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.
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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