solitarily, among algal mats on coral heads in the lagoon. Leliaert (2004) advocated the merging of Phyllodictyon and other genera such as Boodlea under Cladophoropsis. Indeed, some ecomorphs of French Polynesian Boodlea composita are strikingly alike Phyllodictyon anastomosans as circumscribed here, and the validity of this and other related genera might change in future. Genus Siphonocladus F. Schmitz *Siphonocladus tropicus (P. et H. Crouan) J. Agardh, 1887: 105. Somalia: Sartoni, 1992: 315, fig. 12A; Indian Ocean: Silva et al., 1996: 797; Hawaiian Islands: Abbott & Huisman, 2004: 90, fig. 30; Wallis: N’Yeurt & Payri, 2004: 372. (Fig. 37) Basionym: Apjohnia tropica P. et H. Crouan in Schramm et Mazé, 1865: 47 (syntype localities: Guadeloupe, West Indies). Material examined: Rarapai Islet, Rapa, 12 Nov. 2002, leg. J. L. Menou, UPF 2170; between Matarepe and Takaraotaranga Points, Rapa, 9 Nov. 2002, leg. C. E. Payri, UPF 2091. Thallus to 40 mm high, composed of erect subcylindrical vesicles 1-3 mm in diameter, with prominent basal annular constrictions, and well developed, nonseptate basal rhizoids. Each axis consisting of a pseudoparenchymatous filament of appressed, polygonal cells 200-500 µm in diameter. Blunt-ended lateral branchlets in irregularly radial whorls, to 10 mm long, formed from rounded-up protoplast of main axis or previous branch order. Remarks: growing between 12 and 32 m on hard substrata, so far in French Polynesia only reported from the southern Australs (Rapa). Genus Struveopsis Rhyne et H. Robinson *Struveopsis sp. (Fig. 38) Material examined: Marotiri, Australs, 6 Nov. 2002, leg. I. R. D., UPF 2106, RPS 20. Thallus plumose, grass-green, rugose when dry, 20-30 mm high and 5-6 mm broad, including a non-annulated, terete stipe 9-10 mm long and 500-600 µm in diameter, attached to the substratum via cylindrical, blunt-ended rhizoids. Central axis 250-300 µm in diameter, composed of cylindrical to subquadrate cells 300-600 µm long, with thick stratified cell walls. Side branchlets 100-220 µm in diameter are issued alternately, opposite or unilaterally at irregular intervals along the main axis, and are at their base in continuous connection with the parent cell, lacking a cross wall. Some parts of side branchlets apparently with segregative cell division, most portions of thallus with intercalary, asynchronous cell division. Remarks: Dredged from 52 m depth. The genus Struveopsis is intermediate between Cladophoropsis, Struvea, and Phyllodictyon and is conspecific with the genus Pseudostruvea Egerod (1975: 47) (Kraft & Wynne, 1996: 140). The genus could be considered a phenodeme in the highly polymorphic Cladophoropsis complex (Leliaert, 2004: 167). The Marotiri plant clearly belongs in the Siphonocladales, and the presence in some parts of the thallus of segregative cell division would suggest placement in Struvea, following the interpretation of Kraft & Wynne (1996). However, Leliaert (2004) revised this interpretation, finding both segregative and intercalary division in many members of the Cladophorales. The long naked stipe and erect, percurrent main axis of the plant distinguishes it from Cladophoropsis. The Marotiri plant differs from Cladophora (sensu stricto) French Polynesian green algae 29 based on the lateral branches with distinct delayed cross-wall formation, and from Struvea by the total lack of tenacular adhesion between adjacent branchlets (absence of a reticulum), and the asynchronous, partially unilateral, Cladophoropsis-like side branchlets with delayed cross wall formation, features characteristic of Struveopsis (F. Leliaert, pers. com.). Family Valoniaceae Nägeli Genus Valonia C. Agardh, 1822: 429 Key to the French Polynesian species of Valonia 1a. Thallus forming mostly erect clumps . . . . . . . . . . . . . . . . . . . . . . . . V. fastigiata 1b. Thallus forming encrusting mats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Vesicles ovoid to ocellate, solitary or in colonies, mostly unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. macrophysa 2b. Vesicles cylindrical to clavate, much branched . . . . . . . . . V. aegagropila Valonia aegagropila C. Agardh, 1823: 429 (lectotype locality: Venezia, Italy according to Egerod, 1952: 348). French Polynesia: Payri & N’Yeurt, 1997: 881, Payri et al., 2000: 78; Hawaiian Islands: Egerod, 1952: 348, pl. 29b; Abbott & Huisman, 2004: 92, fig. 31A; Indian Ocean: Silva et al., 1996: 801; Rotuma: N’Yeurt, 1996: 372, fig. 15a-b; Papua New Guinea: Leliaert et al., 1998: 192, figs 40-41; Fiji: N’Yeurt, 2001: 708, Littler & Littler, 2003: 206. Material examined: Mokoto, Mangareva, 27 Sep. 1997, leg. J. Starmer, UPF 623; Hotel Intercontinental, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2589. Thallus encrusting, light yellowish to olive-green, composed of cylindrical to clavate vesicles 3-13 mm long and 1.5-2 mm broad, subdichotomously branched from the sides or the ends of the cells. Young plants attached to each other, the older ones more or less free. Remarks: Forming extensive mats at the base of coral heads and on flat surfaces, in the lagoon of high islands. Valonia fastigiata Harvey ex J. Agardh, 1887: 101, pl. I fig. 5 (syntype localities: Sri Lanka; Tonga). French Polynesia: Setchell, 1926: 80, Payri & N’Yeurt, 1997: 881; Payri et al., 2000: 78; Somalia: Sartoni, 1992: 321, fig. 14C; Papua New Guinea: Coppejans et al., 1995: 98, fig. 39, Leliaert et al., 1998: 192, figs 42-44; Indian Ocean: Silva et al., 1996: 801; Samoa: Skelton & South, 2002a: 162, fig. 24A; Fiji: Littler & Littler, 2003: 206. (Fig. 39) Basionym: Conferva utricularis Roth, 1797: 160, pl. I fig. 1 (type locality: Mediterranean Sea). Material examined: Tahiti, Sep. 1875, leg. H. Moseley, BM 841146, 841343; Punaruu, Tahiti, 12 Jul. 1922, leg. W. A. Setchell & H. E. Parks, BM 841322, 841342; Taapuna, Tahiti, 7 May 1996, leg. C. E. Payri, UPF 140, 141; Rurutu, Australs, 18 Aug. 2000, leg. C. E. Payri, UPF 730; Matira, Bora Bora, 15/18 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2491, 2537. Thallus dark olive green, forming erect clumps or cushions to 3 cm high and 10 cm across. Individual vesicles elongate-ovoid, hollow, irregularly branched, to 10 mm long and 5 mm wide, loosely attached to each other. Remarks: Growing as clumps between branches of coral and crevices in hard substratum, in the lagoon and on the reef flat. 30 A. D. R. N’Yeurt & C. E. Payri Valonia macrophysa Kützing, 1843: 307 (type locality: Lessina, Croatia). French Polynesia: Payri et al., 2000: 80; Indian Ocean: Silva et al., 1996: 802; Belize: Littler & Littler, 1997: 91, fig. 121; Fiji: Littler & Littler, 2003: 206. Material examined: Tiahura, Moorea, 3 Nov. 2004, leg. A. D. R. N’Yeurt, in Herb. UPF. Thallus to 10 cm across and 5 cm thick, shiny dark olive green, composed of irregularly ovoid to ocellate, bubblelike macroscopic cells 5-20 mm in diameter and 10-40 mm long, mostly unbranched. Thallus attached to the substratum via rhizoids issued from basal cells. Remarks: Forming tightly adhering mats on flat coral, usually Favites sp. in shallow lagoon waters. Genus Valoniopsis Børgesen Valoniopsis pachynema (G. Martens) Børgesen, 1934: 10-16, figs 1a-f, 2. French Polynesia: Abbott, 1986: 163, figs 3-5; Micronesia: Dawson, 1957: 102, fig. 2; Fiji: N’Yeurt, 2001: 709, fig. 31; Hawaiian Islands: Abbott & Huisman, 2004: 93, fig. 32A. Basionym: Bryopsis pachynema G. Martens, 1868: 24, 62-63, pl. IV fig. 2 (syntype localities: Benkulen (Bengkulu) and Pulau Tikus, near Bengkulu, Sumatra, Indonesia). Material examined: Lake Fauna Nui, Huahine, c. 2003, leg. C. E. Payri, UPF 3967, 3968. Thallus forming cushions up to 5-8 cm across and 3 cm high, composed of coenocytic, erect or arcuate, elongated cylindrical vesicles 1-1.5 mm in diameter. Branching irregular, with up to 5 basally-septated branch segments in an umbellate apical cluster on parent segment. Basal rhizoids irregularly branched and tapered, lacking annular constrictions. Remarks: Growing on the lagoon floor, 0.3 m depth. This unique ball-like form of Valoniopsis, illustrated in Abbott (1986), was found growing about 2 miles from the barrier reef, probably having been washed up from its normal habitat and assuming its unusual habit owing to environmental factors inherent to the semiclosed internal lagoon. Genus Ventricaria J.L. Olsen et J.A. West Ventricaria ventricosa (J. Agardh) J.L. Olsen et J.A. West, 1988: 104, fig. 11. French Polynesia: Payri & N’Yeurt, 1997: 881; Payri et al., 2000: 80; Somalia: Sartoni, 1992: 323, fig. 14E; Indian Ocean: Silva et al., 1996: 800; Rotuma: N’Yeurt, 1996: 372, fig. 23; Belize: Littler & Littler, 1997: 89, fig. 119; Fiji: N’Yeurt, 2001: 708, Littler & Littler, 2003: 204; Hawaiian Islands: Abbott & Huisman, 2004: 94, fig. 32B. (Fig 40) Basionym: Valonia ventricosa J. Agardh, 1887: 96 (syntype localities: St. Croix, Virgin Island; Guadeloupe). French Polynesia: Setchell, 1926: 79, Payri & Meinesz, 1985a: 510; Hawaiian Islands: Egerod, 1952: 347, pl. 29a; Viêt Nam: Dawson, 1954: 388, fig. 8e; Marshall Islands: Dawson, 1956: 28, 1957: 101; Philippines: Trono, 1986: 213, fig. 7. Material examined: Opunohu Bay, Moorea, 30 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 142; Tupai, 3 Jul. 2002, leg. C. Vermenot, UPF 2488; Matira, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2494. French Polynesian green algae 31 Thallus to 5 cm in height, shiny dark green, consisting of a thin-walled, turgid, fluid-filled subspherical to spherical or pyriform marble-like single cell. Attached to the substratum via minute basal rhizoids. The cell does not rupture if the cell wall is punctured. Remarks: Common in the lagoon, growing in interstices on coral micro-atolls or epiphytic on algae such as Galaxaura spp.; or on coral debris to a depth of 20 meters on the outer reef slope (Society). Large, deep-water specimens are often covered with epiphytic crustose coralline algae. Order Bryopsidales Schaffner Family Bryopsidaceae Bory de Saint-Vincent Genus Bryopsis J.V. Lamouroux Key to the French Polynesian species of Bryopsis 1a. Thallus light translucent green, branches plumose, not curved. . B. plumosa 1b. Thallus dark green, branches curved, appearing uniseriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pennata var. secunda Bryopsis pennata J.V. Lamouroux var. secunda (Harvey) Collins et Hervey, 1917: 62. French Polynesia: Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 82; Brazil: Mitchell et al., 1979: 109, pl. 1 figs 6-8; Indian Ocean: Silva et al., 1996: 807; Belize: Littler & Littler, 1997: 97, fig. 134; Fiji: N’Yeurt, 2001: 709, fig. 30, Littler & Littler, 2003: 208; Samoa: Skelton & South, 2002a: 163, fig. 24E; Hawaiian Islands: Abbott & Huisman, 2004: 98, fig. 33C. Basionym: Bryopsis plumosa (Hudson) C. Agardh var. secunda Harvey, 1858a: 31, plate XLV.A. figs 1-3 (syntype localities: Key West & Sand Key, Florida, U.S.A.). Heterotypic synonym (given by Silva et al., 1996: 807): Bryopsis harveyana J. Agardh, 1887: 82 (syntype localities: Florida, U.S.A.; Tonga). French Polynesia: Setchell, 1926: 80; Payri & N’Yeurt, 1997: 882. Material examined: Taharaa, Tahiti, 17 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 701617; Hikueru, 17 Nov. 1996, leg. J. Orempuller, UPF 12; Hotel Beachcomber, Faa’a, Tahiti, estuary, leg. A. D. R. N’Yeurt, 4 May 1997, UPF 14, 15; 5 Jun. 1997, UPF 10, 15; Pirae, Tahiti, 15 Apr. 1998, leg. C. E. Payri, UPF 718; Taharaa, Tahiti, 22 Apr. 1998, leg. A. D. R. N’Yeurt, UPF 719. Thallus dark iridescent green; in compact clumps 12-60 mm high and 120 mm broad; main axis 300-800 µm in diameter and unbranched, with slight upward curvature. Secondary branches cylindrical to clavate, up to 4 mm long and 100 µm broad, with rounded apex and slight constriction at the base. Branchlets occurring in an offset pair of lateral rows on one side of the primary axis, giving a uniseriate appearance to the thallus. Secondary branchlets typically longer in middle of axis, imparting a renoid curvature to the younger blades. Remarks: Growing in clumps, intertidally near the reef crest or in dense clumps on pebbles and rocks in the calm, sediment-rich waters of estuaries (Tahiti). Bryopsis plumosa (Hudson) C. Agardh, 1823: 448. French Polynesia: Payri et al., 2000: 82; Peru: Dawson et al., 1964: 14, pl. 11 fig. D; Kermarrec, 1974: 21, pl. 1 fig. A; Rietema, 1975: 8-24, pls 1-9; Florida: Woelkerling, 1976: 86, figs 9-14; South Australia: Womersley, 1984: 282, figs 96C, 97A; Korea: Lee et al., 1991: 24, figs 1A-E, 4A; Indian Ocean: Silva et al., 1996: 808; Rotuma: N’Yeurt, 1996: 374, figs 49, 60; 32 A. D. R. N’Yeurt & C. E. Payri Belize: Littler & Littler, 1997: 97, fig. 135; Caribbean: Littler & Littler, 2000: 344; Oman: Wynne, 2001: 365, fig. 33. (Fig. 41) Basionym: Ulva plumosa Hudson, 1762: 571 (type locality: Exmouth, Devonshire, England). Material examined: Taharaa, Tahiti, 14 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 16; Matira, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2558; Rarapai Islet, Rapa, 12 Nov. 2002, leg. J. L. Menou, UPF 2158. Thallus to 40 mm high, iridescent translucent green, in lax clumps composed of feather-like, oppositely branched axes. Main axes straight, to 500 µm in diameter, naked below and furnished above with two opposite rows of secondary branchlets to 100 µm in diameter and 2-3 mm long, progressively smaller towards the apex so as to give a plumose outline to the branches. Remarks: Non-staggered, opposite ramuli in straight rows differentiate this species from Bryopsis indica (Gepp & Gepp, 1908: 169, pl. 22 figs 10, 11). However, B. indica from Lord Howe Island (Kraft, 2000: 629) has been reported to vary greatly in respect to the straightness of ramuli rows. The French Polynesian plants (cf. Payri et al., 2000, fig. on p. 83, lower) show axes with straight, opposite ramuli that would conform with B. plumosa as understood here. Family Derbesiaceae Hauck Genus Derbesia Solier Derbesia marina (Lyngbye) Solier, 1846: 453; French Polynesia: Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 84; Marshall Islands: Dawson, 1956: 35, fig. 15a-b; South Australia: Womersley, 1984: 288, figs 98C-G, 99A; Netherlands Antilles: Stegenga & Vroman, 1988: 300, figs 1-4; Philippines: West & Calumpong, 1990: 186; Indian Ocean: Silva et al., 1996: 810; Belize: Littler & Littler, 1997: 99, fig. 137; Lord Howe Island: Kraft, 2000: 621, figs 39A-E; American Samoa: Skelton & South, 2004: 303, fig. 9. Basionym: Vaucheria marina Lyngbye, 1819: 79, pl. 22A (type locality: Kvivig, Strømø, Faeroes). Material examined: Entre deux Baies, Moorea, 23 Nov. 1995, leg. A. D. R. N’Yeurt, S109 UPF 495. Thallus to 30 mm high, dark yellowish green, forming filamentous tufts composed of sparsely branched axes to 20-44 µm in diameter. Chloroplasts circular, 4-6 µm long, pyrenoids absent. Ovoid to pear-shaped sporangia to 130 µm long and 100 µm in diameter occur singly or in pairs on the sides of branches, borne on a small stalk. Double walls sometimes occur at the base of branches or sporangia. Remarks: Epiphtyic on larger algae, to depths of 30 meters. The French Polynesian material has plastids lacking pyrenoids and double septa at the base of sporangia, thus conforming to D. marina and not to the similar species D. tenuissima (Morris et de Notaris) P. Crouan et H. Crouan. Moroever, D. tenuissima has larger axis diameters (up to 80 µm) than D. marina (Womersley, 1984). However, as discussed by Kraft (2000: 621) these characters are sometimes difficult to ascertain, with considerable overlap between the species reported in the literature. The Polynesian plants are most similar to Marshall Islands material described in Dawson (1956: 35), particularly as regards the shape of the sporangia (as illustrated in Payri et al., 2000). French Polynesian green algae 33 Figs 39-46. 39. Valonia fastigiata (UPF 2491). Habit of freshly-collected, split-out thallus, showing elongate vesicles. Scale = 5 mm. 40. Ventricaria ventricosa (UPF 2494). Habit of young thallus. Scale = 5 mm. 41. Bryopsis plumosa (UPF 2158). Habit. Scale = 1 mm. 42. Codium arabicum (UPF 73). Habit of flattened, convoluted thallus. Scale = 5 mm. 43. Codium geppiorum (UPF 2402). Habit of branched, terete thallus. Scale = 3 mm. 44. Codium geppiorum (UPF 2402). Detail of utricle, showing medullary filament plug (arrowhead). Scale = 200 µm. 45. Codium mamillosum (UPF 2111). Habit of large, spherical thallus attached to rhodolith. Note large utricles visible to the naked eye. Scale = 5 mm. 46. Codium mamillosum (UPF 2111). Longitudinal section of thallus, showing solid inner region and dense, rhizoidal basal holdfast (arrowhead). Scale = 5 mm. 34 A. D. R. N’Yeurt & C. E. Payri Family Codiaceae (Trevisan) Zanardini Genus Codium Stackhouse Key to the French Polynesian species of Codium 1a. Thallus applanate, adherent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arabicum 1b. Thallus spherical or terete, erect. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Thallus consisting of spherical balls, utricles large and apparent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mamillosum 2b. Thallus terete and dichotomously branched, utricles smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. geppiorum Codium arabicum Kützing, 1856: 35, pl. 100 fig. 2 (type locality: Tor, Sinai Peninsula, Gulf of Suez, Egypt). French Polynesia: Payri & N’Yeurt, 1997: 883, Payri et al., 2000: 102; Hawaiian Islands: Silva, 1952: 382, pl. 34b figs 11-13, Abbott & Huisman, 2004: 102, fig. 35A-B; Fiji: N’Yeurt, 2001: 718, Littler & Littler, 2003: 210; Marshall Islands: Dawson, 1956: 38, fig. 24; Lord Howe Island: Jones & Kraft, 1984: 255, figs 1-2; Indian Ocean: Silva et al., 1996: 850; Rotuma: N’Yeurt, 1996: 383, figs 41, 43, 59; Kenya: Van den heede & Coppejans, 1996: 391, figs 1, 5, 7. (Fig. 42) Heterotypic synonyms (given by Silva et al., 1996): Codium coronatum Setchell, 1926: 82, pl. 10 figs 2-5, pl. 11 figs 2, 3, pl. 12 figs 1, 5 (type locality: Taharaa, Tahiti). Sri Lanka: Børgesen, 1936: 67. — Codium coronatum var. insculptum Setchell. French Polynesia: Setchell, 1926: 83 (type locality: Arue, Tahiti). Material examined: Taharaa, Tahiti, 9 Jul. 1922, leg. W. A. Setchell & H. E. Parks, !UC 261420; Arue Reef, Tahiti, 30 May 1922, leg. W. A. Setchell & H. E. Parks, !UC 261421; Punaauia, Tahiti, leg. A. D. R. N’Yeurt, 10 May 1997, UPF 73, 6 Jul. 1997, UPF 72; Motu Totegegie, Gambier, 22 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 781; Tarakoi Pt., Rapa, Australs, 6 Nov. 2002, leg. C. E. Payri, UPF 2023; Takaraotaranga Pt., Rapa, 9 Nov. 2002, leg. C. E. Payri, UPF 2087; Mei Pt., Rapa, 18 Nov. 2002, leg. C. E. Payri, UPF 2221; Tauna islet, Rapa, 27 Nov. 2002, leg. C. E. Payri, UPF 2346; Hotel Pearl Beach, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2574; Tiahura, Moorea, 29 Sep. 2004, leg. A. D. R. N’Yeurt, In Herb. UPF. Thallus dark green, aplanate and dorsiventral, up to 15 cm broad and 1 cm thick; adhering strongly to the substratum, assuming a convoluted habit. Medullary filaments 17-23 µm in diameter; peripheral utricles clavate to pyriform, 58-88 µm broad and 380-500 µm long, with rounded apices. Remarks: Growing in shallow water on coralline substratum, on coral heads in the lagoon near the reef front. Setchell (1926: 82, and cited in a letter relating to Sri Lanka material in Børgesen, 1936: 68) separated the Tahitian species C. coronatum from C. arabicum by the thinner thallus, and shape and size of vesicles with crownlike circles of hair. However, Silva et al. (1996) considered both species conspecific, and our own examination of Setchell’s material concurs with this opinion, as the characters used to distinguish the species in question can be highly variable depending on environmental conditions. Throughout the Indo-Pacific, C. arabicum shows great anatomical variability (P. C. Silva, pers. com.). Codium geppiorum O.C. Schmidt, 1923: 50, fig. 33 (“geppii”). French Polynesia: Payri et al., 2000: 102; Sri Lanka: Børgesen, 1936: 68, fig. 3; Viêt Nam: Dawson, 1954: 395, fig. 13k (“geppii”); Marshall Islands: Dawson, 1956: 39, fig. 26 (“geppii”); New Caledonia: Valet, 1968: 41, pl. 10 fig. 5; Thailand: Egerod, 1975: 59, figs 25-26; Indian Ocean: Silva et al., 1996: 856; Kenya: Van den heede & Coppejans, 1996: 400, figs 11, 16; Fiji: N’Yeurt, 2001: 719, fig. 68. (Figs 43-44) French Polynesian green algae 35 Basionym: Codium divaricatum A. Gepp et E.S. Gepp, 1911: 136, 145 plate XXII Figs 195-199, nom. illeg. (syntype localities: Kai Islands and Celebes, Indonesia). Heterotypic synonym (given by Silva et al., 1996): Codium bulbopilum Setchell, 1924: 173, fig. 38 (type locality: Tutuila Island, American Samoa). French Polynesia: Setchell, 1926: 84, pl. 11 fig. 1, pl. 12 fig. 2; Lord Howe Island: Jones & Kraft, 1984: 26, figs 4, 5B-F, Kraft, 2000: 591, fig. 31F; Rotuma: N’Yeurt, 1996: 384, figs 42, 45; Samoa: Skelton & South, 2002a: 165, fig. 25F; Fiji: Littler & Littler, 2003: 212. Material examined: Hikueru, 17 Nov. 1996, leg. J. Orempuller, UPF 75; Afaahiti, Tahiti, 7 May 1997, leg. A. D. R. N’Yeurt, UPF 74; Vairao, Tahiti, 20 Mar. 1998, leg. J. Orempuller, UPF 593; Puka Puka, Aug. 1999, leg. Anonymous, UPF 767, 822; Motu Tarauraroa, Gambier, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 854; Ha’ruei, Rapa, 29 Nov. 2002, leg. J. L. Menou, UPF 2402; Tiahura, Moorea, leg. A. D. R. N’Yeurt, 29 Sep. 2004, In Herb. UPF; 30 Sep. 2004, UPF 2940; 1 Oct. 2004, UPF 2941; 3 Oct. 2004, UPF 2942. Thallus dark-green branched, terete and imbricating axes arching downwards; attached at various points to the substratum. Branching irregularly dichotomous, axes 2-3 mm in diameter and up to 8 cm long. Up to two plugged medullary filaments 35-41 µm in diameter, arising per utricle. Peripheral utricles obovoid, cylindrical to subspherical 140-235 µm broad and 382-500 µm long with rounded apices and occasional hairs up to 29 µm in diameter arising from the apical zone. Remarks: Growing attached to coral, on the fringing and barrier reefs. This is an edible species, being a favourite dish eaten raw, or cooked with fish in coconut milk, by many Pacific islanders (South, 1993: 339). The Codium geppiorum complex of species, which includes several poorly demarcated, dichotomously branched, terete members of the genus, is currently under molecular review with examination of specimens from around the world (P. C. Silva, pers. com.). The relationship of Pacific records of C. geppiorum to C. bulbopilum remain unclear pending molecular studies; the latter species is currently distinguished from C. geppiorum by its imbricating, hummock-like habit (Jones & Kraft, 1984: 26) but intermediate forms do exist. *Codium mamillosum Harvey, 1855: 565; 1858b: plate XLI, figs 1-4 (lectotype locality: Swan River, Western Australia). Japan: Okamura, 1915: 151, plate CXXXV, figs 10-16; Hawaiian Islands: Silva 1952: 389, fig. 16 pl. 35a, Abbott, 1986: 162, fig. 1, Abbott & Huisman, 2004: 109, fig. 39A-B; South Australia: Silva & Womersley, 1956: 269, fig. 6, Womersley, 1984: 230, figs 77A, 78A; Indian Ocean: Silva et al., 1996: 859; Fiji: N’Yeurt, 2001: 720, figs 48, 50, 69, Littler & Littler, 2003: 212; Wallis: N’Yeurt & Payri, 2004: 373; American Samoa: Skelton & South, 2004: 303. (Figs 45-46) Basionym: Lamarckia mamillosa (Harvey) Kuntze, 1891: 900. Material examined: Marotiri, Australs, 6 Nov. 2002, leg. I. R. D., UPF 2111, 2112, 2113, 2114. Plants spherical, solid, firm, 9-30 mm in diameter. Utricles large and visible to the naked eye, 300-620 µm in diameter and up to 3-4 mm long. Thallus with a single basal point of attachment to substratum. Plants lustrous when dry, non-adhering to the paper. Remarks: A common species at depths ranging from 15 to well below 60 m on the outer vertical reef slope in other parts of the Pacific (Fiji, Wallis, possibly Samoa and New Caledonia), in French Polynesia it is only reported dredged from great depths (80 to 120 m) in the southern Australs (Marotiri) where it it forms large dominant covers littering the seafloor. Whereas in shallower waters this species 36 A. D. R. N’Yeurt & C. E. Payri assumes a smaller, gregarious habit (9-12 mm in diameter; N’Yeurt, 2001), the French Polynesian deepwater plants are mostly unattached or epiphytic on freerolling rhodoliths, large (up to 30 mm in diameter) with big utricles visible to the naked eye. Specimens of this unusual growth form were confirmed as belonging to C. mamillosum by Professor P. C. Silva, and it represents one of the deepestinhabiting green algae in the ocean. Family Caulerpaceae Greville ex Kützing Genus Caulerpa J.V. Lamouroux A genus of some 70 tropical to temperate species (Womersley, 1984), many of which are dominant parts of the reef cover in certain areas. Some species (eg. C. racemosa, C. bikinensis, C. cupressoides) are edible and consumed regularly in several tropical localities such as the Philippines (Meñez & Calumpong, 1982), Fiji and Rotuma (South, 1993; N’Yeurt, 1996; South & Skelton, 2003a) and French Polynesia (Conte & Payri, 2002, 2006; this study). Fifteen species are listed for French Polynesia by Payri & N’Yeurt (1997), with one endemic species (C. seuratii). Meinesz et al. (1981), in a monographic study, listed some 11 species of Caulerpa from Moorea and Takapoto. Key to the French Polynesian species of Caulerpa 1a. Branchlets usually swollen, spherical or peltate . . . . . . . . . . . . . . . . . . . . . . 10 1b. Branchlets not swollen, spherical or peltate . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Branchlets strap-like, sub-dichotomous, dentate, or spirally twisted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. serrulata 2b. Branchlets opposite, radial or in multiple rows . . . . . . . . . . . . . . . . . . . . 3 3a. Branchlets spiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Branchlets filamentous, plumose or furry . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Branchlets straight, not basally compressed, spiniform, generally arranged on 3 to 5 opposite rows of densely ranked, sometimes inflated ramuli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cupressoides 4b. Branchlets angular, slightly compressed at the bases, radially arranged in 3 rows with thorn-like, never inflated ramuli . . . . . . . . . . . . C. urvilliana 5a. Branchlets in two or three opposite rows, stolons naked . . . . . . . . . . . . . . . 6 5b. Branchlets radially or verticillately arranged . . . . . . . . . . . . . . . . . . . . . . . . . 7 6a. Branchlets cylindrical and of the same diameter throughout. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sertularioides 6b. Branchlets compressed, basally constricted, with tapered tips . . C. taxifolia 7a. Branchlets slender, in verticillate whorls, stolons naked. . . . . . C. verticillata 7b. Branchlets radial, not in whorls, stolons covered with branchlets. . . . . . . . 8 8a. Branchlets in 3 rows only, thallus spiky. . . . . . . . . . . . . . . . . . . C. seuratii 8b. Branchlets in more than 3 rows, thallus furry. . . . . . . . . . . . . . . . . . . . . . 9 9a. Branchlets forming a continuous, uniform covering on fronds and stolons; thallus to 60 mm high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pickeringii 9b. Branchlets forming a non-continuous covering, stolons partly naked; thallus less than 12 mm high. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. webbiana 10a. Ends of the branchlets terminating abruptly in a single, thin peltate disk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. peltata 10b. Ends of branchlets cylindrical to clavate, if peltate thick and not occuring singly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 French Polynesian green algae 37 11a. Ends of branchlets inflated, subspherical to subpeltate or a mixture of both, laxly or radially beset about foliar axis . . . . . . . . . . . . . . . . . . . . . C. racemosa 11b. Ends of the branchlets elongate, club-shaped, flattened or slightly convex, arranged in an alternate or distichous manner. . . . . . . . . . . . . . C. bikinensis Caulerpa bikinensis W.R. Taylor, 1950: 66, pl. 33 (type locality: Bikini Atoll, Marshall Islands). French Polynesia: Meinesz et al., 1981: 431, figs 5-6; Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 90. (Fig. 47) Material examined: Bikini Atoll, 11 Jul. 1946, leg. W.R. Taylor, BM 701601; Rongelap Atoll, 21 Jun. 1946, leg. W.R. Taylor, BM 701600; Taiaro, 24 Sep. 1992, leg. R. Galzin, UPF 17, SUVA-A 1301; Nihiru, 30 Sep. 1995, leg. J. Orempuller, SUVA-A 546; Ilot aux Oiseaux, Tikehau, 4 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 20; Takapoto, 5 Nov. 1995, leg. L. Addessi, UPF 19, SUVA-A 1202, 1203; 8 Nov. 1995, leg. P. Loret, UPF 21; Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 615; Marokau, 22 Sep. 1996, leg. J. Orempuller, UPF 18, SUVA-A 1201; Haraiki, 15 Nov. 1996, leg. J. Orempuller, UPF 22; Vahitahi, Oct. 2000, leg. Anon., UPF 772, 773; Vairaatea, Oct. 2000, leg. Anon., UPF 768; Fangatau, 12 May 2003, leg. C. E. Payri, UPF 2707, 2708, 2709; Tatakoto, n.d., leg. Anon., UPF 763. Thallus to 15 cm high, greenish-yellow in colour, composed of a creeping stolon to 1 metre long and 10 mm wide, with erect axes 2-3 mm in diameter bearing elongate, club-shaped, flattened or slightly convex branchlets to 10 mm long and 4 mm in diameter, arranged in an alternate or distichous manner. Remarks: Found growing in the lagoon and outer slope of atolls, to depths of 70 m. This species is edible, being a favorite dish as a salad with fish in the Austral Islands. Caulerpa bikinensis appears characteristic of atolls of the Tuamotu archipelago, having been collected in the northern section (Tikehau), eastern part (Fangatau, Takatoto, Takapoto) and as far down as Nengo Nengo and Vairaatea. In Takapoto (north-east), it forms an almost continuous belt around the atoll in the outer slope from a depth of 35 m to over 70 m, with covers exceeding 75 % in parts (Meinesz et al., 1981: 433). However, it has not been reported to date from the more southern Gambier islands. It is reported to be present in the Australs Group (Raivavae Island), where the local population traditionally consumes it, along with C. racemosa (Payri et al., 2000: 90; Conte & Payri, 2002: 171, 2006). Unfortunately, no herbarium specimens of C. bikinensis from the Australs could be located in UPF. The French Polynesian material is in excellent agreement with authentic material from Bikini Atoll examined in BM. Some plants from Bikini Atoll and other Marshall Islands (BM 701599, 701602-701607) originally identified as varieties of Caulerpa racemosa by W.R. Taylor (1950), were subsequently determined to be forms of C. bikinensis by H. Ohba (31 Jul. 1998, annotation on herbarium sheets), and we concur with that opinion after examination of the same. French Polynesian plants also show a variety of ecomorphs. Caulerpa cupressoides (Vahl) C. Agardh, 1823: 441. French Polynesia: Setchell, 1926: 85, Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 90; Indian Ocean: Silva et al., 1996: 815; Lord Howe Island: Kraft, 2000: 597, fig. 32 C-E. Basionym: Fucus cupressoides Vahl, 1802: 38 (type locality: St Croix, Virgin Islands). Remarks: Caulerpa cupressoides being a species with continuously variable phenotypes (Coppejans & Prud’homme van Reine, 1992; Kraft, 2000: 597), it may be futile to assign clearly defined infra-specific combinations; however the following two varieties and forms represent consistently encountered patterns in French Polynesian populations. 38 A. D. R. N’Yeurt & C. E. Payri Caulerpa cupressoides (Vahl) C. Agardh var. lycopodium Weber-van Bosse, 1898: 335, pl. XXVII figs 5-13; pl. XXVIII figs 10-12, 14. French Polynesia: Payri et al., 2000: 90; Fiji: South & N’Yeurt, 1993: 114, fig. 8; Indian Ocean: Silva et al., 1996: 817; Caribbean: Littler & Littler, 2000: 360, fig. on p. 361. (Figs 48-49) Basionym: Caulerpa lycopodium J. Agardh, 1847: 6, nom. illeg. (syntype localities: Brazil & West Indian Ocean). Material examined: Faarapa, Tahiti, 19 Jun. 1922, leg. W. A. Setchell & H. E. Parks, UPF 24, 25; Tiahura, Moorea, 13 Jun. 1984, leg. C. E. Payri, UPF 2796, 2797, 2878; Afaahiti, Tahiti, 7 May 1997, leg. A. D. R. N’Yeurt, UPF 23; Taravai, Gambier, 20 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 874, 880; Motu Totegegie, fringing reef in front of airport, Gambier, 22 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 802; Hotel Intercontinental, Bora Bora, on concrete stilts of bungalows, 19 Aug. 2002, leg A. D. R. N’Yeurt, UPF 2584; Agairoa Bay, Rapa, 2 Nov. 2002, leg. C. E. Payri, UPF 1907, 1908; Tekogoteemu Pt., fringing reef, 5 Nov. 2002, leg. C. E. Payri, UPF 2000; Aruroa Pt., Rapa, 22 Nov. 2002, leg. A. D. R. N’Yeurt, UPF 2246; Ha’urei, Rapa, on first buoy at exit of bay, 29 Nov. 2002, leg. J. L. Menou, UPF 2419, 2420. Thallus forming dense aggregations, with a smooth spreading stolon up to 30 cm long and 3 mm in diameter, anchored by numerous rhizoid-bearing branches spaced at close (0. 5-1 cm) intervals. Erect axes up to 4 cm tall and 2 mm wide, oppositely pinnate and compressed, with upward curving tendency, tapering to a sharp point at the tip, and generally twice as long as the diameter of the supporting axis, spaced at relatively wide (2-3 cm) intervals along spreading stolon. The ramuli usually in ranks of threes, sometimes twos, oppositely pinnate with mucronate, upward-curving branchlets up to 1 mm long. Remarks: This variety of C. cupressoides is edible, being consumed raw as a salad in French Polynesia on the island of Rapa (local name: mama’ga, Fig. 49). It could be confused with Caulerpa lesonii Bory de Saint-Vincent (1828: 193) but the latter species has a pronouncedly wide and flattened rachis, with short, basally constricted pinnules whose length is less than the diameter of the supporting rachis. However, intermediate forms exist between these species, and C. lessonii itself is intermediate between C. cupressoides and C. serrulata (Coppejans & Prud’homme van Reine, 1992: 692). Caulerpa cupressoides (Vahl) C. Agardh var. mamillosa (Montagne) Weber-van Bosse, 1898: 332 pl. 28 fig. 6. Somalia: Sartoni, 1978: 402, fig. 2b; Indonesia: Coppejans & Prud’homme van Reine, 1992: 679, fig. 3A, 8B (as ecad mamillosa); Indian Ocean: Silva et al., 1996: 818; Rotuma: N’Yeurt, 1996: 377, figs 25, 35; Lord Howe Island: Kraft, 2000: 597, fig. 32C-D; Wallis: Payri et al., 2002: 44, pl. 1 fig. 6; Fiji: Littler & Littler, 2003: 216. (Figs 50-51) Basionym: Caulerpa mamillosa Montagne, 1842: pl. 6, fig. 3 (syntype localities: Agalega Islands, Indonesia; Mangareva Islands, Gambier, French Polynesia). Material examined: Agalega Island, Indonesia, n.d., leg. Leduc, BM 515883, PC 10326 (syntype); Mangareva, Gambier, n.d., leg. M. Hombron, PC 10329 (syntype); Haraiki, 14 Nov. 1996, leg. J. Orempuller, UPF 709; Hikueru, 16 Nov. 1996, leg. J. Orempuller, UPF 708; Atituiti, Mangareva, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 776, 19 Nov. 2000, UPF 918; Kirimiro, Mangareva, 21 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 919. Thallus stout and bushy, with the erect foliar axes closely spaced at 0. 5-1 cm intervals along a relatively thick, spreading stolon up to 2.5 mm in diameter. The foliar axes are several times forked very early from the base, bearing mucronate, obvoid to subnavicular, distinctly inflated ramuli in several ranks. Remarks: This species appears in French Polynesia to be confined to atolls of the Tuamotu and the Gambier. It could be confused with C. urvilliana, but our own examination of the relevant type specimens in PC confirmed that C. urvilliana is French Polynesian green algae 39 characterised by angular erect branches with dentate, thorn-like pinnules, contrasting with the straight branches with densely ranked, inflated ramuli of C. cupressoides var. mamillosa (Fig. 51). Coppejans & Prud’homme van Reine (1992: 686) suggested that C. urvilliana could represent an extreme growth form of the continuously variable taxa C. cupressoides; molecular studies alone could provide a definitive answer to these questions. *Caulerpa nummularia Harvey ex J. Agardh, 1873: 38 (syntype localities: Tonga and Nuku-Hiva, Marquesas). Indian Ocean: Silva et al., 1996: 827; Belize: Littler & Littler, 1997: 105, fig. 150; Fiji: Littler & Littler, 2003: 224, South & Skelton, 2003: 542, fig. 3; Hawaiian Islands: Abbott & Huisman, 2004: 121, fig. 44A, Hodgson et al., 2004: 28; South Africa: De Clerck et al., 2005: 66, fig. 35. (Fig. 52) Misapplied names: Caulerpa peltata J.V. Lamouroux f. imbricata (G. Murray) Weber-van Bosse, 1898: 375. French Polynesia: Setchell, 1926: 86. — Caulerpa peltata J.V. Lamouroux. French Polynesia: Payri et al., 2000: 93. Material examined: Tahara, Tahiti, 17 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 841598, 841599, 841607 (f. imbricata); Rurutu, 18 Aug. 2000, leg. C. E. Payri, UPF 755; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 855. Plants small and delicate, occurring as single stolons to 80 mm long and 0.3-0.7 mm in diameter, sparingly provided with short rhizoids. Cylindrical erect branchlets 1-5 mm long occur at 2-3 mm intervals, with consistently thin, terminal irregularly peltate discs 1-3 mm in diameter with crenulate margins; occurring sometimes singly but usually with additional blades proliferating from marginal or central areas of parent blades. Remarks: Caulerpa peltata var. exigua (Weber-van Bosse, 1898: 377; Eubank, 1946; Kraft, 2000: 601, fig. 33F) is likely synonymous with C. nummularia (Abbott & Huisman, 2004: 121). Other workers consider them a form of C. racemosa var. peltata (Coppejans & Prud’homme van Reine, 1992: 696), but the latter species has large, non-crenulated, singly-placed, inconsistently peltate assimilators borne on a stolon thicker than 1 mm in diameter. The status of C. peltata J.V. Lamouroux (1809c: 332) is unclear, pending molecular studies in the C. racemosa – C. peltata complex. Some workers (e.g. Kraft, 2000: 601, fig. 33E) recognize C. peltata for plants with single, peltate assimilators on stolons less than 1 mm in diameter. Payri et al., (2000: 93, upper photo) show a plant from Moorea under C. peltata in fact represents C . nummularia (with slightly crenulated, new disks emerging from the borders of subtending ones). Tahitian “C. peltata f. imbricata” from Setchell’s collection (BM 841598) has thin assimilators which would place it well within C. nummularia as understood here, but differs from the small, dense clavate ramuli of the Sri Lanka type material of C. imbricata G. Murray (Eubank, 1946: 423). A further related species from Tonga, C. stellata Harvey, has distinctly stellate, centrally tiered assimilators and is currently placed under C. peltata var. stellata (Harvey ex J. Agardh) Weber-van Bosse (1898: 377) but its specietal status might need to be resurrected. Caulerpa pickeringii Harvey et Bailey, 1851: 373 (type locality: “Wilson Island” (Manihi), Tuamotu). French Polynesia: Taylor, 1973: 38 (Moorea); Meinesz et al., 1981: 433, figs 11-14, Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 92; Indonesia: Weber-van Bosse, 1898: 272, pl. 21 figs 7, 8; Caroline Islands: Trono, 1968: 167, pl. 15 fig. 5. (Figs 53-57) Material examined: Nihiru, 11 Sep. 1995, leg. J. Orempuller, UPF 27; Kauehi, 25 Sep. 1995, leg. J. Orempuller, SUVA-A 945; Reka Reka, 9 Nov. 1996, leg. J. Orempuller, UPF 26, 28, SUVA-A 1348; Oeno atoll, Pitcairn, 2 Aug. 1997, leg. J. Starmer, UPF 665; Henderson, Pitcairn, 17 Aug. 1997, leg. J. Starmer, UPF 666, 667, 668; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2461, 2462, 2463. 40 A. D. R. N’Yeurt & C. E. Payri Figs 47-53. 47. Caulerpa bikinensis (UPF 2703). Habit. Scale = 7 mm. 48. Caulerpa cupressoides var. lycopodium (UPF 2000). Habit of dried specimen. Scale = 10 mm. 49. Caulerpa cupressoides var. lycopodium (UPF 2419). Habit of freshly-collected thallus. Scale = 2 mm. 50. Caulerpa cupressoides var. mamillosa (MA 10329). Partial habit of Syntype collection from Mangareva, Gambier. Scale = 7 mm. Photographed with permission from the Museum National d’Histoire Naturelle, Paris. 51. Caulerpa cupressoides var. mamillosa (MA 10329). Detail of Syntype showing distinctly inflated, mucronate ramuli (arrowheads). Scale = 2 mm. 52. Caulerpa nummularia (UPF 855). Habit showing thin, disk-like assimilators with crenate margins. Scale = 3 mm. 53. Caulerpa pickeringii (UPF 26). Habit of densely-invested plant from Reka Reka. Scale = 10 mm. French Polynesian green algae 41 Plants dark green in colour, spongy, to 6 cm high, composed of a creeping stolon to 3 mm wide and 15 cm long, bearing profuse, sparsely and irregularly branched erect axes to 3 cm long, with blunt rounded tips. Secondary branchlets are usually radially arranged about both the erect axes and stolon, forming a uniform, unbroken fur-like covering (in some cases, parts of the plant are less densely covered than other portions). The secondary branchlets are up to 700 µm long and 130 µm wide, dichotomously branched 3 or 4 times in a single plane, with rounded, mucronate tips. Remarks: Common on atolls (Tuamotu) on the algal ridge or in the lagoon at depths of 3-5 meters, and less frequently on the outer reef slope at 7-20 metres depth. Less frequent on the barrier reef of high volcanic island. Originally described from Manihi atoll in the Tuamotu archipelago (Harvey & Bailey, 1851), C. pickeringii was long considered endemic to French Polynesia, with records of the species elsewhere (eg. Tanzania, Jaasund, 1976: 19, fig. 40) considered to be misapplied names for C. elongata Weber-van Bosse (1898: 271, pl. xxi, figs 5, 6; syntype localities Indonesia, Tonga) or C. webbiana var. pickeringii (Harvey et Bailey) Eubank (1946: 416; type locality Tuamotu) (Meinesz et al., 1981: 433). As remarked by the latter authors, the original material from French Polynesia has both erect branches and stolons densely covered by ramuli, with the stolons totally hidden and having the superficial appearance of terete species of Codium. This situation is evident in the herbarium material examined in UPF, and differs from the descriptions of “C. pickeringii” in Jaasund (1976), and Littler & Littler, (2003: 226, photo from Fiji) where the stolons are sparsely covered with ramuli, or naked in parts. On the other hand, plants with sparsely covered stolons superficially looking like C. elongata or C. webbiana var. pickeringii have been found in Tatakoto, French Polynesia (UPF 761) and also from Henderson, Pitcairn Island (UPF 666, 668). The latter material from Pitcairn is interesting, because parts of the thallus look like C. pickeringii with the stolon densely covered, and others have a more lax appearance (Fig. 55). All the specimens examined from Polynesia however have ramuli 3-4 times dichotomously branched in a single plane, unlike what is reported for C. elongata, which has dichotomies at right angles to each other (Littler & Littler, 2003: 216). Caulerpa webbianna var. pickeringii can have densely clothed assimilators and a Codium-like habit (Kraft, 2000: 608, fig. 35 E) but these are not as dense as in C. pickeringii, and they are much shorter and also branch at right-angle to each other. C. seuratii, as understood here, has mostly tristichous ramuli with distichous or tristichous, mucronate extremities and a plumose appearance when dry, which differs from C. pickeringii. “Typical” C. pickeringii with fully, densely covered stolons also occurs in Oeno Atoll and Pitcairn Island (UPF 665, 667; Fig. 54). The Oeno Island specimen has very dense arrangement of ramuli, which are the most elongate (1500-2000 µm) and have the smallest diameter (44-90 µm) of all the material examined. Material from Reka Reka (UPF 26, Fig. 53) is intermediate, with a length of 1300-1500 µm and a diameter of 89-178 µm, and a lax plant from Tatakoto (UPF 761) had ramuli lengths of 1300-1780 µm and diameters of 133-244 µm. On the other hand, a specimen from Henderson, Pitcairn Island (UPF 666) had the shortest (1220-1300 µm) and widest (200-355 µm) ramuli. It could be that these variations are related to the degree of exposure, depth or luminosity in the different habitats. These observations raise the question as to whether C. elongata, C. webbiana var. pickeringii and C. pickeringii would represent various ecomorphs of the same continuously variable entity (C. webbiana), as suggested by Eubank (1946: 416); the definitive answer to this would lie in a molecular study of these taxa. 42 A. D. R. N’Yeurt & C. E. Payri Figs 54-59. 54. Caulerpa pickeringii (UPF 665). Habit of large, very densely-invested plant from Oeno Atoll. Scale = 10 mm. 55. Caulerpa pickeringii (UPF 668). Habit of “hybrid” plant from Henderson Island, showing both typically dense, C. pickeringii-like (arrowhead) and lax, C. seuratii-like branches (arrow). Scale = 10 mm. 56. Caulerpa pickeringii (UPF 665). Transverse section of erect branch, showing ramuli 3-4 times dichotomously branched in a single plane. Scale = 100 µm. 57. Caulerpa pickeringii (UPF 761). Detail of apex, showing rounded, mucronate ends. Scale = 20 µm. 58. Caulerpa racemosa var. racemosa (UPF 669). Habit of pressed specimen. Scale = 5 mm. 59. Caulerpa racemosa var. typica (UPF 2592). Detail of freshly-collected plant from intertidal habitat in Bora Bora, showing thick rhizome with clavate to subspherical ramuli. Scale = 2 mm. Caulerpa racemosa (Forsskål) J. Agardh, 1873: 35-36 (var. racemosa). French Polynesia: Setchell, 1926: 85, Taylor, 1973: 38 (Bora Bora); Meinesz et al., 1981: 433, Payri & N’Yeurt, 1997: 882, Payri et al., 2000: 94, Conte & Payri, 2002: 167, fig. 3; Somalia: Sartoni, 1978: 406, fig. 4c; Papua New Guinea: Coppejans & Meinesz, 1988: 191, figs 22, 23, Coppejans, 1992: 397, fig. 4C, Coppejans et al., 1995: French Polynesian green algae 43 78, fig. 7, Littler & Littler, 2003: 226; Indonesia: Coppejans & Prud’homme van Reine, 1992: 698, fig. 18A-B; Indian Ocean: Silva et al., 1996: 832; Rotuma: N’Yeurt, 1996: 378, figs 28, 36; Lord Howe Island: Kraft, 2000: 602, fig. 34A-D; Fiji: N’Yeurt, 2001: 714, fig. 43; Wallis: Payri et al., 2002: 44, pl. 1 fig. 2; Hawaiian Islands: Abbott & Huisman, 2004: 122, fig. 44B-C. (Figs 58-59) Basionym: Fucus racemosus Forsskål, 1775: 191 (type locality: Suez, Egypt). Material examined: Arue, Tahiti, 27 Jul. 1922, leg. W. A. Setchell & H. E. Parks, UPF 33; Marokau, 22 Nov. 1996, leg J. Orempulller, UPF 30; Taharaa, Tahiti, 14 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 29; Afaahiti, Tahiti, 7 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 32; Punaauia, Tahiti, 6 Jul. 1997, leg. C. E. Payri, UPF 35; Tedside, Pitcairn, 28 Aug. 1997, leg. J. Starmer, UPF 669; Arue, Tahiti, 30 Mar. 1998, leg. J. Orempuller, UPF 684; Tubuai, 14 Nov. 2000, leg. C. Monier, UPF 808; Raivavae, Nov. 2000, leg. C. Monier, UPF 807; Rikitea, Mangareva, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 848; Motu Taraururoa, Gambier, 16 Nov. 2000, UPF 891; Atituiti, Mangareva, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 897; Hotel Le Meridien, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2592; Tauna Islet, Rapa, 27 Nov. 2002, leg. C. E. Payri, UPF 2340, 2341. Thallus dark green, up to 15 cm long, with a spreading stolon to 3 mm in diameter beset with ventral rhizoids to 15 mm long. Ascending foliar axes up to 3 cm long, bearing up to 15 radially disposed stipitate ramuli with subspherical inflated ends 2-4 mm in diameter. Remarks: Common on the reef flat, growing on coral heads, sponges and soft corals in shallow waters. The typical form of the taxon includes what was previously called C. racemosa var. clavifera (Turner) Weber-van Bosse, 1898: 361, pl. XXXIII figs 1-3 (see Silva et al., 1996: 832). Caulerpa racemosa is one of the most common and variable of Caulerpa species, with several recognisable ecomorphs or varieties, and a continuum of intergrades between these depending on environmental factors. It is also one of the most favoured edible algal species complex eaten raw as salad in the Asia-Pacific region (Abbott, 1991; South, 1993, 1998; Trono & Toma, 1993; Conte & Payri, 2002, 2006). Aside from the typical form, at least three distinct varieties of C. racemosa occur in French Polynesia. *Caulerpa racemosa (Forsskål) J. Agardh var. peltata (J.V. Lamouroux) Eubank in Stephenson, 1944: 349. Hawaiian Islands: Eubank, 1946: 421, fig. 2r-s; Philippines: Meñez & Calumpong, 1982: 8, pl. 2K; Kenya: Coppejans & Beeckman, 1989: 388, figs 27-29; Indonesia: Coppejans & Prud’homme van Reine, 1992: 696, fig. 17B (as ecad peltata); Rotuma: N’Yeurt, 1996: 378, figs 31, 37. (Figs 60-61) Basionym: Caulerpa peltata J.V. Lamouroux, 1809c: 332 (type locality: Antilles). Material examined: Punaauia, Tahiti, 10 May 1997, leg. A. D. R. N’Yeurt, UPF 31; Vairao, Tahiti, 20 Mar. 1998, leg. J. Orempuller, UPF 590, 591; Nuku Hiva, Marquesas, 1997, leg. J. Orempuller, UPF 576; Rikitea, Mangareva, Gambier, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 848; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 856, 893; Bora Bora, 16 Aug. 2002, leg A. D. R. N’Yeurt, UPF 2531, 2532, 2533, 2534; Ha’urei, Rapa, 11 Nov. 2002, leg. J. L. Menou, UPF 2125, 2126, 2127, 2128, 2135; 29 Nov. 2002, UPF 2425; 13 Nov. 2002, UPF 2184, 2185, 2186, 2187; Rapa Iti, Rapa, 28 Nov. 2002, leg. J. L. Menou, UPF 2403, 2404, 2405; Tiahura, Moorea, 30 Sep. 2004, leg. A. D. R. N’Yeurt, and A. Pham, UPF 2958. Thallus in densely intermingled clumps 5-10 cm across; stolons about 8 cm long and sparingly provided with short rhizoidal branches. Erect foliar axes 1-1.5 cm long at 2-3 mm intervals, with flattened clavate to subpeltate (sunhatshaped) discs 3-5 mm in diameter either singly at the end, or several discs axially arranged around the main foliar branches. Both subsperical and subpeltate ramuli can occur on the same erect branch. 44 A. D. R. N’Yeurt & C. E. Payri Remarks: This variety of C. racemosa with a predominance of large peltate assimilators is often confused in the literature with C. peltata, and vice-versa. We follow Eubank (1946) and Kraft (2000) in assigning the varietal name “peltata” to plants of C. racemosa with thick stolons (1-2 mm diam.) and some (or all) assimilators with peltate ramuli, but never the entire assimilators being small, thin peltate disks on slender stolons to 700 µm in diameter. On the other hand, South & Skelton (2003b) consider Caulerpa racemosa var. peltata a synonym of Caulerpa peltata. In French Polynesia, we have observed plants intermediate between C. racemosa vars peltata, occidentalis (here considered synonymous with var. racemosa) and turbinata, suggesting that all these ecomorphs represent the same continuously variable entity of C. racemosa. Caulerpa racemosa (Forsskål) J. Agardh var. turbinata (J. Agardh) Eubank, 1946: 420, figs 2o-q. French Polynesia: Conte & Payri, 2002: 167, fig. 3; Marshall Islands: Dawson, 1956: 35, fig. 16a; Kenya: Coppejans & Beeckman, 1989: 386, figs 24-26; Indonesia: Coppejans & Prud’homme van Reine, 1992: 698, fig. 19A-B (as ecad turbinata); Fiji: South & N’Yeurt, 1993: 129, fig. 24; Indian Ocean: Silva et al., 1996: 837; Rotuma: N’Yeurt, 1996: 379, fig. 29. (Figs 62-63) Basionym: Caulerpa clavifera (Turner) C. Agardh var. turbinata J. Agardh, 1837: 173 (type locality: near Tor, Sinai Peninsula, Egypt). Material examined: Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 34; Ua Huka, Marquesas, Aug. 1999, leg. E. Conte & C. E. Payri, UPF 800; Tarakoi, Rapa, 5 Nov. 2002, leg. C. E. Payri, UPF 2006, 2007, 2008; Moerai, Rurutu, 20 Jul. 2005, leg. J. Peterano and N. Gebel, UPF 3007, 3008. Thallus lacking a well-defined spreading stolon; erect branchlets to 25 mm long, radially and densely beset with trumpet-shaped, terminally flattened to concave assimilators up to 1.5 mm long and 1-3 mm in diameter. Some subpeltate or clavate ramuli can occur amidst the predominantly turbinate assimilators (Fig. 63). Remarks: This variety (intermediate between vars clavifera and peltata) of C. racemosa has a predominance of characteristic trumpet-shaped assimilators, densely beset about the foliar axis. Formerly known as C. racemosa var. chemnitzia (Esper) Weber-van Bosse (1898: 370), the nomenclature of this variety was reviewed by Eubank (1946: 421). It is one of the preferred varieties of C. racemosa for human consumption (South 1993; Conte & Payri, 2002, 2006). In French Polynesia, it is commonly consumed on the island of Rurutu, Australs (N. Gebel, pers. com.). *Caulerpa racemosa (Forsskål) J. Agardh var. uvifera (C. Agardh) J. Agardh, 1873: 35. Indonesia: Weber-van Bosse, 1898: 362, pl. 33, figs 6-7; fig. 23; 1913: 105; Bikini Atoll: Taylor, 1950: 63; 1960: 153, pl. 17, fig. 3; pl. 18, fig. 4; Marshall Islands: Dawson, 1957: 106; New Caledonia: Valet, 1968: 45, pl. 7, fig. 2; Solomon Islands: Womersley & Bailey, 1970: 276; Philippines: Meñez & Calumpong, 1982: 9, pl. 2D; Rotuma: N’Yeurt, 1996: 380, fig. 27a-b. Basionym: Caulerpa uvifera C. Agardh, 1817: XXIII (type locality: Red Sea). Material examined: Tahiti, 15 Aug. 1912, leg. J. E. Tilden, BM 841501; Arue Point, Tahiti, 27 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 841502, 841509; Paea, Tahiti, 28 Jul. 1928, leg. C. Crossland, BM 841503. Branchlets 10-20 mm high, corpulent; ends of branchlets club to trumpet-shaped, 1-2 mm in diameter, compact, very densely beset about foliar axis. French Polynesian green algae 45 Figs 60-65. 60. Caulerpa racemosa var. peltata (UPF 31). Habit of pressed specimen. Scale = 5 mm. 61. Caulerpa racemosa var. peltata (IFR). Detail of freshly-collected plant from intertidal habitat in Bora Bora, showing relatively thick rhizome with both clavate (arrowhead) and subpeltate (arrow) ramuli. Scale = 3 mm. 62. Caulerpa racemosa var. turbinata (UPF 2006). Habit of pressed plant. Scale = 10 mm. 63. Caulerpa racemosa var. turbinata (UPF 3008). Detail of freshly-collected plant of an edible variety from Rurutu Island, showing odd subpeltate assimilators (arrowhead) on same branch as predominantly turbinate ones. Scale = 5 mm. 64. Caulerpa serrulata (UPF 2557). Habit of untwisted ecomorph. Scale = 2 mm. 65. Caulerpa serrulata (UPF 2556). Habit of twisted ecomorph. Scale = 2 mm. Remarks: This succulent, least fibrous variety is one of the most favoured for human consumption (South, 1993; N’Yeurt, 1996). Caulerpa serrulata (Forsskål) J. Agardh, 1837: 174. French Polynesia: Payri & Meinesz, 1985a: 507; Payri et al., 2000: 94; Viêt Nam: Dawson, 1954: 393, fig. 10a; 1956: 38, fig. 23; Caroline Islands: Trono, 1968: 169, pl. 14 figs 1-2; pl. 16 figs 4, 8; 46 A. D. R. N’Yeurt & C. E. Payri pl. 17 fig. 9; New Caledonia: Valet, 1968: 43, pl. 9, fig. 1; Somalia: Sartoni, 1978: 408, fig. 5a; Philippines: Meñez & Calumpong, 1982: 9, pl. 2E, Trono, 1986: 218, fig. 14; Coppejans & Meinesz, 1988: 192, figs 27-28; Coppejans & Beeckman, 1989: 120, figs 24-25; Indonesia: Coppejans et al., 1992: 701, fig. 20B; Fiji: South & N’Yeurt, 1993: 117, fig. 12, N’Yeurt, 2001: 715, fig. 44, Littler & Littler, 2003: 230; Papua New Guinea: Coppejans, 1992: 403, fig. 7; Coppejans et al., 1995: 78, figs 9-10; Indian Ocean: Silva et al., 1996: 841; Rotuma: N’Yeurt, 1996: 382, figs 30, 40; South Africa: Leliaert et al., 2001: 452, fig. 1; Samoa: Skelton & South, 2002a: 164, figs 24B-D; Hawaiian Islands: Abbott & Huisman, 2004: 123, fig. 45A. (Figs 64-65) Basionym: Fucus serrulatus Forsskål, 1775: 189 (type locality: Mokha, Yemen). Material examined: Tahiti, 15 Aug. 1912, leg. J. E. Tilden, BM 841450; Tikehau, 5 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 36; Haraiki, 15 Nov. 1996, leg. J. Orempuller, UPF 37; Mokoto, Mangareva, 27 Sep. 1997, leg. J. Starmer, UPF 629; Vairaatea, Oct. 2000, leg. Anon, UPF 769; Atituiti, Mangareva, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 897; Taravai, Gambier, 20 Nov. 2000, UPF 872, 889, 871; Kirimiro, Mangareva, blue hole, 21 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 908; Matira, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2556, 2557, 2582; Hotel Intercontinental, Bora Bora, 18 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2546; Tiahura, Moorea, 28 Sep. 2004, leg. A. D. R. N’Yeurt, UPF 2938. Thallus light to dark green, with a spreading stolon up to 20 cm long and 2 mm wide, bearing flattened to compressed erect branches up to 7 cm tall at 1-4 cm intervals along the spreading stolon. The erect branches are several times dichotomously or irregularly branched, terete below up to point of dichotomy, the rest compressed (1-2 mm broad) with moderate to strong twisting and serrated margins; the serrations more pronounced on the outwardly facing edge of the twist. Remarks: One of the commonest Caulerpa species growing in shallow waters, on the fringing and barrier reef of high islands (Society, Gambier) and in the lagoon of the atolls (Tuamotu). Assimilators are characteristically serrated (sawtoothlike) and of variable ecomorphology, ranging from untwisted, single plane branching to highly twisted. Caulerpa sertularioides (S.G. Gmelin) Howe, 1905: 576. French Polynesia: Payri et al., 2000: 96; Hawaiian Islands: Eubank, 1946: 417, fig. 2c-d; Abbott & Huisman, 2004: 124, fig. 45B-C; Somalia: Sartoni, 1978: 410, fig. 5b; Papua New Guinea: Coppejans & Meinesz, 1988: 192, fig. 29, Coppejans et al., 1995: 80, fig. 8; Kenya: Coppejans & Beeckman, 1990: 120, figs 26-27; Indonesia: Coppejans & Prud’homme van Reine, 1992: 704, fig. 21A; Indian Ocean: Silva et al., 1996: 843; Belize: Littler & Littler, 1997: 107, fig. 154; Fiji: South & N’Yeurt, 1993: 121, fig. 16, N’Yeurt, 2001: 716, fig. 42; Littler & Littler, 2003: 232; Wallis: Payri et al., 2002: pl. 1 fig. 4. (Figs 66-67) Basionym: Fucus sertularioides S.G. Gmelin, 1768: 151, pl. XV fig. 4 (type locality: “in coralliis americanis”). Material examined: Tahiti, 15 Aug. 1912, leg. J. E. Tilden, BM 841479; Maraa Point, Tahiti, 24 Sep. 1928, leg. C. Crossland, BM 841482; Taaone, Tahiti, 7 May 1997, leg J. Orempuller, UPF 38, 39; Raivavae, Nov. 2000, leg. C. Monier, UPF 809; Hotel Le Meridien, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2591. Thallus to 6 cm tall, with terete stolons 0.25-1.0 mm in diameter, bearing sparse plumose, pinnate erect branches to 18 mm wide and 70 mm long, simple or occasionally (sub-) dichotomously divided. Ultimate branchlets cylindrical throughout, not constricted at the base, to 8 mm long and 200 µm in diameter, upcurved, with mucronate tips. French Polynesian green algae 47 Remarks: Growing in patches, creeeping in sandy substratum at the bottom of the lagoon or on fallen logs and other surfaces in estuaries. Caulerpa seuratii Weber-van Bosse, 1910: 2, pl. I figs 5-9, pl. II fig. 1 (type locality: Tokaai, Tuamotu, French Polynesia). French Polynesia: Meinesz et al., 1981: 436, figs 22-25; Payri & N’Yeurt, 1997: 883; Payri et al., 2000: 96. (Figs 68-70) Material examined: Raroia, 2 Aug. 1952, leg. M. S. Doty & J. Newhouse, SUVA-A 6742; Kauehi, Sep. 1995, leg. J. Orempuller, UPF 40, SUVA-A 1266; Nihiru, 1 Oct. 1995, leg. J. Orempuller, SUVA-A 947; Hikueru, 27 Sep. 1996, leg. J. Orempuller, UPF 41, 44; Hiti, 11 Nov. 1996, leg. J. Orempuller, UPF 46; Marokau, 22 Nov. 1996, leg. J. Orempuller, UPF 42, 43, 45, SUVA-A 1265; Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 619. Thallus to 8 cm high, with stolons to 40 cm long and 4 mm wide, bearing occasionally irregularly divided erect branches. Stolons spiky, being covered with short mucronate branchlets to 1.3 mm long and 300 µm in diameter. Erect branches bear terete pinnules to 5 mm long and 300 µm in diameter which are disposed radially in 3 or 4 parallel lines around the branch axis, with di- or trichotomously divided, mucronate tips. Remarks: This elegant species has long been considered endemic to the Tuamotu archipelago, French Polynesia, and indeed had not been republished since the original description by Weber-van Bosse (1910) until over 70 years later (Meinesz et al., 1981), although it had been collected in 1952 from Raroia atoll by Maxwell Doty & Jan Newhouse (SUVA-A 6742). It has also been reported from Fiji (Littler & Littler, 2003: 232), but the inconspicuous plants from the latter locality appear predominantly bipinnate with a lax habit and non-pubescent runners, contrasting with the always tristichous, larger French Polynesian plants whose runners are fully invested with short ramuli. Furthermore, the end of the pinnules in French Polynesian plants are always distichously to tristichously divided (Fig. 69), while those reported for the Fiji plants are divided up to five times, and could rather represent a form of C. webbiana. Caulerpa taxifolia (Vahl) C. Agardh, 1817: XXII. French Polynesia: Payri et al., 2000: 98; Somalia: Sartoni, 1978: 410, fig. 5c; Indonesia: Coppejans & Prud’homme van Reine, 1992: 706, figs 6B, 22B; Fiji: South & N’Yeurt, 1993: 122, fig. 17; Littler & Littler, 2003: 234; Papua New Guinea: Coppejans, 1992: 406, fig. 8A-G; Coppejans et al., 1995: 80, fig. 11; Indian Ocean: Silva et al., 1996: 846; Belize: Littler & Littler, 1997: 109, fig. 155; Lord Howe Island: Kraft, 2000: 604, figs 34E, 35A; South Africa: Leliaert et al., 2001: 452, fig. 2; Wallis: Payri et al., 2002: 44, pl. 1 fig. 5; Hawaiian Islands: Abbott & Huisman, 2004: 124, fig. 46A-B. (Figs 71-72) Basionym: Fucus taxifolius Vahl, 1802: 36 (type locality: St. Croix, Virgin Islands). Material examined: Vairao, Tahiti, 3 Oct. 1997, leg. C. E. Payri, UPF 612; 20 Mar. 1998, leg. J. Orempuller, UPF 588, 589; Atituiti, Mangareva, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 799; Taravai, Gambier, 20 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 830, 831, 871; Rapa Island, Australs: Agairoa Bay, 2 Nov. 2002, leg. C. E. Payri, UPF 1892; off Tarakoi islet, 6 Nov. 2002, leg. C. E. Payri, UPF 2044; Tupua’I Bay, 2 Dec. 2002, leg. J. L. Menou, UPF 2444; Akatamiro Bay, 2 Dec. 2002, leg. J. L. Menou, UPF 2452. Thallus to 45 mm tall, yellowish-green, composed of a creeping stolon to 25 cm long and 1 mm in diameter, bearing erect, plumose branches at relatively wide intervals. Erect branches to 8 mm wide, bearing pairs of opposite branchlets to 4 mm long and 1 mm in diameter which are compressed, basally constricted, with tapered, upwardly curved tips. Remarks: Infrequent in the Society islands, more represented in islands of the Gambier and Australs. As discussed in N’Yeurt & Payri (2004: 374), some records 48 A. D. R. N’Yeurt & C. E. Payri Figs 66-71. 66. Caulerpa sertularioides (UPF 39). Habit of pressed plant. Scale = 10 mm. 67. Caulerpa sertularioides (UPF 2591). Detail of erect assimilator of freshly-collected plant. Note terete, pointed branchlets. Scale = 10 mm. 68. Caulerpa seuratii (UPF 41). Habit of typical plant from Hikueru Island, Tuamotu archipelago. Note densely spinulose stolon (arrowhead). Scale = 5 mm. 69. Caulerpa seuratii (UPF 45). Habit of larger, more lax plant from Marokau Island. Note densely spinulose stolon (arrowhead). Scale = 20 mm. 70. Caulerpa seuratii (UPF 41). Detail of pinnule apex, showing distichously divided ramuli with mucronate tips. Scale = 200 µm. 71. Caulerpa taxifolia (UPF 2452). Habit of pressed plant. Scale = 5 mm. of C. taxifolia from the South Pacific have been erronously ascribed to Caulerpa mexicana Sonder ex Kützing, a species genetically distinct from C. taxifolia (Olsen et al., 1998). True Caulerpa mexicana from the Caribbean is characterised by a flattened main axis bearing non-constricted pinnules; the fronds also have a darker green colour with a lustrous sheen not found in C. taxifolia. (Meinesz et al., 1994; Littler & Littler, 2000: 364, fig. on p. 365, upper). A main morphological 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 French Polynesian green algae 49 per shoot in C. mexicana. Hawaiian records of C. mexicana (Taylor, 1977) have been transferred to C. taxifolia (Abbott & Huisman, 2004: 124), and C. mexicana would seem not to occur in the South Pacific (A. Meinesz, pers. com.). *Caulerpa taxifolia (Vahl) C. Agardh ecad. tristichophylla Svedelius, 1906: 112, fig. 5. Fiji: Littler & Littler, 2003: 222 (“Caulerpa mexicana var. pluriseriata”); Wallis: N’Yeurt & Payri, 2004: 374. (Fig. 73) Misapplied names: Caulerpa taxifolia f. asplenioides Greville, 1853: figs 1-2, pl. I. French Polynesia: Weber-van Bosse (1898: 292; 1910: 2). — Caulerpa mexicana var. pluriseriata W.R. Taylor, 1975a: 77, fig. 1. Fiji: Littler & Littler, 2003: 222. Material examined: Taravai, Gambier, on coral heads on fringing reef, 20 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 875, 876, 877; Agairoa Bay, Rapa, in Sargassum beds, 2 Nov. 2002, leg. C. E. Payri, UPF 1893. Thallus to 45 mm tall, yellowish-green, composed of a creeping stolon to 25 cm long and 1 mm in diameter, bearing erect, plumose branches at relatively wide intervals. Erect branches to 8 mm wide, bearing sometimes opposite, mostly three-dimensional, radially arranged pinnules to 4 mm long and 1 mm in diameter which are compressed, basally constricted, with tapered, upwardly curved tips. Remarks: This ecad is distinguished from the typical form of C. taxifolia by its three-dimensional, often radial arrangement of pinnules on erect assimilators. Both opposite and radial arrangement of pinnules can be encountered on the same plant, even the same erect branches. This seems to be a shallow-water ecotype of C. taxifolia, rather than an infraspecific taxon (A. Meinesz, pers. com.). The combination Caulerpa taxifolia f. asplenioides was invalidated by Meinesz et al. (1994: 108) since it was based on Caulerpa asplenioides Greville, a misapplied name for Caulerpa mexicana. Plants from Fiji reported by Littler & Littler (2003) as Caulerpa mexicana var. pluriseriata would best be ascribed to C. taxifolia ecad. tristichophylla (see remarks above for C. taxifolia). Caulerpa urvilleana Montagne, 1845: 21 (“urvilliana”) (type locality: Toud Island, Torres Strait, Australia). French Polynesia: Payri et al., 2000: 98; Indonesia: Weber-van Bosse, 1898: 318, pl. 26 figs 7-12; Coppejans & Prud’homme van Reine, 1992: 686, figs 3B, 11B (as C. cupressoides ecad. urvilliana); Bikini Atoll: Taylor, 1950: 60, pls 31 fig. 1, pl. 32 fig. 1; Indian Ocean: Silva et al., 1996: 848; Fiji: N’Yeurt, 2001: 716, fig. 39 (“urvilliana”); Littler & Littler, 2003: 234. (Figs 74-75) Material examined: Toud Island, Torres Strait, Australia, n.d., leg. D. d’Urville, PC 10388 (holotype); Hao, 23 Dec. 1937, leg. C. H. Maggs, BM 841603; Taiaro, 24 Sep. 1992, leg. R. Galzin, UPF 50; Nengo Nengo, Jun. 1996, leg. J. M. Zanini, UPF 617; Kauehi, 21 Sep. 1995, leg. J. Orempuller, UPF 51; Ilot aux Oiseaux, Tikehau, 4 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 52; Takapoto, 7 Nov. 1995, leg. L. Addessi, UPF 47, 48, 49; Puka Puka, Aug. 1999, leg. Anon., UPF 764; Vairaatea, Oct. 2000, leg. Anon., UPF 771; Atituiti, Mangareva, 18 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 895, 907; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2464, 2465. Thallus to 12 cm high, with a creeping stolon to 40 cm long and 3 mm wide bearing sparse erect branches at irregular intervals. Upright branches slender, angular and terete, a few to several times dichotomously branched, or irregularly branched, 5-12 cm tall with axes smooth below, mamillate and spiny above, 1-2 mm in diameter. Assimilators thorn-like, radially aranged with mucronate tips. Remarks: A common species from the atolls, forms extensive, loose populations on sandy substratum at the bottom of lagoons at depths of 1 to 28 metres, and on the outer reef slope between 3 and 25 meters. Shallow-water populations tend to 50 A. D. R. N’Yeurt & C. E. Payri be smaller and bushier than deeper-water plants (Tuamotu). There has always been a confusion in the literature between C. urvilleana and C. cupressoides var. mamillosa. An examination of the type material of both species held in PC has shown them to be distinct, at least in gross external morphology. Although Montagne’s type of C. urvilleana from Toud Island, Northern Australia (Fig. 74) only consists of a single piece of erect assimilator, the slender branches with characteristic thorn-like, non-inflated mucronate pinnules distinguishes it from C. cupressoides var. mamillosa. The latter species also seems to possess a goldenyellow hue of the stolon and base of erect assimilators which is usually not found in plants of C. urvilleana, which are typically dark grass-green in colour. Caulerpa verticillata J. Agardh, 1847: 6 (type locality: West Indies). French Polynesia: Meinesz et al., 1981: 436, figs 26-27; Indonesia: Weber-van Bosse, 1898: 267, pl. 20 figs 7-10, Coppejans & Prud’homme van Reine, 1992: 708, fig. 21B; Sri Lanka: Svedelius, 1906: 108, fig. 1a-b; Trinidad: Børgesen, 1907: 19, figs 1-3; Ryukyu Islands: Yamada, 1934: 62, figs 31-32; Micronesia: Yamada, 1940: 97, Hodgson et al., 2004: 33; Viêt Nam: Dawson, 1954: 392, fig. 10b; Caribbean: W.R. Taylor, 1960: 138, pl. 10 figs 1-2; Caroline Islands: Trono, 1968: 167, pl. 14 fig. 4; New Caledonia: Valet, 1968: 42, pl. 6 fig. 4; Papua New Guinea: Coppejans & Meinesz, 1988: 194, figs 30-34, Littler & Littler, 2003: 234; Indian Ocean: Silva et al., 1996: 849; Belize: Littler & Littler, 1997: 109, fig. 156; Hawaiian Islands: Abbott & Huisman, 2004: 125, fig. 46C. (Fig. 76) Material examined: Opunohu Pass, Moorea, 22 Aug. 1978, leg. A. Meinesz, UPF 2990. Thallus 30-70 mm high, Branchlets terete and regularly dichotomously branched 3-5 times, clustered in verticillate whorls in distal parts of thallus. Creeping stolons naked, 225-375 µm in diameter, usually interwoven. Vertical axes 220-225 µm in diameter at the base, 70-85 µm in diameter at the top. Apices obtuse. Tufts of ramuli composed of 2-3 regularly disposed series of verticils, each including 3 or 4 ramuli 2.5-3 mm long and 25-30 (top) to 55-65 (base) µm in diameter, with 2 or 3 terminal indentations. Remarks: Growing abundantly on Halimeda sp., 30 metres depth under the ship wreck of Kersaint, in the reef pass. This alga seems to be relatively rare in French Polynesia, having been collected only once in the past on Moorea Island. Caulerpa webbiana Montagne, 1837: 354 (type locality: Arrecife, Isla Lanzarote, Islas Canarias). French Polynesia: Setchell, 1926: 84, Payri & N’Yeurt, 1997: 883, Payri et al., 2000: 100; Philippines: Meñez & Calumpong, 1982: 10, pl. 2G-J; Fiji: South & N’Yeurt, 1993: 124, fig. 15; N’Yeurt, 2001: 718, fig. 74; Littler & Littler, 2003: 236; Papua New Guinea: Coppejans et al., 1995: 81, figs 13-14 (as ecad disticha); Indian Ocean: Silva et al., 1996: 849. (Figs 77-78) Material examined: Punaruu, Tahiti, 11 Jul. 1922, leg. W. A. Setchell & H. E. Parks, BM 841523, SAP 037722; Afaahiti, Tahiti, 7 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 53; Punaauia, Tahiti, 6 Jul. 1997, leg. C. E. Payri, UPF 54; Tekokota, 1996, leg. J. Orempuller, UPF 56; Agairoa Bay, Rapa, 2 Nov. 2002, leg. C. E. Payri, UPF 1895; Tarakoi Islet, Rapa, 6 Nov. 2002, leg. C. E. Payri, UPF 2037; Patagaroa Pt., Rapa, 18 Nov. 2002, leg. J. L. Menou, UPF 2222, 2223; Rapa Iti, north slope, 28 Nov. 2002, leg. J. L. Menou, UPF 2406; Akatamiro Bay, Rapa, 2 Dec. 2002, leg. J. L. Menou, UPF 2453. Thallus dark green and spongy, 5-10 mm high, with a stolon to 7 cm long and 500 µm in diameter. Uprights branches 200-300 µm in diameter, irregularly branched 1-5 times and bearing whorls of fine branchlets to 300 µm long and 80 µm in diameter with mucronate tips, giving the uprights a furry appearance. The branchlets do not completely cover the stolon, which can be partly naked in places. French Polynesian green algae 51 Figs 72-77. 72. Caulerpa taxifolia (UPF 2044). Detail of erect assimilators with flattened, basally constricted branchlets. Scale = 2 mm. 73. Caulerpa taxifolia ecad. tristichophylla (UPF 876). Habit, showing both opposite and three-dimensional (arrow) erect assimilators on the same stolon. Scale = 5 mm. 74. Caulerpa urvilleana (MA 10388). Habit of Montagne’s Type from Toud Island, showing assimilators with thorn-like, non-inflated pinnules. Scale = 5 mm. Photographed with permission from the Museum National d’Histoire Naturelle, Paris. 75. Caulerpa urvilleana (UPF 907). Habit of plant from Mangareva Island, showing similarity to Montagne’s type from Northern Australia. Scale = 10 mm. 76. Caulerpa verticillata (UPF 2990). Habit of pressed plant, showing ramuli arranged in series of verticils. Scale = 3 mm. 77. Caulerpa webbiana (UPF 2037). Habit of pressed thallus. Scale = 3 mm. 52 A. D. R. N’Yeurt & C. E. Payri Remarks: An inconspicuous species, growing in crevices on the fringing and barrier reef and on the outer reef slope to a depth of 4 meters. Family Halimedaceae Link Genus Halimeda J.V. Lamouroux Recent molecular studies on this genus (Kooistra et al., 2002, Verbruggen & Kooistra, 2004, Kooistra & Verbruggen, 2005, Verbruggen et al., 2005a, b, 2006) have created upheavals in taxonomy, and have shown the need to re-describe several common Indo-Pacific Halimeda species. Key to the French Polynesian species of Halimeda (Important note: in order for this morphological key to be meaningful in distinguishing superficially similar-looking species, measurements of anatomical structures must represent the median of 10 measurements of the structures in question from a mid-thallus segment, as discussed in Verbruggen et al., 2005a). 1a. Holdfast always single, frequently massive and bulbous; plants growing erect in sand or mud; nodal filaments fused in a single unit with intercommunicating pores. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 1b. Holdfast(s) single or multiple, small, not bulbous; plants erect on hard substratum, forming massive clumps, or featuring a sprawling habit . . . . . 2 2a. Plant sprawling, holdfasts several, diffuse . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Plant erect or pendant, holdfast single, basal . . . . . . . . . . . . . . . . . . . . . . 6 3a. Secondary utricles clavate, up to 70 µm in diameter, extending into medulla; nodal filaments fused completely in pairs or triplets for a distance of 400-800 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gracilis 3b. Secondary utricles not clavate, less than 40 µm in diameter; nodal filaments fused in pairs (occasionally in triplets) for a distance of less than 150 µm 4 4a. Thallus in dense cushions or clumps, branched in perpendicular planes, segments distinctly ribbed, dull and coarse upon drying . . . . H. opuntia 4b. Thallus sprawling, in lax clumps, branching mostly in one plane . . . . . 5 5a. Segments slightly to distinctly keeled, not distorted, often smooth and shiny, middle and distal segments relatively large and often ivy-leaf shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. distorta f. hederacea 5b. Segments not keeled, often distorted, dull and coarse upon drying, distal segments relatively small, not ivy-leaf shaped . . . . . . . H. distorta f. distorta 6a. Thallus pendant, segments small, ribbed. Nodal filaments mostly fused in pairs for a short distance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. minima 6b. Thallus erect, segments not ribbed. Nodal filaments unfused, or fused in pairs or triplets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7a. Segments often trilobed; daughter segments emerging from isolated pits on distal segment edge; nodal medullary filaments closely adherent, never completely fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Segments never lobed; daughter segments emerging at any point from a continuous band along distal segment edge; nodal medullary filaments unfused or completely fused in pairs or triplets . . . . . . . . . . . . . . . . . . . . . . . 9 8a. Peripheral utricles less than 50 µm in diameter and 60 µm high, segments smooth, never ruffled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. melanesica 8b. Peripheral utricles above 67 µm in diameter and 90 µm high, segments rugose, often ruffled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. heteromorpha French Polynesian green algae 53 9a. Nodal filaments unfused, basal segment large and fan-shaped, carrying many (> 6) daughter segments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. micronesica 9b. Nodal filaments fused in pairs or triplets, basal segments not fan-shaped, carrying only a few daughter segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Secondary utricles not inflated; thallus showing an obvious gradient from large and darker brownish segments near the base to small and lighter, whitish green segments near the apices. . . . . . . . . . H. lacunalis 10b. Secondary utricles distinctly inflated; thallus not showing any obvious gradient in segment size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Thallus flaccid, lightly calcified; segments not wedge-shaped; peripheral utricles 38-43 µm in diameter, often merging into one another H. discoidea 11b. Thallus rigid, heavily calcified, segments wedge-shape to trapezoidal, peripheral utricles 28-32 µm in diameter, not merging into one another . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. taenicola 12a. Plant flaccid, anchored in mostly muddy substratum; branching generally in one plane; segments large, 12-30 mm wide and to 20 mm high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. macroloba 12b. Plant stiff, anchored in sand, branching in many planes; segments small, 6- 8 mm wide and to 6 mm high . . . . . . . . . . . . . . . . . . . . H. borneensis *Halimeda borneensis W.R. Taylor, 1975b: 81, figs 1-2 (type locality: Pulau Gaya, Cleland, North Borneo). Hillis-Colinvaux, 1980: 105, fig. 27; Fiji: Littler & Littler, 2003: 240; Wallis: N’Yeurt & Payri, 2004: 374. (Figs 79-80) Misapplied names (given by Verbruggen et al., 2005a, b): Halimeda incrassata f. ovata. French Polynesia: Payri & Meinesz, 1985b: 642, figs 4, 8, 12, 38-40. — Halimeda simulans Howe. French Polynesia: Setchell, 1926: 81; New Caledonia: Valet, 1968: 48, pl. 9(4) fig. 3; Hillis-Colinvaux, 1980: 103, fig. 26; Fiji: South, 1992: 10, figs 29-31, N’Yeurt, 2001: 724; Rotuma: N’Yeurt, 1996: 390, figs 76, 85; Belize: Littler & Littler, 1997: 113, fig. 163; Papua New Guinea: Coppejans et al., 2001: 411. Material examined: Punaruu, Tahiti, 11 Jul. 1922, leg. W. A. Setchell & H. E. Parks, BM 841073 (as H. simulans); Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2816, 2817; Tiahura, Moorea, 13 Jun. 1984, leg. C. E. Payri, UPF 2787; Ilot aux Oiseaux Tikehau, 6 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 98; Teahupoo, Tahiti, 10 Mar. 2002, leg. A. D. R. N’Yeurt, UPF 2828; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2519; Hotel Bora Bora Lagoon, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2569, Hotel Pearl Beach, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2577; Hotel Intercontinental, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2588; Hotel le Meridien, Bora Bora, 19 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2593; Tiahura, Moorea, leg. A. D. R. N’Yeurt, 28 Sep. 2004, UPF 2867; 29 Sep. 2004, UPF 2868; Punaauia, Tahiti, 9 Oct. 2004, leg. A. D. R. N’Yeurt, and V. Pouira, UPF 2829; Sheraton, Moorea, 18 Oct. 2004, leg. A. D. R. N’Yeurt, and V. Pouira, UPF 2830; Mataia, Tahiti, 11 Nov. 2004, leg. A. D. R. N’Yeurt, UPF 2824, 2825, 2826; Baie de Cook, Moorea, 2 Nov. 2005, leg. S. Hatosy, UPF 2723. Emergent portion (excluding holdfast) about 50 mm high; single basal rhizoidal holdfast bulbous and sand-infiltrated, 15 mm in diameter and 30 mm long (Fig. 79). Colour dark-green to grass-green, heavily calcified and rigid. Texture smooth, surface shiny. Basal segments fan-shaped, 6-15 mm wide and 6-8 mm high; about 1 mm thick, 7-14 mm wide and 6-10 mm high. Peripheral utricles obpyriform and usually flat-topped, 32-(45)-55 µm in diameter and 41-(54.5)-68 µm long (Fig. 80). Nodal medullary filaments 45-64 µm in diameter, fused in groups of three, sometimes two. Remarks: Growing in sandy areas on the fringing and barrier reefs, and in small tidepools. Halimeda borneensis was initially described quite narrowly, without encompassing the full morphological diversity of the species in the Indo-Pacific, 54 A. D. R. N’Yeurt & C. E. Payri leading to misidentifications in the past. The superficial morphological similarities (large bulbous holdfast, large tripartite segments) between H. borneensis from the Indo-Pacific and “true” H. incrassata from the Caribbean and Atlantic is likely a result of convergent evolution, since their DNA sequences and internal anatomy are quite distinct (Verbruggen et al., 2005a, b). Similarly, previous Pacific records of Halimeda simulans are referable to H. borneensis, since H. simulans is strictly an Atlantic species, and no specimens with a H. simulans DNA barcode were found in the Indo-Pacific (Kooistra et al., 2002: 134; Verbruggen et al., 2005a). Halimeda discoidea Decaisne, 1842: 102 (type locality: “Kamschatka”, Russia according to Decaisne, 1842 and Silva et al., 1987, but actual type locality unknown, since Kamschatka is an unlikely cold-temperate locality for this genus (Hillis, 1959: 353; Hillis-Colinvaux, 1980: 138; Lipkin & Silva, 2002: 68)). French Polynesia: Payri & Meinesz, 1985b: 642, figs 1, 5, 9, 35-36, Payri et al., 2000: 108; Hawaiian Islands: Egerod, 1952: 398, pl. 38, fig. 19b-d; Abbott & Huisman, 2004: 131, fig. 49B-E; Hillis, 1959: 352, pl. 2, fig. 5; pl. 5, fig. 11; pl. 6, fig. 11; pl. 7, figs 9-10; pl. 8, figs 5-8; pl. 11; Hillis-Colinvaux, 1980: 136, fig. 41; Fiji: South, 1992: 6, figs 15-17; N’Yeurt, 2001: 721, figs 23a-b, 34; Indian Ocean: Silva et al., 1996: 866; Rotuma: N’Yeurt, 1996: 386, figs 67, 79; American Samoa: Littler & Littler, 2003: 244. (Figs 81-82) Material examined: Maraa, Tahiti, 28 Sep. 1928, leg. C. Crossland, BM 841193; Beachcomber, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2854; Côte Est, Bora Bora, Apr. 1990, leg. C. E. Payri, UPF 88; Tikehau, Oct. 1999, leg. S. Golubic, UPF 2702; Rurutu, 18 Aug. 2000, leg. C. E. Payri, UPF 744; Rikitea, Mangareva, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 845; Motu Taraururoa, Gambier, 16 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 904; Atituiti, Mangareva, 19 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 909; Tiahura, Moorea, 29 Sep. 2004, leg. A. D. R. N’Yeurt, UPF 2860, 30 Sep. 2004 UPF 2988, UPF 2859, 1 Oct. 2004, In Herb. UPF. Thallus to 10 cm tall, with a single short stalk-like segment at the base; lightly calcified, light green to cream in colour (Fig. 81). Segments large and thin (up to 20 mm broad and 15 mm high) and in a single plane, mostly branching dichotomously; plant drying to a papery texture. Daughter segments emerging at any point from a continuous band along distal segment edge. Peripheral utricles hexagonal in surface view, between 38-43 µm in diameter, with neighbouring peripheral utricles often merging with one another (Fig. 82). Cortex generally two-layered; secondary utricles distinctly inflated, up to 128 µm in diameter, supporting 5 to 14 peripheral utricles. Nodal filaments united in pairs or triplets, entangled below the fusion and only slightly adhering. Remarks: Growing on the barrier reef flats and outer reef slope to a depth of 10 metres. Also grows on rocky shores (e.g. Afaahiti). A morphologically highly variable entity, which might contain cryptic entities (H. Verbruggen, pers. com.). Halimeda distorta (Yamada) Hillis-Colinvaux, 1968: 33, figs 4, 6(2) f. distorta f. inedit. French Polynesia: Kooistra & Verbruggen, 2005: 180, fig. 2f-j; HillisColinvaux, 1980: 120, fig. 34; Wallis: N’Yeurt & Payri, 2004: 374. (Figs 83-84) Basionym: Halimeda incrassata f. distorta Yamada, 1941: 119 (type locality: near Ponape, Ant Atoll, Caroline Islands). Micronesia: Yamada, 1944: 28, Pl. 4. Misapplied name (given by Kooistra & Verbruggen, 2005): Halimeda copiosa Goreau et Graham, 1967: 432, figs 1-10. Hawaiian Islands: Abbott & Huisman, 2004: 131, fig. 49A. Material examined: Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2846; îlot aux Oiseaux, Tikehau, 6 Nov 1995, leg. A. D. R. N’Yeurt, UPF 85; Punaauia, Tahiti, 8 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 86; Nuku Hiva, Marquesas, 1997, leg. J. Orempuller, UPF 597, 598, French Polynesian green algae 55 Figs 78-84. 78. Caulerpa webbiana (UPF 2037). Detail of pinnule apex. Scale = 50 µm. 79. Halimeda borneensis (UPF 2867). Habit of plant from Moorea, showing large bulbous holdfast. Scale = 10 mm. 80. Halimeda borneensis. Internal anatomy (c = surface view of cortex; s = medullary siphons; u = peripheral utricles). All scales = 100 µm. 81. Halimeda discoidea (UPF 89). Habit of plant from Marquesas. Scale = 10 mm. 82. Halimeda discoidea. Internal anatomy. All scales = 100 µm. 83. Halimeda distorta f. distorta (UPF 2518). Habit of plant from Bora Bora, showing contorted segments. Scale = 5 mm. 84. Halimeda distorta f. distorta. Internal anatomy. Scales: c, u = 50 µm, s = 100 µm. 56 A. D. R. N’Yeurt & C. E. Payri 600, 601; Nengo Nengo, Jun. 1996, leg. J.-M. Zanini, UPF 618; Tikehau, Oct. 1999, leg. S. Golubic, UPF 2699; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 858; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 902; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2467, 2468; Aquarium, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2503; Hotel Intercontinental, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2518; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2521; Hotel Bora Bora Lagoon, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2555; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2607; Tiahura, Moorea, leg. A. D. R. N’Yeurt, 28 Sep. 2004, UPF TH12, 29 Sep. 2004, UPF 2865, 30 Sep. 2004, UPF 2858. Thallus to 30 cm long, sprawling, branched in multiple planes with both prostrate and erect portions and multiple rhizoidal holdfasts (Fig. 83). Segments heavily calcified, 4-6 mm wide and 3-4 mm high, often ribbed or keeled, contorted, predominantly tripartite with three bundles of medullary siphons. Peripheral utricles hexagonal, 36-51 µm in diameter, separating but not becoming rounded after decalcification; up to 4 supported by secondary utricles (Fig. 84). Secondary utricles 19-31 µm in diameter. Nodal siphons joined usually in pairs or triplets, fused for a distance of about 100-160 µm. Remarks: New molecular data showed that previous Pacific records of H. copiosa are referable to H. distorta, as the former is exclusively an Atlantic Ocean species (Kooistra & Verbruggen, 2005). Two morphs (f. distorta and f. hederacea) are present within this species; recent molecular evidence (Kooistra & Verbruggen, 2005) points to their being conspecific. Within the French Polynesian diversity, however, both morphs are morphologically distinct and therefore we describe them here as forms within a single species. H. distorta f. distorta is superficially characterised by having flat, dull contorted segments, which distinguishes it from f. hederacea which has keeled, shiny uncontorted segments. Halimeda distorta (Yamada) Hillis-Colinvaux f. hederacea (Barton) comb. inedit. (Figs 85-86) Heterotypic synonyms (given by Koistra & Verbruggen, 2005): Halimeda hederacea (Barton) Hillis, 1968: 30, figs 3, 6:1, 6:4-8. French Polynesia: Kooistra & Verbruggen, 2005: 180, fig. 2a-e; Fiji: Littler & Littler, 2003: 242; Wallis: N’Yeurt & Payri, 2004: 375. — Halimeda opuntia f. hederacea Barton 1901: 21, pl. 2, fig. 23 (type locality: Indonesia). — Halimeda opuntia var. hederacea (Barton) Hillis, 1959: 360, pl. 2 fig. 7, pl. 5 fig. 4. Bikini Atoll: Taylor, 1950: 81, pl. 40 fig. 1; Rotuma: N’Yeurt, 1996: 389, figs 70c, 84. Misapplied name (according to H. Verbruggen, pers. com.): Halimeda copiosa Goreau et Graham. Papua New Guinea: Coppejans et al., 1995: p. 84 fig. 21. Material examined: Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2846; Nuku-Hiva, Marquesas, 1997, leg. J. Orempuller, UPF 598; Taapuna, Tahiti, 8 Nov. 1998; leg. C. E. Payri, UPF 2874, 2875; Aquarium, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2503; Hotel Intercontinental, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2518; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2521; Tiahura, Moorea, 28 Sep. 2004, leg. A. D. R. N’Yeurt, UPF 2866, 30 Sep. 2004, UPF 2858. Thallus stiff and sprawling, to 38 cm long, moderately calcified, forming mats (Fig. 85). Branching in one plane, dichotomous or trichotomous. Segments of middle and distal regions relatively large and ivy-leaf in shape, to 10 mm wide and 6 mm high, with those near the base smaller, thick and tripartite. Segments with smooth and shiny surfaces when dried. Peripheral utricles polygonal in surface view, 28-46 µm in diameter, with up to four supported per secondary utricle (Fig. 86). Secondary utricles 19-30 µm in diameter. Nodal filaments united in pair or occasionally in triplets or sextuplets, fused for a distance of 84-116 µm. French Polynesian green algae 57 Figs 85-91. 85. Halimeda distorta f. hederacea (UPF 2866). Habit of plant from Bora Bora, showing characteristic large, ivy-leaf shaped segments (arrowhead). Scale = 5 mm. 86. Halimeda distorta f. hederacea. Internal anatomy. Scales: c, u = 50 µm, s = 100 µm. 87. Halimeda gracilis (UPF 93). Habit, showing straggling moniliform habit with subcuneate, centrally thickened segments and single basal rhizoidal holdfast (arrowhead). Scale = 5 mm. 88. Halimeda gracilis. Internal anatomy. All scales = 100 µm. 89. Halimeda heteromorpha (UPF 2810). Habit, showing typically soft and pliable texture, and single basal matted holdfast (arrowhead). Scale = 5 mm. 90. Halimeda heteromorpha (UPF 2809). Detail of segments of mid-thallus, showing rugose, pitted surface (arrowhead). Scale = 2 mm. 91. Halimeda heteromorpha. Internal anatomy. All scales = 100 µm. Remarks: The relatively large, keeled, ivy-leaf shaped segments branched in one plane are characteristic of this morph. See comments above for f. distorta. Halimeda gracilis Harvey ex J. Agardh, 1887: 82 (type locality: Sri Lanka). Barton, 1901: 22, pl. 3 figs 28-32. French Polynesia: Payri & Meinesz, 1985b: 642, figs 2, 6, 10, 37; Micronesia: Yamada, 1941: 116; Viêt Nam: Dawson, 1954: 396, fig. 13f; Brazil: Joly et al., 1968: 162, figs 2-4; Caroline Islands: Trono, 1968: 186, 58 A. D. R. N’Yeurt & C. E. Payri pl. 18 fig. 9; Hillis-Colinvaux, 1980: 144, fig. 44; Bahamas: Blair & Norris, 1988: 231, fig. 6; Papua New Guinea: Coppejans et al., 1995: 86, fig. 24; Indian Ocean: Silva et al., 1996: 867; South Africa: Leliaert et al., 2001: 453, figs 3, 9-10, De Clerck et al., 2005: 84, figs 57-58; Palau: Dragastan et al., 2002: 9, pl. 1 figs 6-8, pl. 3 figs 1-4, 6, 8; Fiji: Littler & Littler, 2003: 246; Hawaiian Islands: Abbott & Huisman, 2004: 133, fig. 49F. (Figs 87-88) Material examined: Makatea, Apr. 1982, leg. C. E. Payri, In Herb. UPF; îlot aux Oiseaux, Tikehau, 6 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 93; Tikehau, 7 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 92. Thallus with spreading habit, to 15 cm long, straggling with a main basal attachment point and several secondary attachments at points of contact with the substratum (Fig. 87). Segments subcuneate to reniform, moniliform, to 6 mm high and 10 mm broad and 500-750 µm thick, ribbed, seed-like (thick in central portion), moderately to heavily calcified, often with a glossy surface. Cortex 2-3 layered, peripheral utricles 28-53 µm in diameter, 40-90 µm long, slightly adherent in surface view when decalcified, up to 8 per secondary utricle (Fig. 88). Secondary utricles clavate (club-shaped), 23-70 µm in diameter, long and extending to medulla. Nodal filaments completely fused in pairs or triplets for a distance of 400-850 µm; fusion groups not entangled. Remarks: Growing at depths of 10-15 metres on the outer reef slope in atolls of the Tuamotu group. The gracile, straggling habit with small, bead-like segments are characteristic of this species. *Halimeda heteromorpha N’Yeurt in Verbruggen et al., 2006: 351, figs 14-26, 41-43, 51-54, 63, 64 (type locality: between Papetoai and Motu Tiahura, Moorea, French Polynesia). (Figs 89-91) Misapplied names (given by Verbruggen et al., 2006): Halimeda incrassata (Ellis) J.V. Lamouroux, 1816: 307. French Polynesia: Payri & Meinesz, 1985b: 642. — Halimeda incrassata f. ovata. French Polynesia: J. Agardh, 1887: 86; Barton, 1901: 27, pl. IV figs 42, 47; Hillis, 1959: 365. Material examined: Marquesas, n.d., leg. EPHE, UPF 96; Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2808, 2815; Tiahura, Moorea, 13 Jun. 1984, leg. C. E. Payri, UPF 2786, 2789, 2803; Punaauia, Tahiti, 25 Sep. 1995, leg. A. D. R. N’Yeurt, UPF 97; Taapuna, Tahiti, 8 Nov. 1998, leg. C. E. Payri, UPF 2869; Tikehau, Oct. 1999, leg. S. Golubic, UPF 2701; Tupai, 3 Jul. 2002, leg. C. Vermenot, UPF 2478; Aquarium, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2505, 2506; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2520; Hotel Bora Bora Lagoon, Bora Bora, 17 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2570; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2597; Tiahura, Moorea, 28 Sep. 2004, leg. A. D. R. N’Yeurt, 29 Sep. 2004, UPF 2857, 1 Oct. 2004, UPF 2810, 2814; Punaauia PK 18, Tahiti, 9 Oct. 2004, leg. A. D. R. N’Yeurt, and V. Pouira, UPF 2823; Punaauia PK 13, Tahiti, 20 Nov. 2004, leg. A. D. R. N’Yeurt, UPF 2809. Thallus to 55 mm high and 110 mm broad, with a single, circular, basal matted rhizoidal holdfast 15 mm in diameter (Fig. 89). Colour grass-green in upper half of thallus, whitish in lower portions. Moderately to lightly calcified. Basal portion hidden within crevice in the substratum. Texture soft and decumbent, segments highly pliable at nodes; surface rough, dull-pitted (Fig. 90). Branching dense throughout. First two basal segments 6-7 mm wide and 4.5-5 mm high; lower segments fan-shaped to cylindrical, 4-6 mm wide and 5-6 mm high. Middle segments tripartite to reniform, 5-7 mm wide and 4-5 mm high. Daughter segments emerging from isolated pits on distal segment edge. Peripheral utricles in middle portions of thallus elongate obovoid with usually rounded ends, 59-90 µm in diameter and 91-113 µm high (Fig. 91). Nodal medullary filaments 54-90 µm in diameter, adhering French Polynesian green algae 59 into a single unit at segment nodes; connecting pores between neighbouring segments at segment nodes totally absent or smallish, 25-36 µm in diameter. Remarks: Common on the fringing reefs and in the lagoon of high islands, nested amongst coral debris and cavities in ledges. Its flaccid habit with small segments could be confused in the field with H. melanesica, but the latter species has smaller peripheral utricle diameter and height. Based on new morphological and molecular evidence in Verbruggen et al. (2006) there are several species contained in what was previously considered as Halimeda incrassata. One species, which is exclusively present in the Caribbean, conforms to the type material of H. incrassata, while the other two are Pacific species which were described as new (H. kanaloana Vroom from Hawaii, and H. heteromorpha from the Indo-Pacific). This species was first recorded from Nuku-Hiva Island in the Marquesas by J. Agardh (1887) and later documented by Barton (1901), as H. incrassata f. ovata (H. Verbruggen, pers. com.). Halimeda lacunalis W.R. Taylor, 1950: 91, 208, pl. 51 figs 1-2 (type locality: Eniwetok Atoll, Marshall Islands) f. lata (W.R. Taylor) Hillis-Colinvaux, 1980: 129, fig. 38b. French Polynesia: Taylor, 1973: 38 (Tikehau); Payri & Meinesz, 1985b: 643, figs 13, 17, 21, 43, Payri et al., 2000: 110; Hillis-Colinvaux, 1980: 129, fig. 38a; Indian Ocean: Silva et al., 1996: 868. (Figs 92-93) Material examined: Mataiva, Tuamotu, Apr. 1982, leg. C. E. Payri, UPF 2805; Nengo Nengo, Jun. 1996, leg. J.-M. Zanini, UPF 616; Marokau, Tuamotu, leg. J. Orempuller, 22 Nov. 1996, UPF 99, 100. Thallus erect or hanging, flaccid, up to 18 cm tall, fairly compact, arising from a minute holdfast (Fig. 92). Calcification light to moderate, whitish-green. Branching mainly planar and commonly dichotomous, several segments arising from a single one; basal segments small, cylindrical to subcuneate, upper segments discoidal to reniform, to 15 mm long and 20 mm broad, and 0.5-0.7 mm in thickness. Daughter segments emerging at any point from a continuous band along distal segment edge. Dried plants show an obvious gradient from large and darker brownish segments near the base to small and lighter, whitish green segments near the apices. Cortex 2-4 layered, peripheral utricles hexagonal in surface view, 20-55 µm in diameter, remaining attached after decalcification (Fig. 93). Secondary utricles 15-50 µm broad, not inflated, bearing up to 5 peripheral utricles. Nodal filaments united in pairs or triplets for a distance of about 80-150 µm. Remarks: Growing on coral heads in the lagoon of atolls. Two forms within this species were described by Hillis-Colinvaux (1980): f. lacunalis and f. lata. The latter is more compact, and more commonly encountered. The French Polynesian plants conform to f. lata. Halimeda macroloba Decaisne, 1841: 118 (type locality: Red Sea). French Polynesia: Payri & Meinesz, 1985b: 643, figs 14, 19, 22-23, 44-45, Payri et al., 2000: 112; Hillis, 1959: 375, pl. 3 fig. 3, pl. 5 figs 19-20, pl. 6 fig. 17, pl. 12; Thailand: Egerod, 1974: 148, figs 65-68; 1975: 61, fig. 33; Hillis-Colinvaux, 1980: 108-110, fig. 28; Fiji: South, 1992: 8, figs 12-14, N’Yeurt, 2001: 722, fig. 38; Papua New Guinea: Coppejans et al., 1995: 86, fig. 25; Indian Ocean: Silva et al., 1996: 869; American Samoa: Littler & Littler, 2003: 248; Hawaiian Islands: Abbott & Huisman, 2004: 135, fig. 50B. (Figs 94-95) Material examined: Hotel Beachcomber, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2853; 4 May 1997, leg. A. D. R. N’Yeurt, UPF 101, 102; Taapuna, Tahiti, leg. C. E. Payri, UPF 2873. 60 A. D. R. N’Yeurt & C. E. Payri Figs 92-97. 92. Halimeda lacunalis f. lata (UPF 100). Habit, showing gradient from large and darker brownish segments near the base to small and lighter, whitish green segments near the apices. Scale = 10 mm. 93. Halimeda lacunalis f. lata. Internal anatomy. All scales = 100 µm. 94. Halimeda macroloba (UPF 2873). Habit, showing thick large, epiphyte-laden unribbed segments and sandincrusted bulbous holdfast. Scale = 10 mm. 95. Halimeda macroloba. Internal anatomy. All scales = 100 µm. 96. Halimeda melanesica (UPF 2804). Habit of plant from Afaahiti, Tahiti. Scale = 10 mm. 97. Halimeda melanesica. Internal anatomy. Scales: c, u = 50 µm, s = 100 µm. Thallus up to 15 cm high and 20 cm broad, with a large bulbous and incrustated holdfast 30-40 mm long and 15-20 mm wide (Fig. 94). Segments thick and large, subcuneate to reniform, not ribbed, about 1 to 2 mm thick and up to 30 mm wide and 20 mm high. Branching planar to bushy, di- to polychotamous. French Polynesian green algae 61 Calcification moderate; basal segment compressed and supporting several separate or laterally consolidated segments forming a fan-shaped unit. Cortex 3-4 layered, mature peripheral utricles 25-45 µm in diameter, rounded and remaining separate in surface view following decalcification (immature segments may have peripheral utricles remaining attached following decalcification, see Payri & Meinesz 1985b) (Fig. 95). Secondary utricles 20-58 µm in diameter and 40-80 µm long. Two to four peripheral utricles supported per secondary utricles. Nodal filaments completely united for about 44-80 µm, communicating by pores. Remarks: This species has the largest and thickest segments, and is commonly found in calm areas subject to much sedimentation (e.g. in the lagoons and estuaries). Despite its conspicuous large segments and massive holdfast, this species has never been collected to date outside of Tahiti and Moorea (Society Group), perhaps due to the unavailability of its typical habitat elsewhere. The broad segments are often the host to a number of creeping epiphytic algae (e.g. Ceramium spp., Chondria spp.). *Halimeda melanesica Valet, 1966b: 683, fig. 2, pl. 1 figs c-d (type locality: Luengöni, Lifou Island, Loyalty Islands, New Caledonia). New Caledonia: Valet, 1968: 49, pl. 9 fig. 3; Hillis-Colinvaux, 1980: 153, fig. 48; Fiji: Littler & Littler, 2003: 250; Verbruggen & Kooistra, 2004; Verbruggen et al., 2006, figs 23-29, 37-39, 48-49. (Figs 96-97) Material examined: Taiohae, Nuku Hiva, Marquesas, c. 1899, leg. E. Jardin, BM 841150; Marquesas, n.d., leg. EPHE, UPF 95; Nuku Hiva, Marquesas, 1997, leg. J. Orempuller, UPF 599, 602, 603; Afaahiti, Tahiti, 7 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 94; Taiohae, Nuku Hiva, Marquesas, 30 Apr. 2002, leg. S. Sidolle, UPF 2714; Afaahiti, Tahiti, 16 Nov. 2004, leg. A. D. R. N’Yeurt & A. Pham, UPF 2804, 2811, 2812, 2813, 2821, 2832. Thalli in flaccid clumps to 12 cm high, attached to the substratum by means of a firm, dense mat of rhizoids (Fig. 96). Segments lightly calcified, consisting of a basal zone of relatively large segments 5 mm long to 9 mm wide, merging into a fan-like structure that carries several branches, with progressively smaller segments to 4 mm long and 3 mm broad, and 500 µm thick. Daughter segments emerging from isolated pits on distal segment edge. Segments from central and upper thallus parts obovate–cuneate, with the majority trilobed, surface generally smooth and shiny, never ruffled or pitted. Cortex consisting of 3 layers of basally-constricted utricles (Fig. 97). Peripheral utricles 40–50 µm in diameter and 40–53 µm high, adherent after decalcification. Median secondary utricles 32–45 µm in diameter and 49–92 µm high. Nodal filaments closely adherent over a distance of 25-30 µm, showing an adhesion band when dissected; sometimes fused in pairs or triplets. Pores small or absent. Remarks: Growing in clumps on rocky substratum at depths of 2-5 metres, so far in French Polynesia only reported from Nuku Hiva in the Marquesas and Afaahiti District in Tahiti. The species seems to be restricted to wave-washed, exposed habitats, and is known to occur from infralittoral fringe, wave-affected sites to deeper waters at sites with strong swells. Superficially similar to H. heteromorpha, H. melanesica can be distinguished by its restriction to wave-washed, exposed areas or deeper waters exposed to strong swell. The segments appear smoother in H. melanesica. Definite identification relies on measurement of the average diameter of the peripheral utricles, which is less than 50 µm in H. melanesica. Upon drying, H. melanesica also tends to take on a pale brownish-yellow colour whereas H. heteromorpha keeps its green colour in the apical thallus parts. The present distribution of the species at opposite ends of the South Pacific (New Caledonia, Fiji, Tahiti) urges more research to locate it in intervening localities. 62 A. D. R. N’Yeurt & C. E. Payri Halimeda micronesica Y. Yamada, 1941: 121, fig. 15; 1944: 29, pl. 5 (type locality: Ant atoll, Ponape Island, East Carolines). French Polynesia: Payri & Meinesz, 1985b: 643, figs 16, 18, 24, 46, Payri et al., 2000: 112; Bikini Atoll: Taylor, 1950: 89, pl. 46, fig. 2; pl. 47; Hillis, 1959: 364, pl. 3, fig. 1; pl. 5, figs 13-14; pl. 6, fig. 2; pl. 9; Caroline Islands: Trono, 1968: 186, pl. 17 fig. 6; Ryukyu Islands: Itono, 1973: 160, fig. 21, Tsuda & Kamura, 1991: 71, pl. 4 figs 2-3; Hillis-Colinvaux, 1980: 149; Maldives: Wynne, 1993: 22, fig. 10; Indian Ocean: Silva et al., 1996: 869; Rotuma: N’Yeurt, 1996: 387, figs 69, 81-82; Fiji: Littler & Littler, 2003: 250; Wallis: N’Yeurt & Payri, 2004: 375. (Figs 98-99) Heterotypic synonym (given by Silva et al., 1996): Halimeda orientalis Gilbert, 1947: 126, fig. 1 (Philippines). Material examined: Makatea, Apr. 1982, leg. C. E. Payri, UPF 2818; Kauehi, 22 Sep. 1995, leg. J. Orempuller, UPF 103; îlot aux Oiseaux, Tikehau, 6 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 104; îlot Ono, Bora Bora, Apr. 1990, leg. C. E. Payri, UPF 106; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2466, 2469, 2472; Tupai, 3 Jul. 2002, leg. C. Vermenot, UPF 2477, 2480, 2485. Plants compact and bushy, to 10 cm tall (Fig. 98). Rhizoids long and fibrous, occurring in the basal region and often at the tips of the branches. Calcification moderate, colour light green to whitish. Basal segment conspicuous and large, to 18 mm broad and 12 mm high, reniform (fan-shaped), with an undulate margin, supporting numerous smaller segments. Branching mainly trichotomous, in one plane. Other segments 9 mm broad and 7 mm high, frequently trilobed. Cortex 3-4 layered, peripheral utricles rounded in surface view, 28-48 µm in diameter, usually separating after decalcification; 2-4 borne per secondary utricles (Fig. 99). Secondary utricles 15-45 µm in diameter, generally not basally constricted. Nodal filaments unfused, sometimes slightly adherent, with thickened and pigmented filament walls. Remarks: Growing usually in the lagoon and on the inner reef and outer reef slope of atolls in the Tuamotu archipelago, but also found in the Society Group in Bora Bora (outer reef flat) and in the atoll of Tupai, which presumably have similar habitats. Specimens were found forming tight clumps on the reef crest, with waves breaking right on top of them (Rangiroa). The single, fused and fan-shaped basal segment of this species is distinctive in the field. Characteristic rope-like uncorticated medullary filaments extend 5-6 cm from some basal segments over the substratum, bearing a young plant at the end. This represents a mode of vegetative reproduction in this species, as well as a means of added attachment to the substratum (HillisColinvaux, 1980; Wynne, 1993). Halimeda minima (W.R. Taylor) Hillis-Colinvaux, 1968: 32, figs 5-6. French Polynesia: Payri & Meinesz, 1985b: 644, figs 16, 20, 25, 47, Payri et al., 2000: 114, Kooistra & Verbruggen, 2005: 178, fig. 1a-b; Hillis-Colinvaux, 1980: 113, fig. 30; Fiji: South 1992: 8, fig. 34, N’Yeurt, 2001: 723, figs 25, 36, Littler & Littler, 2003: 250; Indian Ocean: Silva et al., 1996: 870. (Figs 100-101) Basionym: Halimeda opuntia (Linnaeus) J.V. Lamouroux f. minima W.R. Taylor, 1950: 82-83, 206, pl. 39 fig. 2 (type locality: Bikini Atoll, Marshall Islands). Material examined: Fare Ute, Tahiti, 30 May. 1922, leg. W. A. Setchell & H. E. Parks, BM 841221; Tiahura, Moorea, Aug. 1978, leg. A. Meinesz, UPF 2847; Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2848; Afaahiti, Tahiti, 7 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 105; Taapuna, Tahiti, 8 Nov. 1998, leg. C. E. Payri, UPF 2870; Motu Totegegie, Gambier, 22 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 786; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 890; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2597; Tiahura, Moorea, leg. A. D. R. N’Yeurt, 29 Sep. 2004, UPF 2851, 30 Sep. 2004, UPF 2850, 2852, 3 Nov. 2004, UPF 2849. French Polynesian green algae 63 Figs 98-103. 98. Halimeda micronesica (UPF 2818). Habit, showing characteristic fan-shaped basal segment (arrow) and rope-like medullary filaments extended from basal segments (arrowhead). Scale = 5 mm. 99. Halimeda micronesica. Internal anatomy. All scales = 100 µm. 100. Halimeda minima (UPF 105). Habit, showing single basal holdfast (arrowhead). Scale = 5 mm. 101. Halimeda minima. Internal anatomy. All scales = 100 µm. 102. Halimeda opuntia (UPF 924). Habit. Scale = 5 mm. 103. Halimeda opuntia. Internal anatomy. All scales = 100 µm. 64 A. D. R. N’Yeurt & C. E. Payri Plants bushy, 5-7 cm tall, of pendant habit, with a single minute holdfast about 6 mm high and 3 mm wide (Fig. 100). Segments relatively small, strongly calcified and brittle, 2-4 mm wide and 1-4 mm high, frequently trilobed in lower portions of thallus; rugose-pitted in appearance. Cortex up to four-layered; peripheral utricles 15-25 µm in diameter, subhexagonal and adhering slightly after decalcification, more than 4 borne per secondary utricle (Fig. 101). Secondary utricles 10-20 µm in diameter. Nodal filaments united in pairs or triplets for a distance of 56-75 µm; occasionally separate. Remarks: Growing in crevices on the barrier reef flat, and pendant from ledges on the outer reef slope to a depth of 20 meters. Halimeda minima can be readily distinguished from members of the H. hederacea-distorta-opuntia complex (with which it shares many characters such as incomplete nodal fusion and small, heavily calcified segments) by its pendant habit and single, basal holdfast. Halimeda opuntia (Linnaeus) J.V. Lamouroux, 1816: 308. French Polynesia: Payri & Meinesz, 1985b: 644, figs 26, 29, 32, 48, 49, Payri et al., 2000: 114; Kooistra & Verbruggen, 2005: 179, fig. 1c-e; Barton 1901: 18, pl. 2 figs 19-27; Hawaiian Islands: Egerod, 1952: 397, pl. 3, fig. 19a, e-f, Abbott & Huisman, 2004: 135, fig. 50C; Bikini Atoll: Taylor, 1950: 80, pl. 39, fig. 1; Viêt Nam: Dawson, 1954: 395, fig. 12; Hillis, 1959: 359, pl. 2, figs 7-8; pl. 5, figs 3-4; pl. 6, fig. 6; pl. 7, fig. 3; pl. 10; Caroline Islands: Trono, 1968: 178, pl. 18 Figs 1-4; Thailand: Egerod, 1974: 147, figs 59-61; Somalia: Sartoni, 1979: 284, fig. 4c; Hillis-Colinvaux, 1980: 110, figs 19, 51, 92; Philippines: Trono, 1986: 234, fig. 35; Ryukyu Islands: Tsuda & Kamura, 1991: 65, pl. 2 fig. 4; Maldives: Wynne, 1993: 23, fig. 11; Papua New Guinea: Coppejans et al., 1995: 86, fig. 27; Indian Ocean: Silva et al., 1996: 870; Rotuma: N’Yeurt, 1996: 389, figs 70b, 83; Belize: Littler & Littler, 1997: 111; Fiji: South, 1992: 9, figs 26-28; N’Yeurt, 2001: 723, Littler & Littler, 2003: 252; Samoa: Skelton & South, 2002a: 165, fig. 25C-D. (Figs 102-103) Basionym: Corallina opuntia Linnaeus, 1758: 805 (type locality: Jamaica). Material examined: Tahiti, Sep. 1875, leg. H. Moseley, BM 841222, 841231; Hao, 10 Jan. 1905, leg. L. G. Seurat, BM 841220; Tahiti, Oct. 1909, leg. J. E. Tilden, BM 701523; Maraa, Tahiti, 24 Sep. 1928, leg. C. Crossland, BM 701522; Tiahura, Moorea, Jun. 1980, leg. C. E. Payri, UPF 2845; Taapuna, Tahiti, 16 May 1997, leg. A. D. R. N’Yeurt, UPF 107; Mt. Mokoto, Mangareva, 27 Sep. 1997, leg. J. Starmer, UPF 630; Aukena, Gambier, 21 Sep. 1997, leg. J. Starmer, UPF 632; Hikueru, 16 Nov. 1996, leg. J. Orempuller, UPF 706; Raivavae, 18 Oct. 2000, leg. V. Clouard, UPF 787; Mekiro, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 911; Mangareva, Gambier, leg. A. D. R. N’Yeurt, 18 Nov. 2000, UPF 783, 21 Nov. 2000, UPF 785, 905; Taravai, Gambier, 20 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 925, 930; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2471; Povai, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2544; Tikehau, Oct. 1999, leg. S. Golubic, UPF 2699; Tiahura, Moorea, leg. A. D. R. N’Yeurt, 28 Sep. 2004, UPF 2862, 2863, 29 Sep. 2004, UPF 2864, 1 Oct. 2004, UPF 2861. Thallus forming dense or lax bushy clumps to 30 cm in diameter and 6 cm high, with multiple rhizoidal attachment points, light to dark green in colour (Fig. 102). Branching dense and in perpendicular planes. Segments 10 mm broad to 5 mm high, reniform, ribbed and trilobed at base of plant. Basal segments often bleached white. Cortex up to 5-layered; peripheral utricles 10 to 12 µm in diameter, small, rounded and slightly adhering in surface view following decalcification (Fig. 103). Secondary utricles 11-35 µm in diameter, slender and fork-shaped, arising as dichotomies of the medullary filaments. Nodal filaments united in pairs, occasionally in triplets, for a distance of 25-70 µm. French Polynesian green algae 65 Remarks: Growing in shallow waters on the reef flat. One of the commonest Halimeda species, recognized in the field by its clumped habit and reniform, ribbed segments which are characteristically branched in perpendicular planes Halimeda taenicola W.R. Taylor, 1950: 86, pl. 46 fig. 1. French Polynesia: Payri & Meinesz, 1985b: 645, figs 28, 31, 34, 51, Payri et al., 2000: 116; Hillis, 1959: 354, pl. 2 fig. 6, pl. 5 fig. 12, pl. 6 fig. 14, pls 11, 14; Trono, 1968: 182, pl. 16 fig. 3; HillisColinvaux, 1980: 139, fig. 42; Philippines: Trono, 1986: 236, fig. 37; Maldives: Wynne, 1993: 23, fig. 12; Indian Ocean: Silva et al., 1996: 872; Rotuma: N’Yeurt, 1996: 390, figs 72, 74, 86; Fiji: South, 1992: 10, figs 32-33; Littler & Littler, 2003: 252. (Figs 104-105) Material examined: Tiahura, Moorea, Aug. 1978, leg. A. Meinesz, UPF 2855; Maeva, Tahiti, Apr. 1982, leg. C. E. Payri, UPF 2856; Makatea, Apr. 1982, leg. C. E. Payri, UPF 2806, 2807; Kauehi, 22 Sep. 1995, leg. J. Orempuller, UPF 108; îlot aux Oiseaux, Tikehau, 6 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 109; Taapuna, Tahiti, 8 Nov. 1998, leg. C. E. Payri, UPF 2871, 2872; Tikehau, Oct. 1999, leg. S. Golubic, UPF 2704; Akamaru, Gambier, 17 Nov. 2000, leg. A. D. R. N’Yeurt, UPF 903; Motu aux récifs, Rangiroa, 20 Jul. 2002, leg. A. D. R. N’Yeurt, UPF 2470; Tupai, 3 Jul. 2002, leg. C. Vermenot, UPF 2476, 2479, 2486; Fangatau, 17 May 2003, leg. C. E. Payri, UPF 2706. Thallus light green to yellowish, to 8 cm tall, with a small fibrous holdfast (Fig. 104). Branching in one plane; segments to 2 mm thick. Lower segments often fused; upper segments to 8 mm wide and 10 mm high, deltoid to reniform, becoming concave when dried. Daughter segments emerging at any point from a continuous band along distal segment edge. Peripheral utricles hexagonal in surface view, 28-32 µm in diameter, remaining attached after decalcification (Fig. 105). Secondary utricles often markedly inflated, 26-60 µm in diameter and 50-120 µm long, bearing 4-6 peripheral utricles. Nodal filaments completely fused in pairs or triplets for a distance of 40-70 µm. Remarks: Growing on the barrier reef, on the reef crest, in the passes and on the outer slope to a depth of 10-25 meters. The thick, basally fused and heavily calcified segments which become concave when dry, are characteristic of this species. Family Udoteaceae (Endlicher) J. Agardh Genus Avrainvillea Decaisne Key to the French Polynesian species of Avrainvillea 1a. Stipe slender and prominent, 2-3 mm in diameter and 20-40 mm long; blades gregarious, holdfast matted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1b. Stipe stout to almost lacking, 5-6 mm in diameter and 10-20 mm long; blades solitary, holdfast bulbous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Blade woven, never tufted; siphons bright orange, 30-50 µm in diameter, apices rounded, never clavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. erecta 2b. Blade usually tufted; siphons yellowish green or brown, not bright orange, 66-72 µm in diameter, apices rounded or clavate . . . A. obscura 3a. Blade thin and papery, margins lacerate; siphons 13-28 µm in diameter, some siphon apices pointed or torn away, others rounded; pseudocortex absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lacerata 3b. Blades spongiose, margins entire; all apices rounded; pseudocortex present, stipes poorly differentiated from blades, siphons 28-38 µm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ridleyi 66 A. D. R. N’Yeurt & C. E. Payri Figs 104-109. 104. Halimeda taenicola (UPF 2807). Habit, showing thick wedge-shaped segments. Scale = 10 mm. 105. Halimeda taenicola. Internal anatomy. All scales = 100 µm. 106. Avrainvillea erecta (UPF 774). Habit. Scale = 5 mm. 107. Avrainvillea erecta (UPF 774). Detail of siphon dichotomy, showing deeply constricted, truncated dichotomy (arrowhead). Scale = 30 µm. 108. Avrainvillea lacerata (BM 515992). Habit of isotype from Vavau, Tonga. Scale = 5 mm. 109. Avrainvillea lacerata (BM 515992). Detail of blade anatomy of isotype, showing slightly constricted equal dichotomy (double arrowheads), pointed or torn-away apices (arrows) and rounded apex (arrowhead). Scale = 15 µm. French Polynesian green algae 67 Avrainvillea erecta (Berkeley) A. Gepp et E.S. Gepp, 1911: 29-32, pl. X fig. 89. French Polynesia: Payri et al., 2000: 118; Ryukyu Islands: Yamada, 1934: 73, fig. 41; Palau: Kanda, 1944: 744, fig. 4; Viêt Nam: Dawson, 1954: 395, fig. 13a; Caroline Islands: Trono, 1968: 174, pl. 19 fig. 9; New Caledonia: Valet, 1968: 50, pl. 11(6) fig. 5; Thailand: Egerod, 1975: 60, figs 30-32; Olsen-Stojkovich, 1985: 22, fig. 11, pl. 3a; Indonesia: Coppejans & Prud’homme van Reine, 1989a: 123, pl. 2 figs 18-37; Indian Ocean: Silva et al., 1996: 876; Fiji: N’Yeurt, 2001: 725, fig. 71; Solomon Islands: Littler & Littler, 2003: 238. (Figs 106-107) Basionym: Dichonema erectum Berkeley, 1842: 157, pl. VII fig. 11 (type locality: Philippines Islands). Material examined: Faa’a Channel, Tahiti, 18 Jan. 1926, leg. C. Crossland 6704, UC 667813; Botanical gardens, Papeari, Tahiti, 27 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 7; Temae, Moorea, 12 Nov. 2000, leg. C. E. Payri, UPF 774. Thallus solitary, to 60 mm high, consisting of a flabellate, relatively thick and felted reniform blade 3-4 cm wide and a stout unbranched stipe 10-20 mm long and 4-5 mm wide (Fig. 106). Rhizomatous holdfast bulbous and sandencrusted, 10-15 mm in diameter and 40-80 mm long. Margins smooth, faintly zonate and not lacerate. Siphons bright orange to yellowish brown, 30-50 µm in diameter; cylindrical with moderately to deeply constricted, truncated equal dichotomies (Fig. 107). Siphon apices rounded. Remarks: Growing in sandy substratum in the lagoon and outer reef (Moorea). The solitary habit, large bulbous holdfast and distinct yellow-orange siphons of this species set it apart from the other French Polynesian members of the genus. Coppejans et al., (2001) group A. erecta with A. obscura, pending further studies. Avrainvillea lacerata Harvey ex J. Agardh, 1887: 54 (type locality: Tonga). French Polynesia: Setchell, 1926: 81, Payri et al., 2000: 118; Marshall Islands: Dawson, 1957: 108, fig. 11b; Caroline Islands: Trono, 1968: 175, pl. 19 fig. 8; New Caledonia: Valet, 1968: 50, pl. 10(5) fig. 6; Indonesia: Coppejans & Prudhomme van Reine, 1989a: 125, pl. 2 f igs 1-17; Bahamas: Norris & Olsen, 1991: 321, fig. 5; Indian Ocean: Silva et al., 1996: 877; Papua New Guinea: Coppejans et al., 2001: 419; Fiji: Littler & Littler, 2003: 238. (Figs 108-112) Material examined: Vavau, Tonga, c. 1855, leg. W. H. Harvey, BM 515989 (co-type); Otepa, 28 Nov. 1904, leg. L. G. Seurat, BM 701631; Otepa, 29 Nov. 1904, leg. L. G. Seurat, BM 701632; Hikueru, Feb. 1905, leg. L. G. Seurat, BM 701633; Faarapa, Tahiti, 5 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 841682; Faarapa, Tahiti, 19 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 841687, 841690; Tahiti, May 1925, leg. C. Crossland 6663, UC 667654; Pa’ea, Tahiti, leg. C. Crossland 7027, 25 Sep. 1928, UC 341210; 28 Sep. 1928, Crossland 7066, UC 341172; Maraa, Tahiti, 28 Sep. 1928, leg. C. Crossland, BM 841684; Tahiti, 9 Sep. 1929, leg. C. Crossland 7293, UC 667852; Pa’ea, Tahiti, leg. C. Crossland, 7 Nov. 1929, UC 7245; Tahiti, 7 Dec. 1929, leg. C. Crossland 7246, UC 791864; Taapuna, Tahiti, 7 May 1997, leg. C. E. Payri, UPF 2, 5; Afaahiti, Tahiti, 7 Jun. 1997, leg. A. D. R. N’Yeurt, UPF 1; entre deux baies, Moorea, 11 Dec. 1995, leg. A. D. R. N’Yeurt, UPF 3, 4; Marokau, 22 Sep. 1996, leg. J. Orempuller, UPF 6; Aquarium, Bora Bora, 15 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2501; Mohio, Bora Bora, 21 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2599, 2600; Tiahura, Moorea, 29 Sep. 2004, leg. A. D. R. N’Yeurt, UPF 2965. Thallus gregarious, 50-130 mm high, greyish-green, consisting of papery thin, tightly woven wedge-shaped blades 25-50 mm high and 22-30 mm wide, borne from irregularly branched, distinct stipes 2-3 mm in diameter and 20-40 mm long arising out of a diffuse, spongy encrusting holdfast. Blade margins are lacerate, zonation sometimes present (e.g. BM 515990, Tongan isotype, Fig. 108). 68 A. D. R. N’Yeurt & C. E. Payri Siphons yellowish green, slightly torulose, 13-28 µm in diameter, cylindrical with slightly constricted equal dichotomies and mostly tapering, non-clavate pointed or rounded apices. Parts of the thallus with predominantly pointed or torn-away apices, but at least some siphons with rounded apices present throughout (Fig. 109). Remarks: Growing deeply wedged in crevices of coral heads in the lagoon, and on the outer reef slope to depths of 10 metres. According to Olsen-Stojkovich (1985), the paper-thin, tightly woven lacerate blades and pointed or torn-away siphon apices of A. lacerata distinguish it from A. amadelpha (Montagne) A. et E.S. Gepp, which has more loosely woven, never zonate, felt-like spongy blades with entire margins, and rounded siphon apices. Avrainvillea lacerata is also reported to differ by lacking a pseudo-cortex. However, these characters were found to be very variable in the French Polynesian material, as well as in Indonesian (Coppejans & Prud’homme van Reine, 1989a: 125) and Papua NewGuinean (Coppejans et al., 2001: 418), material of A. lacerata, since both rounded and pointed apices were found to co-exist in the same blade, which can at times be zonate. Both species form gregarious mats deeply lodged in cracks and crevices in coral. We have examined type material of A. lacerata from Vavau, Tonga in BM (515989), which had been cited by Olsen-Stojkovich (1985), and indeed found that both rounded and pointed apices are present (Fig. 109), prompting a re-evaluation of species concepts in Olsen’s paper. The same held true for a more recent specimen from Moorea Island (UPF 2965, Fig. 110). The only reliable characters seem to be the absence of pointed apices in A. amadelpha, and the consistently paper-like, non-spongy lacerate nature of the blade in A. lacerata. A specimen from Tahiti (UC 791865, Crossland 7245) determined as A. amadelpha by W. A. Setchell and cited as such by OlsenStojkovich (1985), was examined by us and found to consist of papery blades with some clearly pointed siphon apices present (Figs 111, 112) and was thus placed in A. lacerata, together with other Tahitian collections by C. Crossland determined as A. amadelpha. Interestingly, Coppejans et al. (2001) place together A. amadelpha with A. lacerata pending further studies, and both species could be merged in future, with A. lacerata being the earlier name. Previously considered a shallow-water species, this alga has been found growing as deep as 73 m in the Bahamas (Norris & Olsen, 1991). *Avrainvillea obscura (C. Agardh) J. Agardh, 1887: 53. Marshall Islands: W.R. Taylor, 1950: 67-68, pl. 34 fig. 1; Olsen-Stojkovich, 1985: 19, figs 9-10, pl. 2; Papua New Guinea: Coppejans et al., 1995: 88, fig. 29; 2001: 419; Indian Ocean: Silva et al., 1996: 877. (Figs 113-114) Basionym: Anadyomene obscura C. Agardh, 1823: 401 (type locality: Guam, Mariana Islands). Synonym (according to Olsen-Stojkovich, 1985): Avrainvillea capituliformis Tanaka, 1967: 14, figs 2-3 (type locality: Philippines). Japan: Tanaka & Itono, 1969: 4, pl. II figs 4-5, pl. III. Material examined: Tahiti, c. 1928, leg. C. Crossland 7199, UC 791872. Thallus solitary, 30-70 mm high including stout, short stipe 3-8 mm long (Fig. 113). Rhizomatous holdfast well-developed, 20-40 mm long and 6-15 mm wide, encrusted with sand. Blade olive to dark green, capitulate, 10-20 mm wide and 12-20 mm high, composed of loose-tufted, intermittently torulose filaments. Siphons olive to light brown, loosely woven to free, 66-72 µm in diameter, with deep, acutely constricted dichotomies. Apices rounded to clavate (Fig. 114). French Polynesian green algae 69 Remarks: The specimen label in UC lacks exact date, locality and habitat information. Initially determined by W. A. Setchell as a new species of Chlorodesmis (C. taitensis?), this specimen was later determined as a form of Avrainvillea obscura by J. O. Stojkovich (1979, annotation label on herbarium sheet). It represents the loose-tufted ecomorph of the species, and is distinguished from A. erecta by its larger diameter filaments with clavate ends, which are never yellow. It has not been recollected in Tahiti since Crossland’s survey. Interestingly, Coppejans et al. (2001) temporarily placed A. erecta under A. obscura. The French Polynesian material agrees well with Marshall Islands material described as A. obscura forma by W.R. Taylor (1950: 68) and Southern Japanese material initially described as A. capituliformis by Tanaka & Itono (1969). Avrainvillea ridleyi A et E.S. Gepp, 1911: 33, pl. XI figs 94-96 (type locality: Christmas Island, Indian Ocean). French Polynesia: Olsen-Stojkovich, 1985: 47, text-fig. 25, pl. 10A. Indian Ocean: Silva et al., 1996: 878. Material examined (according to Olsen-Stojkovich, 1985: 47): Tahiti, 7 Dec. 1929, leg. C. Crossland (7246), in UC. Thallus gregarious from emergent rhizoidal mat, 5 to 60 mm tall including highly variable, poorly differentiated stipe to 40 mm long. Stipes branched once or twice; blades spongiose, olive-brown, elongate-cuneate or rotundate, not zonate, with entire, rounded or lobed margins. Siphons olive green or brown in colour, 28-38 µm in diameter, becoming torulose in cortex, with rounded to subclavate apices. Dichotomies shallowly constricted. Remarks: This unusual species has not been recollected in French Polynesia since it was found in Tahiti by Crossland in 1929. The latter material, then in UC, was examined and confirmed as such by Olsen-Stojkovich (1985) but has unfortunately since been misplaced or lost (P. C. Silva, pers. com.) and no photographic record could be obtained either. Genus Chlorodesmis Harvey et Bailey Chlorodesmis fastigiata (C. Agardh) Ducker, 1969: 17, fig. 1. French Polynesia: Payri et al., 2000: 120; Indonesia: Coppejans & Prud’homme van Reine, 1989a: 127, pl. 3 figs 1-4, 12; Papua New Guinea: Coppejans et al., 2001: 421, figs 22-31; Indian Ocean: Silva et al., 1996: 879; Fiji: N’Yeurt, 2001: 726, fig. 73, Littler & Littler, 2003: 238; Wallis: Payri et al., 2002: 44, pl. 1 fig. 8; Samoa: Skelton & South, 2002a: 165, fig. 26B. (Figs 115) Basionym: Vaucheria fastigiata C. Agardh, 1824: 176 (type locality: Mariana Islands). Synonymy: Chlorodesmis comosa Harvey et Bailey, 1851: 373. French Polynesia: Taylor, 1973: 38 (Bora Bora, Moorea). Material examined: Papara, Tahiti, 4 Aug. 1912, leg. J. E. Tilden, BM 701544; Tahiti, 15 Aug. 1912, leg. J. E. Tilden, BM 841566; Papeete, Tahiti, 26 May 1922, leg. W. A. Setchell & H. E. Parks, BM 841569; Arue, Tahiti, 27 Jun. 1922, leg. W. A. Setchell & H. E. Parks, BM 841570; Maraa, Tahiti, 28 Sep. 1928, leg. C. Crossland, BM 841571; Taharaa, Tahiti, 14 Apr. 1997, leg. A. D. R. N’Yeurt, UPF 58, 59; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2524; Tiahura, Moorea, 1 Oct. 2004, leg. A. D. R. N’Yeurt, UPF TH89. Thallus up to 30 cm high, consisting of bright green tufts of free filaments, with a short basal spongy filamentous holdfast. Filaments cylindrical, 60-100 µm in diameter, distally dichotomously to trichotomously branched with unequal constrictions above dichotomies. 70 A. D. R. N’Yeurt & C. E. Payri Figs 110-115. 110. Avrainvillea lacerata (UPF 2965). Habit of dried plant from Moorea, showing thin, papery blades. Scale = 5 mm. 111. Avrainvillea lacerata (UC 791865). Habit of large plant from Tahiti, showing gregarious disposition of blades. Scale = 10 mm. 112. Avrainvillea lacerata (UC 791865). Detail of blade anatomy of specimen from Tahiti once placed in A. amadelpha (Olsen-Stojkovich 1985: 64, pl. 7A), showing slightly constricted equal dichotomy (double arrowheads), pointed apices (arrows) and rounded apex (arrowhead). Compare with Fig. 96. Scale = 15 µm. 113. Avrainvillea obscura (UC 791872). Habit of Tahitian loose-tufted ecomorph. Scale = 5 mm. 114. Avrainvillea obscura (UC 791872). Detail of blade anatomy, showing clavate apex (arrowhead). Scale = 70 µm. 115. Chlorodesmis fastigiata (UPF 59). Habit. Scale = 5 mm. Remarks: Growing in small crevices and tide pools, essentially on the exposed forereef flat and on the reef crest. In French Polynesia, this species was never found to reach more than 30 mm in height, although it is reported to attain sizes of up to 80 mm in other South Pacific localities (e.g. Fiji, Littler & Littler, 2003). This apparent form of dwarfism could be explained by the fact that in southeastern Polynesia (including the Cook Islands, N’Yeurt & Payri, in prep.) this species has never been found other than in wave-washed, exposed habitats on the fore-reef and reef crest. French Polynesian green algae 71 Genus Rhipidosiphon Montagne Rhipidosiphon javensis Montagne, 1842: 14-15 (type locality: Leiden Island (Nyamuk-besar), near Jakarta, Java, Indonesia). French Polynesia: Payri et al., 2000: 120; Caribbean: Littler & Littler, 1990: 35; Maldives: Wynne, 1993: 23, fig. 14; Indian Ocean: Silva et al., 1996: 882; Rotuma: N’Yeurt, 1996: 394, fig. 61; Papua New Guinea: Coppejans et al., 2001: 422, figs 32-34; Fiji: N’Yeurt, 2001: 726, fig. 33, Littler & Littler, 2003: 254; Hawaiian Islands: Abbott & Huisman, 2004: 140, fig. 52C-D. (Figs 116-117) Homotypic synonym: Udotea javensis (Montagne) A. et E.S. Gepp, 1904: 363. Hawaiian Islands: Egerod, 1952: 379, fig. 10; Viêt Nam: Dawson, 1954: 395, fig. 13b-c; Thailand: Egerod, 1975: 58, figs 28-29; Meinesz, 1980: 82, figs 1-2; Indonesia: Coppejans & Prud’homme van Reine, 1989a: 139, pl. 10 figs 3-9. Material examined: Tikehau, 7 Nov. 1995, leg. A. D. R. N’Yeurt, UPF 482L; Hikueru, 29 Sep. 1996, leg. J. Orempuller, UPF 131; Taaone, Tahiti, 7 May 1997, leg. C. E. Payri, UPF 130. Thallus yellow-green, up to 7 mm high and 5 mm broad, consisting of a calcified, fan-shaped blade with rounded outer margins and wedge-shaped base (Fig. 116); structure consisting of a single layer of parallel filaments 35-40 µm broad (outer margins) to 80-105 µm broad (base of thallus) with unequal constrictions above each dichotomy (Fig. 117). Stipe up to 200 µm in diameter, filamentous and uncorticated, mostly uncalcified and monosiphonous, anchored by fine, translucent rhizoids with clavate ends. Remarks: Growing on coral debris in the lagoon, at depths of 1-10 meters. The monostromatic blade with unequal siphon dichotomies is characteristic of the species. Order Dasycladales Pascher Family Dasycladaceae Kützing Genus Neomeris J.V. Lamouroux Key to the French Polynesian species of Neomeris 1a. Gametangia oblong, radially cohering in annular calcified rows. . N. annulata 1b. Gametangia spherical, individually calcified in discrete beads. . N. van-bosseae *Neomeris annulata Dickie, 1874: 198 (type locality: Mauritius). Sri Lanka: Svedelius, 1923: 452, figs 1-8; Ryukyu Islands: Yamada, 1934: 51, figs 16-17; Philippines: Gilbert, 1943: 19; Viêt Nam: Dawson, 1954: 396, fig. 13e; New Caledonia: Valet, 1969: 593, pl. 15 fig. 5, pl. 34 fig. 5, Indonesia: Coppejans & Prud’homme van Reine, 1989b: 127, figs 44-48; Indian Ocean: Silva et al., 1996: 889; Belize: Littler & Littler, 1997: 121, fig. 179; South Africa: Leliaert et al., 2001: 457, fig. 22; Samoa: Skelton & South, 2002: 166, fig. 26C; Fiji: Littler & Littler, 2003: 262; Hawaiian Islands: Abbott & Huisman, 2004: 144, fig. 55A-B. (Fig. 118) Material examined: Rarapai Islet, Rapa, 12 Nov. 2002, leg. J. L. Menou, UPF 2157. Thallus light-green, erect and cylindrical, 10-15 mm high and 3-5 mm in diameter, with discontinuous transverse basal rows of calcification, formed of rings of laterally-cohering gametangia. Individual gametangia elongate-ovoid, 100-180 µm long and 40-80 µm in diameter, pedicellate on a stalk cell with paired diamond-shaped assimilatory cells. Basal plug of stalk cell cap-like, without tit-like projection. 72 A. D. R. N’Yeurt & C. E. Payri Figs 116-121. 116. Rhipidosiphon javensis (UPF 130). Habit. Scale = 1 mm. 117. Rhipidosiphon javensis (UPF 482L). Detail of basal portion of plant from Tikehau, showing unequally constricted dichotomies (d). Scale = 50 µm. 118. Neomeris annulata (UPF 2157). Detail of elongate gametangia (g) atop stalk cell. Scale = 25 µm. 119. Neomeris vanbosseae (UPF 2722). Habit in situ, showing basal calcification around each individual gametangium (arrowhead). Scale = 4 mm. 120. Neomeris van-bosseae (UPF 2722). Detail of spherical gametangia (g) atop stalk cell, and plug with tit-like projection (arrowhead) into supporting cell. Scale = 25 µm. 121. Parvocaulis parvula (UPF RPS 55). Habit, showing loosely joined clavate segments and corona superior. Scale = 500 µm. French Polynesian green algae 73 Remarks: Growing at 12 m depth, on coralline substratum. This species is characterised by the annular rows of calcified gametangia, and the elongate shape of the gametangial cysts. So far in French Polynesia, it only occurs in the southern Australs, in Rapa. Neomeris van-bosseae M.A. Howe, 1909: 80, pl. 1 figs 4, 7; pl. 5 figs 17-19 (“van bosseae”) (type locality: Sikka, Flores, Indonesia). French Polynesia: Setchell, 1926: 80; Philippines: Gilbert, 1943: 17; Valet, 1969: 596, pl. 153 figs 4-7; Indonesia: Coppejans & Prud’homme van Reine, 1989b: 127, figs 63-71; Payri et al., 2000: 122; Hawaiian Islands: Egerod, 1952: 405, pl. 41, fig. 22b, Abbott & Huisman, 2004: 144, fig. 55C; New Caledonia: Valet, 1968: 52; Papua New Guinea: Coppejans et al., 1995: 100, fig. 41; Indian Ocean: Silva et al., 1996: 890; Rotuma: N’Yeurt, 1996: 396, figs 58, 90; Fiji: N’Yeurt, 2001: 731, Littler & Littler, 2003: 264; South Africa: De Clerck et al., 2005: 92, fig. 68. (Figs 119-120) Material examined: Hiti, 11 Nov. 1996, leg. J. Orempuller, UPF 125; Tekokota, 26 Sep. 1996, leg. J. Orempuller, UPF 126; Taharaa, Tahiti, 14 Apr. 1997, leg. A. D. R. N’Yeurt & C. E. Payri, UPF 124; Rurutu, 18 Aug. 2000, leg. C. E. Payri, UPF 728; Matira, Bora Bora, 16 Aug. 2002, leg. A. D. R. N’Yeurt, UPF 2523; Tiahura, Moorea, 1 Oct. 2004, leg. A. D. R. N’Yeurt, UPF 2728; Trou du Souffleur, Papenoo, Tahiti, 3 Mar. 2005, leg. A. D. R. N’Yeurt & A. Pham, UPF 2722. Thallus dark-green, erect and cylindrical, up to 20 mm tall and 4 mm in diameter, in a broad, 120-130° curve (Fig. 119). Basal portion whitish and moderately calcified around each gametangium giving a beady appearance, upper portion light to dark green with hair-like whorls of radial branchlets at the tip. Surface cells not organized in any distinct ring pattern. Gametangia spherical, 75-100 µm in diameter (Fig. 120), pedicellate on a stalk cell between paired diamond-shaped assimilatory cells. Plug at base of gametangial stalk cell discoid, usually with a tit-like projection into the supporting cell. Remarks: Growing at the side of tide pools and on coral debris in shallow water, on the reef flat and on the outer reef slope to a depth of 10 metres, in the Society and Tuamotu archipelagos. The individually-calcified, spherical cysts with plugs showing a basal projection are characteristic of this species. Family Polyphysaceae Kützing Genus Parvocaulis S. Berger, U. Fettweiss, S. Gleissberg, L.B. Liddle, U. Richter, H. Sawitsky et G.C. Zuccarello *Parvocaulis parvula (Solms-Laubach) S. Berger, U. Fettweiss, S. Gleissberg, L.B. Liddle, U. Richter, H. Sawitsky et G.C. Zuccarello, 2003: 559. (Fig. 121) Basionym: Acetabularia parvula Solms-Laubach, 1895: 29, pl. 2 figs 3, 5 (syntype localities: “Tropical India”; Celebes, Indonesia). New Caledonia: Valet, 1969: 621, pl. 11 figs 1-7; pl. 12 fig. 7; pl. 19 figs 2-4; pl. 20 figs 5-8; pl. 22 figs 1, 4, 7; pl. 29; pl. 38 figs 1, 4-5; pl. 45 figs 5-7; Indian Ocean: Silva et al., 1996: 893; Samoa: Skelton & South, 2002: 167, figs 26D-E, 27A-C; Hawaiian Islands: Abbott & Huisman, 2004: 147, fig. 56C. Homotypic synonym (given by Berger et al., 2003): Polyphysa parvula (SolmsLaubach) Schnetter et Bula-Meyer, 1982: 42, pl. 7 figs c-f (type locality: Celebes, Indonesia). French Polynesia: Payri et al., 2000: 122; Rotuma: N’Yeurt, 1996: 396, fig. 89; Fiji: N’Yeurt, 2001: 733; Berger et al., 2003: 533, figs 11a-h, 25. Material examined: Moruroa, Jun. 1995, leg. V. Stiger, UPF 497; Iri Bay, Rapa Island, 16 Nov. 2002, leg. J. L. Menou, UPF 3221 RPS 25. 74 A. D. R. N’Yeurt & C. E. Payri Thallus up to 6 mm high, light green, with a monoplanar reproductive disc 2.5-3 mm in diameter, borne atop a slender stalk. Disc composed of 14 cylindrically clavate segments with rounded apices, the segments loosely joined together by light calcification. Corona superior present, corona inferior lacking. Remarks: Growing on coral debris in the lagoon. The complicated taxonomy of this species is discussed in Womersley (1984: 295), Silva et al., (1996: 893-894) and Berger et al. (2003). The genus Parvocaulis was created to accommodate remaining members of the section Polyphysa of the Polyphysaceae, after transfer of the type species from Polyphysa to Acetabularia based on new molecular and morphological information. DISCUSSION This study has brought the total number of Chlorophyceae known from French Polynesia to 81 taxa. Of these, 23 (or about 29 %) represent new records, one (Halimeda heteromorpha) is a new species, and two (Microdictyon sp. and Struveopsis sp.) need more detailed studies to describe as new taxa. Most of the species belong to the Indo-Pacific biogeographic province (Veron, 1995). The species distribution within the French Polynesian islands is very similar to what has been documented for the brown algal flora (N’Yeurt & Payri, 2006). The Society, Tuamotu and Austral groups display the richest floristic diversity (74, 50 and 48 % of the total flora, respectively). For the latter archipelago, Rapa and Marotiri islands (27° S), close to temperate waters, appear unique with 8 species (or 10 % of the total Chlorophyta flora) only recorded in French Polynesia from these islands (Tab. 1), including a number of species such as Cladophora aokii, C. feredayoides and C. ohkuboana known from the northern hemisphere in the western Pacific, and the temperate island of Lord Howe. Other species such as Neomeris annulata, Codium mamillosum, and Siphonocladus tropicus are widely distributed and present in the Hawaiian Islands and other tropical areas. The flora from the Marquesas and Gambiers (15 and 30 % of the total flora, respectively) are less rich due in part to a limited sampling effort, in addition to the lack of typical reef habitats in the Marquesas which could explain the low recorded number of Halimeda and Caulerpa species. Most of the taxa (77, or 96 %) recorded in French Polynesian are species with wide distribution in the tropical Pacific, mostly shared with the Central Pacific islands of Fiji and Micronesia (69 and 64 % respectively), while 60 (or 75%) of species are shared with the Indian Ocean in general. Slightly less than half (45%) of the species are shared with the Hawaiian Islands (Tab. 2). While it is difficult to discuss about endemism in highly dispersive organisms like marine algae, 3 (or about 4 %) of the species seem restricted to French Polynesia. On the other hand, the similarity between the flora calculated using Sørensen’s index (SI = 2x / (2x + y +z) where x is the number of shared species, y the number of the total species of the first island and z is the total species of the second island or group), shows the greatest affinities with the nearest archipelago of the Cook Islands. Similar species richness and biogeographical affinities had previously been discussed with the Phaeophyta (N’Yeurt & Payri, 2006), and reveal the same trends of relatively high diversity compared to other tropical Pacific areas. This may simply reflect a better sampling effort, but it also highlights the high diversity French Polynesian green algae 75 Table 1. Distribution of taxa within the different island groups. Australs Avrainvillea erecta Avrainvillea lacerata Avrainvillea obscura § Avrainvillea ridleyi Boodlea composita Bryopsis pennata var. secunda Bryopsis plumosa Caulerpa bikinensis Caulerpa cupressoides Caulerpa cupressoides var. lycopodium Caulerpa cupressoides var. mamillosa Caulerpa nummularia § Caulerpa pickeringii Caulerpa racemosa var. racemosa Caulerpa racemosa var. peltata § Caulerpa racemosa var. turbinata Caulerpa racemosa var. uvifera § Caulerpa serrulata Caulerpa sertularioides Caulerpa seuratii Caulerpa taxifolia Caulerpa taxifolia ecad. tristichophylla § Caulerpa urvilleana Caulerpa verticillata Caulerpa webbiana Chaetomorpha antennina Chaetomorpha basiretrorsa Chaetomorpha fibrosa Chaetomorpha linum § Chlorodesmis fastigiata Cladophora aokii* § Cladophora catenata § Cladophora feredayoides* § Cladophora herpestica § Cladophora ohkuboana* § Cladophora patentiramea Cladophora sericea Cladophora socialis Cladophoropsis sundanensis Codium arabicum Codium geppiorum Codium mamillosum* § Derbesia marina Dictyosphaeria cavernosa Dictyosphaeria versluysii Halimeda borneensis § Halimeda discoidea Halimeda distorta f. distorta Halimeda distorta f. hederacea Halimeda gracilis Halimeda heteromorpha § Halimeda lacunalis f. lata Halimeda macroloba Halimeda melanesica § 1 Gambier Marquesas 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Society 1 1 Tuamotu 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 A. D. R. N’Yeurt & C. E. Payri 76 Table 1. Distribution of taxa within the different island groups. Halimeda micronesica Halimeda minima Halimeda opuntia Halimeda taenicola Microdictyon sp.* § Microdictyon okamurae Microdictyon umbilicatum Neomeris annulata* § Neomeris van-bosseae Parvocaulis parvula § Phyllodictyon anastomosans Rhipidosiphon javensis Rhizoclonium africanum Rhizoclonium riparium § Siphonocladus tropicus* § Struveopsis sp.* § Ulva clathrata § Ulva compressa Ulva flexuosa Ulva lactuca Ulva rigida Valonia aegagropila Valonia fastigiata Valonia macrophysa Valoniopsis pachynema Ventricaria ventricosa Verdigellas peltata § 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Total of 81 taxa 38 24 12 60 1 1 40 * taxa present only in Rapa and Marotiri islands in Australs (8 taxa) § new records for French Polynesia (23 taxa) Table 2. Number and percentages of French Polynesian Chlorophyta species in common with selected tropical localities. Nr of species shared with FP % Sørensen Index Fiji & Rotuma 55 69 0.34 N’Yeurt et al., 1996; N’Yeurt 2001; Littler & Littler, 2003; South & Skelton, 2003b Guam, Pohnpei, Ant atoll & Micronesia 51 64 0.42 Lobban & Tsuda, 2003; Hodgson & McDermid, 2000 Tropical locality Sources Hawaiian Islands 45 56 0.33 Abbott & Huisman, 2004 Solomon Islands 39 49 0.27 Womersley & Bailey, 1970; Littler & Littler, 2003 Cook Islands 35 44 0.43 Chapman 1977; N’Yeurt & Payri, in prep. Samoa 31 39 0.31 Skelton & South, 1999, 2002; Littler & Littler, 2003 Lord Howe Island 27 34 0.26 Millar & Kraft, 1994; Kraft, 2000 Norfolk Island 19 24 0.24 Millar, 1999 Kermadec 13 16 0.20 Nelson & Adams, 1984 Indian Ocean 60 75 0.15 Silva et al., 1996 French Polynesian green algae 77 of available habitats, ranging from high volcanic islands to coral atolls, and the wide latitudinal distribution of the islands which encompass tropical waters in the Marquesas to temperate-cold waters in the southern part of the Australs. Acknowledgements. AN and CEP duly acknowledge financial support from the French government and the Territorial government of French Polynesia. We thank Dr. Loana Addessi, Professor René Galzin, Dr. Pascale Loret, Professor Alexandre Meinesz, Mr. Joel Orempuller, Mr. André Pham, Miss Ahuura Pouira, Miss Hinano Pouira, Miss Vaimiti Pouira, Miss Vaite Pouira, Mr. John Starmer, Dr. Valérie Stiger and Mr. JeanMarc Zanini for help with collections. Dr. Jennifer Bryant (British Museum of Natural History, BM) is warmly thanked for letting AN examine collections of Polynesian algae under her care, as is Professor Michio Masuda (Hokkaido University, SAP). Professor Bruno De Reviers (Muséum National d’Histoire Naturelle, Paris, PC) is thanked for the loan of type collections of Caulerpa, and for kindly allowing these to be photographed and used in this publication. Dr. Heroen Verbruggen is sincerely thanked for his generous assistance with reviewing the Halimeda section of this study, as is Dr. Frederik Leliaert for members of the Cladophorales. We are indebted to Professor Paul C. Silva (University of California at Berkeley) for his kind assistance with Codium taxonomy, and for his unfailing help in locating and lending to us early Polynesian collections held in UC. Mr. Johnson Seeto (The University of the South Pacific Marine Studies Programme, SUVA-A) is thanked for granting access to herbarium collections. Mr. André Pham (Université de la Polynésie française, UPF) is thanked for granting access to his photographs of certain herbarium sheets. Professor G. Robin South is thanked for his detailed review and most useful comments on an earlier version of this manuscript. REFERENCES ABBOTT I. A., 1986 — A unique form of Valoniopsis pachynema (Chlorophyta) from French Polynesia. Cryptogamie, Algologie 7: 161-167. ABBOTT, I. A., 1991 — Polynesian uses of seaweed In: Cox P. A. & Bannack S. (eds), Islands, Plants, and Polynesians. Portland, Dioscorides Press, pp. 135-145. ABBOTT I. A. & HUISMAN J. M., 2003 — New records and notes on Hawaiian marine benthic Chlorophyta, including Pseudochlorodesmis abbreviata (Gilbert) n. comb., (Udoteaceae) and Cladophora luxurians (Gilbert), n. comb., (Cladophoraceae). Pacific science 57: 275-285. ABBOTT I. A. & HUISMAN J. M., 2004 — Marine Green and Brown Algae of the Hawaiian Islands. Bishop Museum Press, Honolulu, xi + 259 + [1] p, 95 figs. AGARDH C. A., 1811 — Dispositio algarum Sueciae, quam publico examini subjiciunt Carl Adolph Agardh. Part II. Lund, Sweden, pp. 17-26. AGARDH C. A., 1817 — Synopsis algarum Scandinaviae, adjecta dispositione universali algarum. Lund, xl + 135 p. AGARDH C. A., 1823 — Species algarum rite cognitae cum synonymis, differentis specifis et descriptionibus succintis. Volume 1, part 2. Lund, pp. 169-531. AGARDH C. A., 1824 — Systema algarum. Lund, xxxviii + 312 p. AGARDH J. G., 1837 — Novae species algarum, quas in itinere ad oras maris rubri collegit Eduardus Rüppell, cum observationibus nonnullis in species rariores antea cognitas. Museum Senckenbergianum, Abhandlungen aus dem Gebiete der beschreibenden Naturgeschichte 2: 169-174. AGARDH J. G., 1847 — Nya alger frän Mexico. Öfversigt af Kongelige (Svenska) vetenskaps akademiens förhandlingar 4: 5-17. AGARDH J. G., 1873 — Till algernes systematik. I. Caulerpa. II. Zonaria. III. Sargassum. Lunds universitets Årsskrift, Afdelningen för mathematik och naturvetenskap 9: 1-71. AGARDH J. G., 1883 — Till algernes systematik, Nya bidrag (Tredje afdelningen). Lunds universitets Årsskrift, Afdelningen för mathematik och naturvetenskap 19: 1-177. AGARDH J. G., 1886-1887 — Till Algernes Systematik, Nya bidrag Femte afdelningen. Lunds universitets Årsskrift, Afdelningen för mathematik och naturvetenskap 232: 1-174. 78 A. D. R. N’Yeurt & C. E. Payri ANAND P. L.,1940 — Marine algae from Karachi. Part I. Chlorophyceae. University of Punjab, Lahore, 52 p. BALLANTINE D. L. & NORRIS J. N., 1994 — Verdigellas, a new deep-water genus (Tetrasporales, Chlorophyta) from the tropical western Atlantic. Cryptogamic botany 4: 368-372. BALLANTINE D. L. & APONTE N. E., 1996 — Verdigellas nektongammea (Tetrasporales, Chlorophyta), a new deep-water species from the Bahamas. Nova Hedwigia 62: 425-429. BARTON E. S., 1901 — The genus Halimeda. Monographs of the Siboga Expedition 60: 32 pp. BERGER S., FETTWEISS U., GLEISSBERG. S., LIDDLE L. B., RICHTER U., SAWITZKY. H. & ZUCCARELLO G., 2003 — 18S rDNA phylogeny and evolution of cap development in Polyphysaceae (formerly Acetabulariaceae; Dasycladales, Chlorophyta). Phycologia 42: 506-561. BERKELEY M. J., 1842 — Enumeration of fungi, collected by H. Cuming, Esq. F.L.S. in the Philippines Islands. London journal of botany 1: 142-157, pl. VI figs 4, 4; pl. VII figs 6-11. BLAIR S. M. & NORRIS J. N., 1988 — The deep-water species of Halimeda Lamouroux (Halimedaceae, Chlorophyta) from San Salvador Island, Bahamas: species composition, distribution and depth records. Coral reefs 6: 227-236. BLIDING C., 1963 — A critical survey of European taxa in Ulvales. Part I. Capsosiphon, Percursaria, Blidingia, Enteromorpha. Opera botanica 8: 1-160. BLOMSTER J., MAGGS C. A. & STANHOPE M. J., 1998 — Molecular and morphological analysis of Enteromorpha intestinalis and E. compressa (Chlorophyta) in the British Isles. Journal of phycology 34: 319-340. BONNEAU E. R., 1977 — Polymorphic behavior of Ulva lactuca (Chlorophyta) in axenic culture. I. Occurrence of Enteromorpha-like plants in haploid clones. Journal of phycology 13: 133-140. BØRGESEN F., 1905 — Contributions à la connaissance du genre Siphonocladus Schmitz. Oversiigt over det Kongelige Danske videnskabernes selskabs forhandlinger 1905: 259-291, 13 figures. BØRGESEN F., 1907 — An ecological and systematic account of the Caulerpas of the Danish West Indies. Det Kongelige Danske videnskabernes selskab skrifter 7 Række, naturvidenskabelig og mathematisk afdeling 4: 337-392. BØRGESEN F., 1913 — The marine algae of the Danish West Indies, Part 1: Chlorophyceae. Dansk botanisk arkiv 1: 1-160. BØRGESEN F., 1924 — Marine algae from Easter Island. In: Skottsberg C. (ed.), The Natural History of Juan Fernandez and Easter Island. Uppsala, Almqvist and Wiksells, pp. 247-309. BØRGESEN F., 1925 — Marine algae from the Canary Islands. I. Chlorophyceae. Det Kongelige Danske videnskabernes selskab, biologiske meddelelser 5: 1-123. BØRGESEN F., 1932 — A revision of Forsskål’s algae mentioned in Flora Aegyptiaco-Arabica and found in his herbarium in the Botanical Museum of the University of Copenhagen. Dansk botanisk arkiv 8: 1-14. BØRGESEN F., 1934 — Some marine algae from the northern part of the Arabian Sea with remarks on their geographic distribution. Det Kongelige Danske videnskabernes selskab, biologiske meddelelser 11(6): 1-72. BØRGESEN F., 1935 — A list of marine algae from Bombay. Det Kongelige Danske videnskabernes selskab, biologiske meddelelser 12(2): 1-64. BØRGESEN F., 1936 — Some marine algae from Ceylon. Ceylon journal of science (A) 12: 57-96. BØRGESEN F., 1940 — Some marine algae from Mauritius. I. Chlorophyceae. Det Kongelige Danske videnskabernes selskab, biologiske meddelelser 15 (4): 1-81. BØRGESEN F., 1952 — Some marine algae from Mauritius. Additions to the parts previously published, IV. Det Kongelige Danske videnskabernes selskab, biologiske meddelelser 18: 3-72. BORY DE SAINT-VINCENT J. B. G. M., 1804 — Voyage dans les quatre principales îles des mers d’Afrique ... Atlas. Volume 2. Paris, 431 p., pls. I—XIV. BORY DE SAINT-VINCENT J. B. G. M., 1826-1829 — Cryptogamie. In: Duperrey M. L. L. (ed.), Voyage autour du Monde, exécuté par ordre du Roi, sur la Corvette de Sa Majesté, la Coquille, pendant les années 1822-5. Bertrand, Paris, pp. 1-96 (1827); pp. 97-200 (1828); pp. 201-301 (1829), pls 1-39. BRAND F., 1904 — Über die Anheftung der Cladophoraceen und über verschiedene polynesische Formen dieser Familie. Beihefte zum botanischen Centralblatt 18: 165-193, pls. V, VI. BRAVIN I. C., TORRES J., GURGEL C. F. D. & YONESHIGUE Y., 1999 — Novas ocorrências de clorofíceas marinhas de profundidade para o Brasil. Hoehnea 26: 121-133. CHAPMAN V. J., 1977 — Marine algae of Norfolk Island and the Cook Islands. Botanica marina 20: 161-165. COLLINS F. S. & HERVEY A. B., 1917 — The algae of Bermuda. Proceedings of the American academy of arts and sciences 53: 1-195, VI pls. French Polynesian green algae 79 CONTE E. & PAYRI C. E., 2002 — La consommation des algues en Polynésie française: premiers résultats d’une enquête. Journal de la société des océanistes 114-115: 165-172. CONTE E. & PAYRI C. E., 2006 — Present day consumption of edible algae in French Polynesia: a study of the survival of pre-European practices. Journal of the Polynesian society 115: 77-93. COPPEJANS E. & MEINESZ A., 1988 — Marine algae of Papua New Guinea (Madang Prov.) 1. Caulerpaceae (Chlorophyta – Caulerpales). Blumea 33: 181-196. COPPEJANS E. & BEECKMAN T., 1989 — Caulerpa section Sedoideae (Chlorophyta, Caulerpales) from the Kenyan coast. Nova Hedwigia 49: 381-393. COPPEJANS E. & PRUD’HOMME VAN REINE W. F., 1989a — Seaweeds of the Snellius-II Expedition. Chlorophyta: Caulerpales (Except Caulerpa and Halimeda). Blumea 34: 119-142. COPPEJANS E. & PRUD’HOMME VAN REINE W. F., 1989b — Seaweeds of the Snellius-II Expedition. Chlorophyta: Dasycladales. Netherlands journal of sea research 23: 123-129. COPPEJANS E. & BEECKMAN T., 1990 — Caulerpa (Chlorophyta, Caulerpales) from the Kenyan coast. Nova Hedwigia 50: 111-125. COPPEJANS E., 1992 — Marine algae of Papua New Guinea (Madang Prov.) 2. A revised and completed list of Caulerpa (Chlorophyta – Caulerpales). Blumea 36: 383-410. COPPEJANS E. & PRUD’HOMME VAN REINE W. F., 1992 — Seaweeds of the Snellius-II Expedition (E. Indonesia): the genus Caulerpa (Chlorophyta – Caulerpales). Bulletin des séances de l’académie des sciences d’Outre-Mer 37: 667-712. COPPEJANS E., DE CLERCK O. & VAN DEN HEEDE C., 1995 — Annotated and illustrated survey of the marine macroalgae from Motupore Island and vicinity (Port Moresby area, Papua New Guinea). I. Chlorophyta. Biologisch Jaarboek Dodonaea 62: 70-108. COPPEJANS E, LELIAERT F., DARGENT O. & DE CLERCK O., 2001 — Marine green algae (Chlorophyta) from the north coast of Papua New Guinea. Cryptogamie, Algologie 22: 375-443. COPPEJANS E., LELIAERT F., VERBRUGGEN H., DE CLERCK O., SCHILS T., DE VRIESE T. & MARIE D., 2004 — The marine green and brown algae of Rodrigues (Mauritius, Indian Ocean). Journal of natural history 38: 2959-3020. DAWSON E. Y., 1954 — Marine Plants in the Vicinity of the Institut Océanographique de Nha Trang, Viêt Nam. Pacific science 8: 373-469. DAWSON E. Y., 1956 — Some marine algae of the Southern Marshall Islands. Pacific science 10: 25-66. DAWSON E. Y., 1957 — An annotated list of marine algae from Eniwetok Atoll, Marshall Islands. Pacific science 11: 92-132. DAWSON E. Y., 1959 — Some algae from Clipperton Island and the Danger Islands. Pacific naturalist 1: 2-8. DAWSON E. Y., ACLETO C. & FOLDVIK N., 1964 — The seaweeds of Peru. Nova Hedwigia 13: 1-111, pls 1-81. DE CLERCK O., BOLTON J. J., ANDERSON R. J. & COPPEJANS E., 2005 — Guide to the seaweeds of Kwazulu-Natal. Scripta botanica belgica 33: 1-294. DE REVIERS B., 2003 — Biologie et Phylogénie des Algues. Tome 2. Paris, Belin, 255 p. DECAISNE M. J., 1841 — Plantes de l’Arabie heureuse, recueillies par M. P. E. Botta et décrites par M. J. Decaisne. Archives du Muséum d’histoire naturelle (Paris) 2: 89-199, pls V-VII. DECAISNE M. J., 1842 — Mémoires sur les corallines ou polypiers calcifères. Annales des sciences naturelles, Botanique, series 2, 17: 297-380, pls 14-17; 18, 96-128. DICKIE G., 1874 — On the algae of Mauritius. Journal of the Linnean society (Botany) 14: 190-202. DICKIE G., 1877 — Notes on algae from the island of Mangaia, South Pacific. Journal of the Linnean society (Botany) 15: 30-33. DIXIT S. C., 1968 — Species list of Indian marine algae - II. Journal of the university of Bombay 36: 9-24. DRAGASTAN O. N., LITTLER D. S. & LITTLER M. M., 2002 — Recent vs. fossil Halimeda species of Angaur Island, Palau and adjacent western Pacific areas. Acta palaeontologica Romaniae 1: 1-22. DUCKER S. C., 1969 — Additions to the genus Chlorodesmis (Chlorophyta). Phycologia 8: 17-20. EGEROD L. E., 1952 — An analysis of the siphonous Chlorophycophyta with special reference to the Siphonocladales, Siphonales and Dasycladales of Hawai’i. University of California Publications in botany 25: 325-454. EGEROD L. E., 1974 — Report of the marine algae collected on the fifth Thai-Danish Expedition of 1966. Chlorophyceae and Phaeophyceae. Botanica marina 17: 130-157. EGEROD L. E., 1975 — Marine algae of the Andaman coast of Thailand: Chlorophyceae. Botanica Marina 18: 41-66. ENDLICHER S. L., 1843 — Mantissa botanica altera: Sistens generum plantarum supplementum tertium. Vienna, 111 p. 80 A. D. R. N’Yeurt & C. E. Payri EUBANK L. L., 1946 — Hawaiian representatives of the Genus Caulerpa. University of California Publications in botany 18: 409-432. FELDMANN J., 1937 — Les algues marines de la côte des Albères. I-III. Cyanophycées, Chlorophycées, Pheophycées. Revue algologique 9: 141-334. FORSSKÅL P., 1775 — Flora Aegyptiaco-Arabica. Copenhagen, 219 p. FURNARI G., CORMACI M. & SERIO D., 1999 — Catalogue of the benthic marine macroalgae of the Italian coast of the Adriatic Sea. Bocconea 12: 1-214. GEPP A. & GEPP E. S., 1904 — Rhipidosiphon and Callipsygma. Journal of botany 42: 363-366, plate 467. GEPP A. & GEPP E. S., 1908 — Marine algae (Chlorophyceae and Phaeophyceae) and marine phanerogams of the “Sealark” Expedition, collected by J. Stanley Gardiner, M.A., F.R.S., F.L.S. Transactions of the Linnean society of London, 2nd Series, Botany, 7: 163-188, pls 22-24. (reprinted 1909). GEPP A. & GEPP E. S., 1911 — The Codiaceae of the Siboga Expedition, including a monograph of Flabellariae and Udoteae. Siboga-Expeditie Monographie 62: 1-150, 22 pls. GILBERT W. J., 1943 — Studies on Philippines Chlorophyceae. I. The Dasycladaceae. Papers of the Michigan academy of science, arts, and letters 28: 15-35. GILBERT W. J., 1947 — Studies on Philippines Chlorophyceae. III. The Codiaceae. Bulletin of the Torrey botanical club 74: 121-132. GILBERT W. J., 1962 — Contribution to the marine Chlorophyta of Hawaii, I. Pacific science 16: 135-144. GILBERT W. J., 1965 — Contribution to the marine Chlorophyta of Hawaii, II. Additional records. Pacific science 19: 483-492. GMELIN S. G., 1768 — Historia fucorum. Petropolis (Leningrad), (8) + 239 = 6 p., pls IA, IB, IIA, IIB, III = xxxiii. GOREAU T. F. & GRAHAM E. A., 1967 — A new species of Halimeda from Jamaica. Bulletin of marine science 17: 432-441. GRAY J. R., 1866 — On Anadyomene and Microdictyon, with the description of three new allied genera, discovered by Menzies in the Gulf of Mexico. Journal of botany 4: 41-51, 65-72, plate XLIV. GREUTER W., MCNEILL J., BARRIE F. R., BURDET H. M., DEMOULIN V., FILGUEIRAS T. S., NICOLSON D. H., SILVA P. C., SKOG J. E., TREHANE P., TURLAND N. J. & HAWKSWORTH D. L., 2000 — International Code of Botanical Nomenclature (Saint Louis Code) adopted by the Sixteenth International Botanical Congress St. Louis, Missouri, July - August 1999. (Regnum Vegetabile, Volume 138). Königstein, Koeltz Scientific Books, 474 p. GREVILLE R. K., 1830 — Algae britannicae. Edinburgh, lxxxviii + 218 p., 9 pls. GREVILLE R. K., 1853 — Remarks on some algae belonging to the genus Caulerpa. Transactions of the botanical society (Edinburgh) 4: 197-199, pls III, IV. HANYUDA T., WAKANA I., ARAI S., MIYAJI K., WATANO Y.& UEDA K., 2003 — Phylogenetic relationships within Cladophorales (Ulvophyceae, Chlorophyta) inferred from 18s rRNA gene sequences, with special reference to Aegagropila linnaei. Journal of phycology 38: 564-571. HARPER J. T. & GARBARY D. J., 1997 — Marine algae of northern Senegal: the flora and its biogeography. Botanica marina 40: 129-138. HARVEY W. H., 1834 — Notice of a collection of algae, communicated to Dr. Hooker by the late Mrs. Charles Telfair, from “Cap Malheureux”, in the Mauritius; with descriptions of some new and little known species. Journal of botany (Hooker) 1: 147-157, pls. CXXV, CXXVI. HARVEY W. H., 1847-1849 — Nereis australis. London, viii + 124 pp., L pls. [1847: pp. i-viii + 1-64, pls. I-XXV; 1849: pp. 65-124, pls. XXVI-L]. HARVEY W. H. & BAILEY J. W., 1851 — Descriptions of seventeen new species of algae collected by the United States Exploring Expedition. Proceedings of the Boston society of natural history 3: 370-373. HARVEY W. H., 1855 — Some account of the marine botany of the colony of Western Australia. Transactions of the royal Irish academy 22: 525-566. HARVEY W. H., 1858a — Nereis boreali-americana ... Part III. Chlorospermeae. Smithsonian contributions to knowledge 10: 1-140, pls XXXVII-L. HARVEY W. H., 1858b — Phycologia australica. Volume 1. London, [i]-xi + v-viii (Index) p., pls I-LX. HARVEY W. H., 1859 — Phycologia australica. Volume 2. London, viii p., pls LXI-CXX. HAYDEN H. S., BLOMSTER J., MAGGS C. A., SILVA P. C., STANHOPE M. J. & WAALAND J. R., 2003 — Linnaeus was right all along: Ulva and Enteromorpha are not distinct genera. European journal of phycology 38: 277-294. HILLIS L., 1959 — A revision of the genus Halimeda (order Siphonales). Publications of the institute of marine science, University of Texas 6: 321-403. French Polynesian green algae 81 HILLIS-COLINVAUX L., 1968 — New species of Halimeda: a taxonomic reappraisal. Journal of phycology 4: 30-35. HILLIS-COLINVAUX L., 1980 — Ecology and taxonomy of Halimeda primary producers of coral reefs. Advances in marine biology 17: 1-327. HODGSON L. M. & MCDERMID K. J., 2000 — Marine plants of Pohnpei and Ant Atoll: Chlorophyta, Phaeophyta and Magnoliophyta. Micronesica 32: 289-307. HODGSON L. M., TRI P. H., LEWMANOMONT K. & MCDERMID K. J., 2004 — Annotated checklist of species of Caulerpa and Caulerpella (Bryopsidales, Caulerpaceae) from Viêt Nam, Thailand, and the Hawaiian Islands. In: Abbott I. A. & McDermid K. J. (eds), Taxonomy of Economic Seaweeds. With reference to the Pacific and other locations. Honolulu, Hawaii Sea Grant College Program, pp. 21-38. HOLMES E. M., 1896 — New marine algae from Japan. Journal of the Linnean society (London), Botany 31: 248-260, pls. 7-12. HOWE M. A., 1905 — Phycological studies II. New Chlorophyceae, new Rhodophyceae and miscellaneous notes. Bulletin of the Torrey botanical club 35: 563 586, pls 23-29. HOWE M. A., 1909 — Phycological Studies, IV. The Genus Neomeris and notes on other Siphonales. Bulletin of the Torrey botanical club 36: 75-104, pls 1-8. HOWE M. A., 1914 — The Marine Algae of Peru. Memoirs of the Torrey botanical club 15: 1-185, 44 figs, 66 pls. HUDSON W., 1762 — Flora anglica. Editio altera. Londinum (London), xxxviii + 690 pp. ITONO H., 1973 — Notes on marine algae from Heteruma Island, Ryukyu. Botany magazine (Tokyo) 86: 155-168. JAASUND E., 1976 — Seaweeds in Tanzania. 1st edition. Tromsø, University of Tromsø, 160 p. JOLY A. B., DE OLIVEIRA-FIGUEIREDO M. A., UGADIM Y., PINHEIRO F. C., FERREIRA M. M. & CORDEIRO M., 1968 — Additions to the marine flora of Brazil - VIII. Rickia 3: 161-170. JONES R. & KRAFT G. T., 1984 — The Genus Codium (Codiales, Chlorophyta) at Lord Howe Island (N.S.W.). Brunonia 7: 255-276. KANDA T., 1944 — Ecological studies on marine algae from Korôru and adjacent island in the south sea islands. Palao tropical biological station studies 2: 733-800. KERMARREC A., 1974 — Reproduction et Cycle de développement des Bryopsis. Ph. D. Thesis, University of Paris VI, 102 p. + 15 pls. KOEMAN R. P. T. & VAN DEN HOEK C., 1980 — The taxonomy of Ulva (Chlorophyceae) in the Netherlands. British phycological journal 16: 9-53. KOOISTRA W. H. C. F., 1992 — Biogeography of Cladophoropsis membranacea (Chlorophyta) based on comparisons of nuclear rDNA ITS sequences. Journal of phycology 28: 660-668. KOOISTRA W. H. C. F., OLSEN J. L., STAM W.T. & VAN DEN HOEK C., 1993 — Problems relating to species sampling in phylogenetic studies: and example of non-monophyly in Cladophoropsis and Struvea (Siphonocladales, Chlorophyta). Phycologia 32: 419-428. KOOISTRA W. H. C. F., COPPEJANS E. & PAYRI C., 2002 — Molecular systematics, historical ecology, and phylogeography of Halimeda (Bryopsidales). Molecular phylogenetics and evolution 24: 121-138. KOOISTRA W. H. C. F. & VERBRUGGEN H., 2005 — Genetic patterns in the calcified tropical seaweeds Halimeda opuntia, H. distorta, H. hederacea and H. minima (Bryopsidales, Chlorophyta) provide insights in species boundaries and interoceanic dispersal. Journal of phycology 41: 177-187. KRAFT G. T. & WYNNE M. J., 1996 — Delineation of the Genera Struvea Sonder and Phyllodictyon J.E. Gray (Cladophorales, Chlorophyta). Phycological research 44: 129-142. KRAFT G. T., 2000 — Marine and estuarine benthic green algae (Chlorophyta) of Lord Howe Island, south-western Pacific. Australian systematic botany 13: 509-648. KUNTZE O., 1891 — Revisio generum plantarum ... Part 2. Leipzig, pp. 375-1011. KÜTZING F. T., 1843 — Phycologia generalis. Leipzig, xxxii + 458 p., 80 pls. KÜTZING F. T., 1845 — Phycologia germanica. Nordhausen, x + 340 p. KÜTZING F. T., 1846 — Tabulae Phycologicae. Volume I. Nordhausen, 8 p., 10 pls. KÜTZING F. T., 1847 — Diagnosen und Bemerkungen zu neuen oder kritischen Algen. Botanische Zeitung 5: 1-5, 22-25, 33-38, 52-55, 164-167, 177-180, 193-198, 219-223. KÜTZING F. T., 1849 — Species Algarum. Leipzig, vi + 922 p. KÜTZING F. T., 1853 — Tabulae Phycologicae. Volume III. Nordhausen, 28 p., 100 pls. KÜTZING F. T., 1856 — Tabulae Phycologicae. Volume VI. Nordhausen, iv + 35 p., 100 pls. KÜTZING F. T., 1866 — Tabulae Phycologicae. Volume XVI. Nordhausen, iv + 35 p., 100 pls. LAMARCK J. B., 1816 — Histoire naturelle des animaux sans vertèbres ... . Vol. 2. Paris, 568 p. LAMOUROUX J. V. F., 1809a — Mémoire sur trois nouveaux genres de la famille des algues marines. Journal de botanique (Desvaux) 2: 129-135, pl. I; pl. III: fig. 1 LAMOUROUX J. V. F., 1809b — Mémoires sur les Caulerpes, nouveau genre de la famille des algues marines. Journal de botanique (Desvaux) 2: 136-146. 82 A. D. R. N’Yeurt & C. E. Payri LAMOUROUX J. V. F., 1809c — Observation sur la physiologie des algues marines, et description de cinq nouveaux genres de cette famille. Nouveau bulletin des sciences de la société philomatique de Paris 1: 330-333. LAMOUROUX J. V. F., 1812 — Extrait d’une mémoire sur la classification des polypiers coralligènes non entièrement pierreux. Nouveau bulletin des sciences de la société philomatique de Paris 3: 181-188. LAMOUROUX J. V. F., 1816 — Histoire des polypiers coralligènes flexibles, vulgairement nommés zoophytes. Caen, F. Poisson, lxxxiv + 560 p., 1 table, 19 pls. LEE W. J., BOO S. M. & LEE I. K., 1991 — Notes on the genus Bryopsis (Bryopsidaceae, Chlorophyta) from Ullungdo Island, Korea. Korean journal of phycology 6: 23-29. LELIAERT F., COPPEJANS E. & DE CLERCK O., 1998 — The Siphonocladales sensu Egerod (Chlorophyta) from Papua New Guinea and Indonesia (Snellius-II Expedition). Belgian journal of botany 130: 177-197. LELIAERT F., DE CLERCK O., BOLTON J. J. & COPPEJANS E., 2001 — New records of the Chlorophyta from South Africa, with the emphasis on the marine benthic flora of KwaZulu-Natal. South African journal of botany 67: 450-459. LELIAERT F. & COPPEJANS E., 2003 — The marine species of Cladophora (Chlorophyta) from the South African East Coast. Nova Hedwigia 76: 45-82. LELIAERT F., ROUSSEAU F., DE REVIERS B. & COPPEJANS E., 2003 — Phylogeny of the Cladophorophyceae (Chlorophyta) inferreed from partial LSU rRNA gene sequences: is the recognition of a separate order Siphonocladales justified? European journal of phycology 38: 233-246. LELIAERT F., 2004 — Taxonomic and Phylogenetic Studies in the Cladophorophyceae (Chlorphyta). Ph. D. Thesis, Gent University, 294 p. LELIAERT F. & COPPEJANS E., 2006 — A revision of Cladophoropsis Børgesen (Siphonocladales, Chlorophyta) Phycologia 45: 657-679. LINK H. F., 1820 — Epistola de algis aquatics in genera disponendis. In: Nees von Esenbeck C. G. (ed.), Horae Physicae Berolinensis. Bonn, pp. 1-7. LINNAEUS C., 1753 — Species plantarum. Vol. 2. Stockholm, pp. 561-1200. LINNAEUS C., 1753 — Systema naturae per regna tria naturae ... Editio decima ... Vol. 1. Stockholm, IV+823 (824=Errata) p. LIPKIN Y. & SILVA P.C., 2002 — Marine algae and seagrasses of the Dahlak Archipelago, southern Red Sea. Nova Hedwigia 75: 1-90, 7 figs. LITTLER D. S. & LITTLER M. M., 1990 — Reestablishment of the green algal genus Rhipidosiphon Montagne (Udoteaceae, Bryopsidales) with a description of Rhipidosiphon floridensis sp. nov. British phycological journal 25: 33-38. LITTLER D. S. & LITTLER M. M., 1997 — An illustrated marine flora of the Pelican Cays, Belize. Bulletin of the biological society of Washington 9: 1-149. LITTLER D. S. & LITTLER M. M., 2000 — Caribbean Reef Plants. Washington, OffShore Graphics, Inc., 542 p. LITTLER D. S. & LITTLER M. M., 2003 — South Pacific Reef Plants. Washington, OffShore Graphics, Inc., 331 p. LLUCH, J. R., 2002 — Marine benthic algae of Namibia. Scientia marina 66 (Suppl 3): 5-256. LOBBAN C. S. & TSUDA R. T., 2003 — Revised checklist of benthic marine macroalgae and seagrasses of Guam and Micronesia. Micronesica 35-36: 54-99. LYNGBYE H. C., 1819 — Tentamem hydrophytologia danicae. Hafnia (Copenhagen), xxxii + 248 p., 70 pls. MARTENS G. VON, 1868 — Die Tange. In: Die Preussische Expedition nach Ost-Asien. Nach amtlichen Quellen. Botanischer Theil. Berlin, 152 p., VIII pls. MEINESZ A., 1980 — Sur la reproduction de l’Udotea javensis A. et E.S. Gepp (Udotéacées, Caulerpale). Phycologia 19: 82-84. MEINESZ A., JAUBERT J. & DENIZOT M., 1981 — Distribution of the algae belonging to the genus Caulerpa in French Polynesia (atoll of Takapoto and island of Moorea). Proceedings of the Fourth International Coral Reef Symposium, Manila 2: 431-437. MEINESZ A., PIETKIEWICZ D., KOMATSU T., CAYE G., BLACHIER J., LEMEE R. & RENOUX-MEUNIER A., 1994 — Notes taxinomiques préliminaires sur Caulerpa taxifolia et Caulerpa mexicana. In: Boudouresque C-F, Meinesz A. & Gravez V. (eds), First International Workshop on Caulerpa taxifolia. GIS Posidonie, Université de la Mediterranée, Marseille, pp. 105-114. MEÑEZ E. G. & CALUMPONG H. P., 1982 — The genus Caulerpa from Central Visayas, Philippines. Smithsonian Contributions to Marine Science 17: 1-21. MILLAR A. J. K., 1999 — Marine benthic algae of Norfolk Island, South Pacific. Australian Systematic Botany 12: 479-547. French Polynesian green algae 83 MILLAR A. J. K. & KRAFT G. T., 1994 — Catalogue of marine benthic green algae (Chlorophyta) of New South Wales, including Lord Howe Island, south-western Pacific. Australian Systematic Botany 7: 419-453. MITCHELL G. J. P., DE SZÉCHY M. T. M. & MITSUYA L. A., 1979 — Synopsis of the marine benthic Chlorophyta from Rio de Janeiro coast. Leandra 8-9: 91-123. MONTAGNE C., 1837 — Centurie de plantes cellulaires exotiques nouvelles. Annales des Sciences Naturelles, Botanique, séries 2, 8: 345-370. MONTAGNE C., 1939-1842 — Plantae cellulares. In: Barker-Webb P. & Berthelot S. (eds), Histoire naturelle des Iles Canaries, volume 3 (part 2, section 4). Paris, xv + 208 p., 9 pls. (1839 = pp. 1-16; 1840 = 17-160, pls 1-4, 6; 1841 = 161-208, i-xv, pls. 5, 7, 8; 1842 = pl. 9). MONTAGNE C., 1842 — Prodromus generum specierumque phycearum novarum, in itinere ad Polum Antarcticum... collectarum. Parisii (Paris), 16 p. MONTAGNE C., 1845 — Plantes cellulaires. In: Hombron J. B. & Jacquinot H. (eds), Voyage au Pôle Sud et dans l’Océanie sur les Corvettes l’Astrolabe et la Zelée... pendant les années 1837-18381839-1840, sous le commandement de M. J. Dumont-D’Urville: Botanique. Volume 1. Atlas: Botanique: Cryptogamie. Paris, xiv + 349 p. 20 pls. MÜLLER O. F., 1778 — Icones plantarum... Florae danicae. Volume 5, fascicle 13. Havnia (Copenhagen), 8 p., pls 721-780. MURRAY G., 1889 — On Boodlea, a new genus of Siphonocladaceae. Journal of the Linnaean Society of London, Botany 25: 243-245, pl. 49. NÄGELI C., 1847 — Die neueren Algensysteme und Versuch zur Begrundung eines eigenen Systems der Algen und Florideen. Neue Denkschriften der Allg. Schweizerischen Gesellschaft für die Gesammten Naturwissenschaften 9: 1-275, X pls. NELSON W. A. & ADAMS N. M., 1984 — Marine algae of Kermadec Island. A list of species. National Museum of New Zealand Miscellanous Series 10: 1-29. NELSON W. A. & RYAN K. G., 1986 — Palmophyllum umbracola sp. nov. (Chlorophyta) from offshore islands of northern New Zealand. Phycologia 25: 168-177. NIENHUIS P. H., 1975 — Biosystematics and ecology of Rhizoclonium riparium (Roth) Harv. (Chlorophyceae, Cladophorales) in the estuarine area of the rivers Rhine, Meuse and scheldt. Rotterdam, Riijksuniversiteit te Groningen, Bronder Offset B. V., pp. 56-93. NIZAMUDDIN M. & BEGUM M., 1973 — Revision of the marine Cladophorales from Karachi. Botanica Marina 16: 1-18. NORRIS J. N. & OLSEN J. L., 1991 — Deep-water green algae from the Bahamas, including Cladophora vandenhoekii sp. nov. (Cladophorales). Phycologia 30: 315-328. N’YEURT A. D. R., 1996 — A preliminary floristic survey of the benthic marine algae of Rotuma Island. Australian Systematic Botany 9: 361-490. N’YEURT A. D. R., 2001 — Marine algae from the Suva Lagoon and reef, Fiji. Australian Systematic Botany 14: 689-869. N’YEURT A. D. R., SOUTH G. R. & KEATS D. W., 1996 — A revised checklist of the benthic marine algae of Fiji (including the island of Rotuma) Micronesica 29: 49-96. N’YEURT A. D. R. & PAYRI C. E., 2004 — A preliminary annotated checklist of the marine algae and seagrasses of the Wallis Islands (French overseas territory of Wallis and Futuna), South Pacific. Australian Systematic Botany 17: 367-397. N’YEURT A. D. R. & PAYRI C. E., 2006 — Marine algal flora of French Polynesia. I. Phaeophyceae (brown algae). Cryptogamie, Algologie 27: 111-152. OKAMURA K., 1907-1909 — Icones of Japanese Algae. Volume I. Kazamashobo, Tokyo, 258 p., pls I-L. OKAMURA K., 1913-1915 — Icones of Japanese Algae. Volume III. Kazamashobo, Tokyo, 218 p., pls CI-CL. OLSEN J. L. & WEST J. A., 1988 — Ventricaria (Siphonocladales-Cladophorales complex, Chlorophyta), a new genus for Valonia ventricosa. Phycologia 27: 103-108. OLSEN-STOJKOVICH J., 1985 — A systematic study of the genus Avrainvillea Decaisne (Chlorophyta, Udoteaceae). Nova Hedwigia 41: 1-68. OLSEN J. L., VALERO M., MEUSNIER I., BOELE-BOS S. & STAM W. T., 1998 — Mediterranean Caulerpa taxifolia and C. mexicana (Chlorophyta) are not conspecific. Journal of Phycology 34: 850-856. PAPENFUSS G. F., 1950 — On the identity of Spongocladia and Cladophoropsis. Pacific Science 4: 208-213. PAYRI C. E., 1987 — Variabilité Spatiale et Temporelle de la Communauté des Macrophytes des Récifs Coralliens de Moorea (Polynésie Française) : Contribution des Algues au Métabolisme du Carbone de l’Ecosystème Récifal. Doctoral Thesis, Académie de Montpellier, Université des Sciences et Techniques du Languedoc. 285 p. PAYRI C. E., 1988 — Halimeda contribution to organic and inorganic production in a Tahitian reef system. Coral Reefs 6: 251-2. 84 A. D. R. N’Yeurt & C. E. Payri PAYRI C. E. & MEINESZ A., 1985a — Algae. Proceedings of the Fifth International Coral Reef Congress, Tahiti 6: 498-518. PAYRI C. E. & MEINESZ A., 1985b — Taxonomy and distribution of the genus Halimeda (Chlorophyta, Caulerpales) in French Polynesia. Proceedings of the Fifth International Coral Reef Congress, Tahiti 6: 641-648. PAYRI C. E. & N’YEURT A. D. R., 1997 — A revised checklist of Polynesian benthic marine algae. Australian Systematic Botany 10: 867-910. PAYRI C. E., N’YEURT A. D. R. & OREMPULLER J., 2000 — Algues de Polynésie française / Algae of French Polynesia. Editions Au Vent des Iles, Papeete, 380 p., 250 colour photographs. PAYRI C. E., PICHON M., BENZONI F., N’YEURT A. D. R., VERBRUGGEN H. & ANDREFOUËT S. M., 2002 — Atelier Marin Wallis 2002. Contribution à l’Etude de la Biodiversité dans les Récifs Coralliens de Wallis. Scléractiniaires et Macrophytes. Université de la Polynésie française, Outumaoro, 54 p., 6 pls with colour photographs, Landsat map. Available in electronic format from the authors. PICCONE A., 1884 — Crociera del Corsaro alle Isole Madera e Canarie del Capitano Enrico D’Albertis: Alghe. Genova (Genoa), 60 p., 5 figs on plate. PROVASOLI L. & PINTNER J. J., 1980 — Bacteria-induced polymorphism in an axenic laboratory strain of Ulva lactuca (Chlorophyceae). Journal of Phycology 16: 196-201. RABENHORST L., 1868 — Flora europaea algarum aquae dulcis et submarinae. Sectio III. Algae chlorophyllophyceas, melanophyceas et rhodophyceas complectens. Lipsiae (Leipzig), XX + 461 p., (51) figs. REINBOLD T., 1905 — Einige neue Chlorophyceen aus dem Ind. Ocean (Niederl. Indien), gesammelt von A. Weber-van Bosse. Nuova Notarisia 16: 145-149. REYES A. Y., 1976 — The littoral benthic algae of Siquijor Province I. Cyanophyta and Chlorophyta. The Philippine Journal of Science 105: 133-191. RHYNE C. F. & ROBINSON H., 1968 — Struveopsis, a new genus of green algae. Phytologia 17: 467-472. RICHARDSON W. D., 1975 — The marine algae of Trinidad, West Indies. Bulletin of the British Museum (Natural History) 5: 71-143, pls 16-27. RIETEMA H., 1975 — Comparative Investigations on the Life-Histories and Reproduction of Some Species in the Siphoneous Green Algal Genera Bryopsis and Derbesia. Ph. D. Thesis, Groningen University, 132 p. ROTH A. W., 1797 — Catalecta botanica .... . Lipsiae (Leipzig), fascicle 1, (VIII) + 244 (+10) p., VIII pls. ROTH A. W., 1800 — Tentamen florae germanicae. Vol. 3, part 1. Lipsiae (Leipzig). ROTH A. W., 1806 — Catalecta botanica ... . Lipsiae (Leipzig), fascicle 3, (VIII) + 350 (+9) p., XII pls. SAKAI Y., 1964 — The species of Cladophora from Japan and its vicinity. Scientific Papers of the Institute of Algological Research, Faculty of Science, Hokkaido Imperial University 5: 1-104. SARTONI G., 1978 — Ricerche sulla flora algale della Somalia centro-meridionale. 1. Il genere Caulerpa. Webbia 32: 397-416. SARTONI G., 1979 — Ricerche sulla flora algale della Somalia centro-meridionale. 2. I generi Halimeda ed Udotea. Webbia 33: 279-300. SARTONI G., 1986 — Algal flora and his [sic.] vertical distribution on the Gesira cliff (centralsouthern Somalia). Webbia 39: 355-377. SARTONI G., 1992 — Research on the marine algae of south-central Somalia. 3. The Siphonocladales-Cladophorales complex. Webbia 46: 291-326. SCHMIDT O. C., 1923 — Beiträge zur Kennthis der Gattung Codium Stack. Bibliotheca Botanica 23: iv + 68 p., 44 figs. SCHMITZ F. R., 1879 — Ueber grüne Algen im Golf von Athen. Sitzungsber. Naturf. Gesell. Halle 1878: 17-23 SCHNETTER R. & BULA-MEYER G., 1982 — Marine algen der Pazifikküste von Kolombien: Chlorophyceae, Phaeophyceae, Rhodophyceae. Bibliotheca Phycologia 60: xvii + 287 p., 37 pls, 1 figure. SCHRAMM A. & MAZE H., 1865 — Essai de Classification des Algues de la Guadeloupe. Basse Terre, 52 p. SEGAWA S., 1938 — On the marine algae of Susaki, Prov. Izu, and its vicinity III. Scientific Papers of the Institute of Algological Research, Faculty of Science, Hokkaido Imperial University 2: 131-153. SETCHELL W. A., 1924 — American Samoa, Part I: Vegetation of Tutuila Island. Carnegie Institution of Washington, Publications 341: 1-188, pls 18-20, 36-37. SETCHELL W. A., 1925 — Notes on Microdictyon. University of California Pubications in Botany 13: 101-107. French Polynesian green algae 85 SETCHELL W. A., 1926 — Tahitian algae collected by W. A. Setchell, C. B. Setchell, and H. E. Parks. University of California Publications in Botany 12: 61-142. SETCHELL W. A., 1929 —The genus Microdictyon. University of California Publications in Botany 14: 453-588. SETCHELL W. A. & GARDNER N. L., 1937 — The Templeton Crocker Expedition of the California Academy of Sciences 1932, N° 31, A Preliminary Report on the Algae. Proceedings of the California Academy of Science 22: 65-98, pls 3-25. SHIMADA S., HIRAOKA M., NABATA S., IIMA M. & MASUDA M., 2003 — Molecular phylogenetic analyses of the Japanese Ulva and Enteromorpha (Ulvales, Ulvophyceae), with special reference to the free-floating Ulva. Phycological Research 51: 99-108. SILVA P. C., 1952 — Codium Stackhouse. In: Egerod L. E. (ed.), An analysis of the siphonous Chlorophycophyta with special reference to the Siphonocladales, Siphonales and Dasycladales of Hawaii. University of California Publications in Botany 25: 325-454; figs 11-18, pls 34b, 35, 36. SILVA P. C. & WOMERSLEY H. B. S., 1956 — The genus Codium (Chlorophyta) in southern Australia. Australian Journal of Botany 4: 261-289. SILVA P. C., MEÑEZ E. G. & MOE R. L., 1987 — Catalog of the marine benthic algae of the Philippines. Smithsonian Contributions in Marine Science 27: iv + 179 p. SILVA P. C., BASSON P. W. & MOE R. L., 1996 — Catalogue of the Benthic Marine Algae of the Indian Ocean. University of California Publications in Botany 79: 1-1259. SKELTON P. A. & SOUTH G. R., 1999 — A preliminary checklist of the benthic marine algae of the Samoan Archipelago. The University of the South Pacific Marine Studies Technical Report 99/1: 30 p. SKELTON P. A. & SOUTH G. R., 2002a — Annotated catalogue of the benthic marine algae of the Palolo Deep National Marine Reserve of Samoa. Australian Systematic Botany 15: 135-179. SKELTON P. A. & SOUTH G. R., 2002b — Mangrove-associated algae from Samoa, South Pacific. Constancea 83: 1-16. SKELTON P. A. & SOUTH G. R., 2004 — New records and notes on marine benthic algae of American Samoa – Chlorophyta and Phaeophyta. Cryptogamie, Algologie 25: 291-312. SOLIER A. J. J., 1846 — Sur deux algues zoosporées formant le nouveau genre Derbesia. Revue Botanique (Duchartre) 1: 452-454. SOLMS-LAUBACH H., 1895 — Monograph of the Acetabularieae. Transactions of the Linnaean Society of London (Bot. ser 2) 5: 1-39, pls 1-4. SOUTH G. R., 1992 — Contributions to a catalogue of the marine algae of Fiji. 1. Halimeda (Chlorophyceae). University of the South Pacific Marine Studies Programme Technical Report 1992 (2): 1-19. SOUTH G. R., 1993 — Edible seaweeds of Fiji, an ethnobotanical study. Botanica Marina 36: 335-349. SOUTH G. R., 1998 — The seaweed resources of the South Pacific Islands. In: Critchley A. T. & Ohno M. (eds), Seaweed Resources of the World. JICA, Yokusaka, pp. 146-155. SOUTH G. R. & N’YEURT A. D. R., 1993 — Contributions to a catalogue of benthic marine algae of Fiji. II. Caulerpa and Caulerpella (Chlorophyta – Caulerpales). Micronesica 26: 109-138. SOUTH G. R. & SKELTON P. A., 2003a — Revisions and additions to Caulerpa (Chlorophyta, Caulerpaceae) from the Fiji Islands, South Pacific. Australian Systematic Botany 16: 539-548. SOUTH G. R. & SKELTON P. A., 2003b — Catalogue of the marine benthic macroalgae of the Fiji Islands, South Pacific. Australian Systematic Botany 16: 699-758. STACKHOUSE J., 1795-1801 — Nereis britannica. Bathonia (Bath), xl + 112 p., xvii pls. (1795 = Fascicle 1, pp. i-viii + 1-30, pls i-viii; 1797 = Fascicle 2, pp. ix-xxiv +31-70, pls ix-xii; 1801 = Fascicle 3, pp. xxv-xl + 71-112, pls xiii-xvii). STEGENGA H. & VROMAN M., 1988 — Additions to the marine algal flora of Curaçao, Netherlands Antilles. Blumea 33: 299-311. STEPHENSON T. A., 1944 — The constitution of the intertidal fauna and flora of South Africa. Part II. Annals of the Natal Museum 10: 261-358, pls XII-XIV. SVEDELIUS N., 1906 — Ecological and systematic studies of the Ceylon species of Caulerpa. Ceylon Marine Biological Reports 1: 111-112. SVEDELIUS N., 1923 — Zur Kenntnis der Gattung Neomeris. Svensk Botanisk Tidskrift 17: 449-471. TANAKA T., 1967 — Some marine algae from Batan and Camiguin islands, Northern Philippines. I. Memoirs of the Faculty of Fisheries, Kagoshima University 16: 13-27. TANAKA T. & ITONO H., 1969 — On two species of Avrainvillea from Southern Japan. Memoirs of the Faculty of Fisheries, Kagoshima University 18: 1-6. TAYLOR W. R., 1950 — Plants of Bikini and other Northern Marshall Islands. University of Michigan Press, Ann Harbor, xv + 227 p., 79 pls. TAYLOR W. R., 1960 — Marine Algae of the Eastern Tropical and Subtropical Coasts of the Americas. University of Michigan Press, Ann Harbor. 86 A. D. R. N’Yeurt & C. E. Payri TAYLOR W. R., 1973 — Marine algae of the Smithsonian-Bredin Expedition to the Society and Tuamotu Islands. Pacific Science 27: 37-43. TAYLOR W. R., 1975a — A noteworthy variant Caulerpa. Contributions from the University of Michigan Herbarium 11: 77-79. TAYLOR W. R., 1975b — A new species of Halimeda from Malaysia. Contributions from the University of Michigan Herbarium 11: 81-83. TAYLOR W. R., 1977 — Notes on plants of the genus Caulerpa in the Herbarium of Maxwell S. Doty at the University of Hawai’i. Atoll Research Bulletin 208: 1-7. TRONO G. C. JR., 1968 — The marine benthic algae of the Caroline Islands, I. Introduction, Chlorophyta, and Cyanophyta. Micronesica 4: 137-206. TRONO G. C. JR., 1971 — Some new species of marine benthic algae from the Caroline Islands, western-Central Pacific. Micronesica 7: 45-77. TRONO G. C. JR., 1986 — Philippines seaweeds. In: Guide to Philippines Flora and Fauna. Ministry of Natural Resources and University of the Philippines, Manilla, pp. 203-287. TRONO G. C JR. & TOMA T., 1993 — Cultivation of the green alga Caulerpa lentillifera. In: Ohno M. & Critchley A. T. (eds), Seaweed Cultivation and Marine Ranching. Kanagawa International Fisheries Training Center and Japan International Cooperation Agency (JICA), Yokosuka, pp. 17-23. TSUDA R. T. & KAMURA S., 1991 — Floristics and geographic distribution of Halimeda (Chlorophyta) in the Ryukyu Islands. Japanese Journal of Phycology (Sôrui) 39: 57-76. VAN DEN HEEDE C. & COPPEJANS E., 1996 — The genus Codium (Chlorophyta, Codiales) from Kenya, Tanzania (Zanzibar) and the Seychelles. Nova Hedwigia 62: 389-417. VAN DER STRATE H. J., BOELE-BOS S. A., OLSEN J. L., VAN DE ZANDE L. & STAM W. T., 2002 — Phylogeographic studies in the tropical seaweed Cladophoropsis membranacea (Chlorophyta, Ulvophyceae) reveal a cryptic species complex. Journal of Phycology 38: 572-582. VAHL M., 1802 — Endeel Kryptogamiske planter (fuci) fra St. Croix. Skrifter af Naturhistorie Selskabet 5: 29-47. VALET G., 1966a — Les Dictyosphaeria du groupe versluysii (Siphonocladales, Valoniacées). Phycologia 5: 256-260. VALET G., 1966b — Sur une espèce rare et une nouvelle espèce d’Halimeda de Mélanésie. Revue Générale de Botanique 73: 680-685. VALET G., 1968 — Les algues marines de la Nouvelle-Calédonie. I, Chlorophycées. Nova Hedwigia 15: 29-63. VALET G., 1969 — Contributions à l’étude des Dasycladales, 2 et 3. Nova Hedwigia 17: 551-644. VAN DEN HOEK C., 1963 — Revision of the European species of Cladophora. E. J. Brill, Leiden, 248 p. + 55 pls. VAN DEN HOEK C., 1982 — A taxonomic revision of the American species of Cladophora (Chlorophyceae) in the North Atlantic Ocean and their geographic distribution. NorthHolland Publishing Company, Amsterdam. 236 p. VAN DEN HOEK C. & CHIHARA M., 2000 — A taxonomic revision of the marine species of Cladophora (Chlorophyta) along the coasts of Japan and the Russian Far-east. National Science Museum (Tokyo) Monographs 19: 1-242. VAN DEN HOEK C. & WOMERSLEY H. B. S., 1984 — Genus Cladophora. In: Womersley H. B. S. (ed), The marine benthic flora of southern Australia. Part I. Government Printer, South Australia, Adelaide, pp. 185-213. VELLEY T., 1800 — Description of Conferva umbilicata, a new plant, from New South Wales. Transactions of the Linnean Society (London) 5: 169-170, pl. 7 VERBRUGGEN H. & KOOISTRA W. H. C. F., 2004 — Morphological characterization of lineages within the calcified tropical seaweed genus Halimeda (Bryopsidales, Chlorophyta). European Journal of Phycology 39: 1-16. VERBRUGGEN H., DE CLERCK O., SCHILS T., KOOISTRA W. H. C. F. & COPPEJANS E., 2005 — Evolution and phylogeography of Halimeda section Halimeda (Bryopsidales, Chlorophyta). Molecular Phylogenetics and Evolution 37: 789-803. VERBRUGGEN H., DE CLERCK O. & COPPEJANS E., 2005 — Deviant segments hamper a morphometric approach towards Halimeda taxonomy. Cryptogamie, Algologie 26: 259-274. VERBRUGGEN H., DE CLERCK O., N’YEURT A. D. R., SPALDING H. & VROOM P. S., 2006 — Phylogeny and taxonomy of Halimeda incrassata, including descriptions of H. kanaloana and H. heteromorpha spp. nov. (Caulerpales, Chlorophyta). European Journal of Phycology 41: 337-362. VERON J. E. N., 1995 — Corals in Space and time. The biogeography and evolution of the Scleractinia. Sydney, University of New South Wales Press, 321 p. WEBER-VAN BOSSE A., 1898 — Monographie des Caulerpes. Annales du Jardin de Buitenz 15: 243-401. French Polynesian green algae 87 WEBER-VAN BOSSE A., 1905 — Note sur le genre Dictyosphaeria Dec. Nuova Notarisia 16: 142-144. WEBER-VAN BOSSE A., 1910 — Note sur les Caulerpa de l’île Taiti et sur un nouveau Caulerpa de la nouvelle Hollande. Annales de l’institut Océanographique de Monaco 2: 1-8, 2 pls. WEBER-VAN BOSSE A., 1913 — Liste des algues du Siboga. I. Myxophyceae, Chlorophyceae, Phaeophyceae avec le concours de M. Th. Reinbold. Siboga-Expeditie Monographie 59a: 1-186 + 52 figures and 5 pls. WEST J. A. & CALUMPONG H. P., 1990 — New records of marine algae from the Philippines. Micronesica 23: 181-190. WOELKERLING W. J., 1976 — South Florida Benthic Marine Algae, Keys and Comments. Sedimenta V. Comparative Sedimentology Laboratory, University of Miami, 149 p. WOMERSLEY H. B. S., 1984 — The Marine Benthic Flora of Southern Australia. Part I. (Chlorophyta). Handbook of the Flora and Fauna of South Australia Series, Government Printer, South Australia, 329 p. WOMERSLEY H.B.S. & BAILEY A., 1970 — Marine algae of the Solomon Islands. Philosophical Transactions of the Royal Society, London Biological Series B 259: 257-352. WULFEN F. X., 1803 — Cryptogama aquatica. Archiv für die Botanik 3: 1-64, 1 plate. WYNNE M. J., 1993 — Benthic marine algae from the Maldives, Indian Ocean, collected during the R/ V Te Vega Expedition. Contributions to the University of Michigan Herbarium 19: 5-30. WYNNE M. J., 1995 — Benthic marine algae from the Seychelles collected during the R/V Te Vega Indian Ocean Expedition. Contributions to the University of Michigan Herbarium 20: 261-346. WYNNE M. J., 2001 — New records of benthic marine algae from the Sultanate of Oman, northern Arabian Sea. II. Nova Hedwigia 72: 347-374. WYSOR B., 2002 — Biodiversity and Biogeography of Marine Green Algae of the Republic of Panama. Ph.D. Thesis, University of Louisiana at Lafayette, 292 p. YAMADA Y., 1925 — Studien über die meeresalgen von der Insel Formosa. 1. Chlorophyceae. Botanical Magazine, Tokyo 39: 77-95, V figs. YAMADA Y., 1934 — The marine Chlorophyceae from Ryukyu, especially from the vicinity of Nawa. Journal of Faculty of Science, Hokkaido Imperial University, Series V, 3: 33-88. YAMADA Y., 1940 — On the species of Caulerpa from Micronesia. Kagaku Nanyo 3: 95-107. YAMADA Y., 1941 — Species of Halimeda in the South Seas. Kagaku Nanyo 4: 108-121. YAMADA Y., 1944 — New Caulerpas and Halimedas from Micronesia. Scientific Papers of the Institute of Algological Research, Faculty of Science, Hokkaido Imperial University 3: 27-29. YOSHIDA T., 1977 — Nomenclatural notes on some Japanese marine algae (2). Bulletin of the Japanese Society of Phycology 25: 71-74. ZANARDINI G., 1862 — Scelta di Ficee nuove o più rare del mare Adriatico. Decade terza. Memorie del Reale Istituto Veneto di Scienze, Lettere ed Arti 10: 447-484, pls XXVI (XVII)XXXIII(XXIV). 88 A. D. R. N’Yeurt & C. E. Payri Taxonomic Index Avrainvillea erecta. . . . . . . . . . . . . . . Avrainvillea lacerata . . . . . . . . . . . . . Avrainvillea obscura . . . . . . . . . . . . . Avrainvillea ridleyi . . . . . . . . . . . . . . Boodlea composita . . . . . . . . . . . . . . Bryopsis pennata var. secunda. . . . . Bryopsis plumosa . . . . . . . . . . . . . . . Caulerpa bikinensis. . . . . . . . . . . . . . Caulerpa cupressoides . . . . . . . . . . . Caulerpa cupressoides var. lycopodium . . . . . . . . . . . . . . . . . . . . . . . Caulerpa cupressoides var. mamillosa . . . . . . . . . . . . . . . . . . . . . . . . Caulerpa nummularia. . . . . . . . . . . . Caulerpa pickeringii . . . . . . . . . . . . . Caulerpa racemosa var. racemosa . . Caulerpa racemosa var. peltata . . . . Caulerpa racemosa var. turbinata . . Caulerpa racemosa var. uvifera . . . . Caulerpa serrulata . . . . . . . . . . . . . . . Caulerpa sertularioides . . . . . . . . . . . Caulerpa seuratii . . . . . . . . . . . . . . . . Caulerpa taxifolia . . . . . . . . . . . . . . . Caulerpa taxifolia ecad. tristichophylla . . . . . . . . . . . . . . . . . . . . . . Caulerpa urvilleana. . . . . . . . . . . . . . Caulerpa verticillata . . . . . . . . . . . . . Caulerpa webbiana . . . . . . . . . . . . . . Chaetomorpha antennina . . . . . . . . . Chaetomorpha basiretrorsa . . . . . . . Chaetomorpha fibrosa . . . . . . . . . . . Chaetomorpha linum . . . . . . . . . . . . Chlorodesmis fastigiata. . . . . . . . . . . Cladophora aokii . . . . . . . . . . . . . . . Cladophora catenata . . . . . . . . . . . . . Cladophora feredayoides . . . . . . . . . Cladophora herpestica . . . . . . . . . . . Cladophora ohkuboana . . . . . . . . . . Cladophora patentiramea. . . . . . . . . Cladophora sericea . . . . . . . . . . . . . . Cladophora socialis. . . . . . . . . . . . . . Cladophoropsis sundanensis . . . . . . 67 67 68 68 23 31 31 37 37 38 38 39 39 42 43 44 44 45 46 47 47 49 49 50 50 14 14 15 15 69 19 19 20 20 21 21 21 23 25 Codium arabicum . . . . . . . . . . . . . . . Codium geppiorum . . . . . . . . . . . . . . Codium mamillosum. . . . . . . . . . . . . Derbesia marina . . . . . . . . . . . . . . . . Dictyosphaeria cavernosa. . . . . . . . . Dictyosphaeria versluysii . . . . . . . . . Halimeda borneensis. . . . . . . . . . . . . Halimeda discoidea . . . . . . . . . . . . . . Halimeda distorta f. distorta . . . . . . . Halimeda distorta f. hederacea. . . . . Halimeda gracilis. . . . . . . . . . . . . . . . Halimeda heteromorpha. . . . . . . . . . Halimeda lacunalis . . . . . . . . . . . . . . Halimeda macroloba. . . . . . . . . . . . . Halimeda melanesica . . . . . . . . . . . . Halimeda micronesica. . . . . . . . . . . . Halimeda minima . . . . . . . . . . . . . . . Halimeda opuntia . . . . . . . . . . . . . . . Halimeda taenicola . . . . . . . . . . . . . . Microdictyon okamurae . . . . . . . . . . Microdictyon umbilicatum . . . . . . . . Microdictyion sp. . . . . . . . . . . . . . . . . Neomeris annulata. . . . . . . . . . . . . . . Neomeris van-bosseae. . . . . . . . . . . . Parvocaulis parvula. . . . . . . . . . . . . . Phyllodictyon anastomosans . . . . . . Rhipidosiphon javensis . . . . . . . . . . . Rhizoclonium africanum . . . . . . . . . Rhizoclonium riparium . . . . . . . . . . Siphonocladus tropicus. . . . . . . . . . . Struveopsis sp. . . . . . . . . . . . . . . . . . . Ulva clathrata. . . . . . . . . . . . . . . . . . . Ulva compressa . . . . . . . . . . . . . . . . . Ulva flexuosa . . . . . . . . . . . . . . . . . . . Ulva lactuca . . . . . . . . . . . . . . . . . . . . Ulva rigida . . . . . . . . . . . . . . . . . . . . . Valonia aegagropila . . . . . . . . . . . . . Valonia fastigiata . . . . . . . . . . . . . . . . Valonia macrophysa . . . . . . . . . . . . . Valoniopsis pachynema . . . . . . . . . . Ventricaria ventricosa . . . . . . . . . . . . Verdigellas peltata . . . . . . . . . . . . . . . 34 34 35 32 25 26 53 54 54 56 57 58 59 59 61 62 62 64 65 11 11 13 71 73 73 76 71 16 18 28 28 7 8 8 10 10 29 29 30 30 30 6

RELATED PAPERS

RELATED TOPICS

Log In

Marine algal flora of french polynesia. II. Chlorophyceae (green algae)

Marine algal flora of french polynesia. II. Chlorophyceae (green algae)

Related Papers

RELATED PAPERS

RELATED TOPICS