CSIRO PUBLISHING
www.publish.csiro.au/journals/asb
Australian Systematic Botany 17, 367–397
A preliminary annotated checklist of the marine algae and seagrasses of
the Wallis Islands (French Overseas Territory of Wallis and Futuna),
South Pacific
Antoine D. R. N’YeurtA,C and Claude E. PayriB
A
Laboratoire Terre-Ocean, Université de la Polynesié Française, BP 6570 Faa’a 98702, Tahiti, French Polynesia.
UMR 7138, Systématique, Adaptation, Evolution, Equipe Biodiversité Marine Tropicale, IRD-Nouméa – BPA5,
98848 Nouméa cedex, New Caledonia.
C
Corresponding author; email: nyeurt@upf.pf
B
Abstract. A total of 194 species of marine algae (14 Cyanobacteria, 41 Chlorophyta, 11 Heterokontophyta and 128
Rhodophyta), as well as three species of seagrasses, represent the first published records for the isolated island of
Wallis, South Pacific. The flora has its strongest affinities with Fiji and Rotuma, followed by Samoa and French
Polynesia. The lack of diverse habitats and its geographical location are invoked to explain the relatively low species
richness compared with localities such as Fiji and Samoa. The flora has a typically tropical component dominated
by encrusting coralline red algae, the calcified green algal genera Halimeda, and assemblages of Cyanobacteria.
Normally ubiquitous species such as Halimeda discoidea, and the brown algal genera Hydroclathrus, Colpomenia,
Rosenvingea, Asteronema, and Chnoospora are notably absent from the island, perhaps due to seasonality and the
lack of suitable habitats. The minute epiphytic red alga Acrochaetium kurogii is reported for the first time outside
of its type locality in Japan, while two as yet unidentified species of red algae (Gracilaria sp. and Sebdenia sp.)
could represent new taxa if further useful material is found.
SB0327
MAa. rDi.nRe.pNal’YnsteufrotamndWaCl. iE.sPayri
Introduction
The Wallis Islands (Fig. 1) include Wallis Island, also known
as Uvea Island, and 19 small islets that are located in the
south-western Pacific between 13°18′S and 176°124′W,
two-thirds of the way between Hawaii and New Zealand
(Fig. 2). The Wallis Islands form an archipelago that also
includes the islands of Futuna and Alofi. Wallis Island is a
volcanic high island, of generally elongate shape, which
represents the emerged portions of ancient craters. It is 15
km wide in a north–south direction, and 7.8 km long in an
east–west direction. Emerged land area comprises 79.12
km2, with 129 km of coastline for a total reef-lagoon area of
219.5 km2. The highest point is 155 m tall Mt Lulu; the
island’s geology is composed of volcanic tufs and basalts.
The island was formed in several stages, the last being in the
middle Pleistocene. The human population is approximately
10000, distributed in villages mainly on the eastern coast of
the island.
There is no previous information in the literature
concerning the marine algae or seagrasses of Wallis Island,
although there exists a report on the lagoons and reefs of
Wallis and Futuna (Salvat 1982) with an excellent treatment
of the corals, but which unfortunately only hints at the
© CSIRO
26 August 2004
existence of two genera of marine algae. A few years later, a
study was made of the terrestrial flora of Wallis, including
mangroves (Morat and Veillon 1985). At the request of the
Environmental Service of the French Overseas Territory of
Wallis and Futuna, a survey was undertaken from 24 May to
8 June 2002, so as to establish a preliminary inventory of the
coral fauna and benthic macroalgae/seagrass flora, and
determine the ecological characteristics of the coral reef and
lagoon habitats of Wallis (Payri et al. 2002). This paper deals
with the marine benthic algae and seagrasses encountered in
that study. Scattered algal floras and checklists exist for
neighbouring islands such as Rotuma and Fiji (N’Yeurt
1996a, 1996b; N’Yeurt 2002; South and Skelton 2003,
2004), Samoa (Skelton and South 1999, 2002a, 2000b), the
Solomon Islands (Womersley and Bailey 1970), Nauru
(South and Yen 1992), Phoenix Group, Kiribati (South et al.
2001), the Cook Islands (Chapman 1977; N’Yeurt in prep.)
and French Polynesia (Payri and N’Yeurt 1997; Payri et al.
2000). Recent monographs on the Lord Howe Island flora
(e.g. Allender and Kraft 1983, Kraft 2000) and Norfolk
Island (Millar 1999) have also proved very useful in
identifying species occurring in more northerly islands such
as Fiji, Rotuma and Wallis. Other areas of interest such as
10.1071/SB03027
1030-1887/04/040367
368
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Fig. 1.
Map of the Wallis Islands, showing the 42 collecting stations. See Table 1 for co-ordinates.
Tonga, Tokelau, Vanuatu, and much of Kiribati still remain
largely unknown phycologically. The biogeographical
implications of algal distribution in the region has been
discussed by Millar (1990) and N’Yeurt and South (1997).
Recently, Littler and Littler (2003) published a useful
illustrated guide to South Pacific algae, adding many new
records mainly to the Fijian flora, but also some for
American Samoa, the Solomon Islands, the Cook Islands,
French Polynesia and Papua New Guinea. Unfortunately,
exact localities of collections are not mentioned in that
publication.
Materials and methods
All material was collected using SCUBA, snorkeling or reef-walking.
The position of each of the 42 collecting stations (Table 1) was
determined with a Garmine 12 portable GPS receiver. Station numbers
in the text correspond to the localities mentioned in Table 1. The 42
sites studied have been clustered into five large groups according to
geomorphological criteria (fringing reef, barrier reef, lagoon, pinnacle
and outer reef slopes; see Table 2 in the Results section). At the base
station on land, specimens were sorted out and systematically
photographed with a digital camera (Nikon Coolpix E-995, Nikon
Corporation, Toyko, Japan) before processing. Microphotographs were
taken with an Olympus C-4000Z digital camera (Olympus Optical Co.
Marine plants from Wallis
Fig. 2.
Australian Systematic Botany
369
Map showing the position of the Wallis Islands (arrow) in relation to neighbouring island groups.
(Europa) GMBH, Hamburg, Germany) mounted on an Olympus BH-2
microscope (Olympus Optical Co. Ltd., Tokyo, Japan). Voucher
herbarium specimens were pressed by standard techniques, and
representative parts of thalli and turf algae were stored in 4% buffered
formalin in sealed plastic bags packed in a light-proof container, for
shipment and later anatomical examination in the laboratory.
Specimens are housed at the Phycological Herbarium of the Université
de Polynésie française in Tahiti (UPF), with ‘WSS’ referring to slide
collections.
Results
Geomorphology
There is an almost continuous reef system around the main
island of Uvea, composed of a barrier reef 63 km long
encircling a lagoon 24 km wide in a north–south direction
and 15 km long in an east–west direction, and fringing reefs
extending up to the barrier reefs in the northern and western
parts of the island. The lagoon communicates with the open
sea through four passes, one in the South and three on the
Western part of the barrier reef. Average depth of the lagoon
varies between 10 and 20 m, with maximum depths of 47 m
and 52 m in the Eastern and Southern parts, respectively. A
total of 19 islets surround the main island, some of volcanic
or coralline origin are present on the Eastern barrier reef, and
volcanic islets with fringing reefs are present in the Eastern
and Southern parts of the lagoon. Maximum tidal
fluctuations are of 2 m, and water exchange in the lagoon
occurs through the passes and over the barrier reefs.
Systematic listing of species
The taxonomy adopted generally follows that of Silva et al.
(1987, 1996) supplemented by de Reviers (2003), and
updated by other sources where necessary. To conserve
space, type locality and basionym information are not given
as these can be readily found in the sources cited above and
in the text. For similar reasons, figures for species are
omitted, but many of these can be found in Payri et al.
(2002), upon request to the authors.
Cyanobacteria
This section mostly follows the classification of Velasquez
(1962) and Silva and Pienaar (2000).
Chroococcales
Dermocarpellaceae
Sphaenosiphon olivaceus Reinsch, 1875: 17. (=
Dermocarpa olivacea (Reinsch) Tilden; Desikachary 1959:
174, pl. 33 figs 13, 14; Nagarkar 1998: 533). Vouchers: WSS
06 St. 5, WSS 53 St. 29, WSS 88 St. 1, 127 St. 8
370
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Table 1.
List of collecting stations around Wallis, as shown in Fig. 1
Station no.
Latitude
Longitude
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
13°17'30.3"S
13°17'8.04"S
13°18'20.2"S
13°22'11"S
13°22'6"S
13°22'0.84"S
13°21'30"S
13°15'52.4"S
13°16'26.3"S
13°16'28.2"S
13°17'28.3"S
13°17'46.9"S
13°16'15.8"S
13°17'4.38"S
13°17'11.6"S
13°15'3.3"S
13°15'20.6"S
13°15'22.2"S
13°16'15"S
13°16'2.04"S
13°16'25.1"S
13°20'0.9"S
13°19'0.3"S
13°19'34.6"S
13°21'22.4"S
13°21'28"S
13°21'9.6"S
13°21'5.55"S
13°22'41.4"S
13°21'39.8"S
13°11'17.8"S
13°13'1.38"S
13°11'6.84"S
13°16'5.4"S
13°10'5.1"S
13°14'17.4"S
13°11'14.2"S
13°13'0.06"S
13°14'25.6"S
13°23'10.9"S
13°22'50"S
13°13'07.4"S
176°15'35.9"W
176°16'0.84"W
176°15'34"W
176°13'26.1"W
176°13'15.5"W
176°13'12.5"W
176°12'40"W
176°9'40.36"W
176°07'29"W
176°07'30"W
176°08'30.3"W
176°08'17.8"W
176°08'18.9"W
176°10'5.52"W
176°07'29"W
176°15'5.52"W
176°15'22.3"W
176°15'17.6"W
176°15'14.3"W
176°14'28.7"W
176°14'33.4"W
176°16'5.28"W
176°16'14.8"W
176°16'16.4"W
176°13'27.8"W
176°12'51.68"W
176°13'10.9"W
176°13'07.71"W
176°14'34.1"W
176°16'6.96"W
176°12'7.62"W
176°11'5.64"W
176°12'0.54"W
176°09'24.4"W
176°11'5.1"W
176°15'3.36"W
176°13'6.72"W
176°14'24"W
176°15'6.78"W
176°12'51.8"W
176°12'43.4"W
176°11'14.0"W
Comments: this characteristically dumbell-shaped alga is
commonly found as epiphytic colonies on macroalgae such
as Laurencia, Hypnea.
Nostocales
Habitat
Lagoon
Outer slope
Lagoon
Fringing reef islet
Fringing reef islet
Fringing reef islet
Fringing reef coastline
Fringing reef coastline
Inner barrier reef
Inner barrier reef
Lagoon
Lagoon
Lagoon
Fringing reef coastline
Inner barrier reef
Outer slope
Inner barrier reef
Inner barrier reef
Inner barrier reef
Pinnacle
Pinnacle
Outer slope
Inner barrier reef
Inner barrier reef
Pinnacle
Fringing reef coastline
Fringing reef coastline
Fringing reef coastline
Outer slope
Outer slope
Inner barrier reef
Fringing reef coastline
Inner barrier reef
Fringing reef coastline
Outer slope
Outer slope
Inner barrier reef
Inner barrier reef
Inner barrier reef
Pass
Lagoon
Fringing reef coastline
Scytonemataceae
Scytonema polycystum Bornet et Flahault, 1887: 90. Silva
and Pienaar 2000: 26, figs 48, 88–89. Vouchers: WSS
20 St. 26, 60 St. 31, 156 St. 7, 183 St. 40
Nostocaceae
Oscillatoriales
Nostoc calcicola Brebisson in Meneghini, 1842: 121.
Velasquez 1962: 341, pl. 8 fig. 100. Vouchers: WSS 58
St. 31
Oscillatoriaceae
Rivulariaceae
Calothrix confervicola (Dillwyn) C. Agardh, 1824: 70.
Velasquez 1962: 353, pl. 10 fig. 125. Vouchers: WSS 123
St. 22
Lyngbya majuscula (Dillwyn) Harvey, 1833: 370. Dawson
1954: 380, fig. 3d; Velasquez 1962: 319, pl. 4 fig. 68; Littler
and Littler 1997: 122, fig. 182; N’Yeurt 2001: 692; Albert
et al. 2002. Vouchers: WSS 88, 230 St. 1, 33 St. 2; UPF
1648, 1679 St. 3; WSS 158 St. 7, 125, 128 St. 8; UPF 1615
St. 16, 1407 St. 20; WSS 224 St 20; UPF 1511 St. 29, 1450
St. 30; WSS 57, 58 St. 31, 235 St. 35; UPF 1740 St. 36,
Marine plants from Wallis
Australian Systematic Botany
Table 2. Distribution of the species within the five major
geomorphological zones
OS: outer slope (stations 2, 16, 22, 29, 30, 35, 36), BR: barrier reef (9,
10, 15, 17, 18, 19, 23, 24, 31, 33, 37, 38, 39), L: Lagoon (1, 3, 11, 12,
13, 41), P: Pinnacles (20, 21, 25, 40), FR: fringing reef (4, 5, 6, 7, 8, 14,
26, 27, 28, 32, 34, 42)
Rhodophyta
Chlorophyta
Heterokontophyta
Cyanobacteria
Magnoliophyta
Total
OS
BR
L
P
FR
67
13
05
05
00
90
068
031
004
012
000
115
067
021
009
008
001
106
13
07
01
03
00
24
080
030
010
007
003
130
1517, 1518 St. 41; WSS 226 St. 14.1; UPF 1603 St. 4; WSS
06 St. 5; UPF 1532 St. 10, 1498 St. 11
Comments: this species is found at most sites, its colour
ranging from greenish to brownish-black.
Lyngbya polychroa (Meneghini) Rabenhorst, 1847: 83.
Velasquez 1962: 316, pl. 3 fig. 64 (‘Lynbya sordida’); Littler
and Littler 1997: 183. Vouchers: UPF 1686 St. 3; WSS 206
St. 31
Comments: this species (often reported under the synonym
L. sordida Gomont, see Silva et al. 1996) is distinguished
from Lyngbya majuscula by the smaller average diameter of
its filaments, which are between 16 and 30 µm as opposed to
20–60 µm in L. majuscula (Velasquez 1962). The sheath is
also thinner, and cell contents appears less dense, in L.
polychroa.
Oscillatoria bonnemaisonii (P. Crouan et H. Crouan)
P. Crouan et H. Crouan, 1860: 371. Velasquez 1962: 288, pl.
1 fig. 16. Voucher: WSS 206 St. 31; UPF 1522 St. 41
Oscillatoria margaritifera (Kützing) Gomont, 1892b: 216.
Dawson 1954: 380, fig. 3f; Velasquez 1962: 293, pl. 2
fig. 23. Voucher: WSS 206 St. 31
Phormidiaceae
Phormidium nigroviride (Thwaites)
Komárek, 1988: 405.
Anagnostidis et
Comments: = Oscillatoria nigroviride Thwaites; Dawson
1954: 380, fig. 3g; Velasquez 1962): 293, pl. 2 fig. 24 .
Vouchers: WSS 59 St. 31; UPF 1519 St. 41
Phormidium papyraceum Gomont, 1890: 355. Velasquez
1962: 310, pl. 3 fig. 57. Vouchers: UPF 1553, WSS 85 St. 1;
UPF 1707 St. 20; WSS 227 St. 14.1
Comments: forms thin sheets on sandy substrata in the
lagoon. Filaments interwoven, 3–4 µm in diameter, with
prominent transverse walls. When dry, specimens become
translucent with a papery texture.
371
Spirulina subsalsa Oersted, 1842: 566, pl. 7 fig. 4.
Velasquez 1962: 284, pl. 1 fig. 13; Nagarkar 1998: 534,
fig. 16. Vouchers: WSS 57, 206 St. 31
Comments: this characteristically coil-shaped minute
Cyanobacteria is often found mixed with other algae in turf
populations.
Symploca atlantica Gomont, 1892a: 109, pl. 2 fig. 5.
Velasquez 1962: 312; Littler and Littler 1997: 123, fig. 184.
Vouchers: UPF 1617 St. 16; WSS 213 St. 23, 216 St. 36, 138
St. 4
Comments: forming erect fascicles; filaments 4–5 µm in
diameter.
Symploca hydnoides (Harvey) Kützing, 1849: 272. Dawson
1954: 380, fig. 3o, p; Velasquez 1962: 312, pl. 3 fig. 59;
Islam and Aziz 1982: 650, pl. 2 figs 13–16, pl. 7 figs 34–35;
Payri et al. 2000: 293; Payri et al. 2002: 53, pl. 6 fig. 1; Abed
et al. 2003: 867, fig. 7a–d. Vouchers: WSS 88 St. 1, 124
St. 8; UPF 1568 St. 16, 1437 St. 25; WSS 62 St. 31; UPF
1527 St. 41, 1287 St. 4, 1479 St. 5, 1541 St. 12
Comments: this species forms characteristic erect fascicles
with a clay-like texture; filaments 6–7 µm in diameter.
Symploca laeteviridis Gomont, 1892b: 109, pl. 2 figs 6–8.
Velasquez 1962: 311, pl. 3 fig. 58; Nagarkar 1998: 539,
fig. 25. Voucher: WSS 30 St. 1; UPF 1490 St. 5
Comments: this species is distinguished from S. hydnoides
by the smaller diameter of the filaments (3–4 µm, never
reaching 5 µm) and the absence of adhesion of filaments into
erect fascicles.
Chlorophyta
Ulvales
Ulvaceae
Ulva clathrata (Roth) C. Agardh, 1811: 23. (= Enteromorpha
clathrata (Roth) Greville; Dawson 1954: 384, fig. 6d–e;
Egerod 1974: 134, fig. 4; Kraft 2000: 525, fig. 8; N’Yeurt
2001: 693, figs 2, 3a–b; Skelton and South 2002a: 160,
fig. 22A–B; 2002b: 8, pl. 5 figs 29–33, pl. 6 fig. 34; Abbott
and Huisman 2004: 46, fig. 5A–C). Vouchers: WSS 60 St. 31
Comments: based on culture and molecular studies, Shimada
et al. (2003) suggested that Japanese records of
Enteromorpha should be transferred to Ulva. Most species
previously listed under the genus Enteromorpha Link (1820)
have been transferred to Ulva by Hayden et al. (2003) based
on their own molecular studies. Ulva clathrata is readily
distinguished by multiseriate laterals ending in bulbous
expansions.
Ulva flexuosa Wulfen subsp. paradoxa (Dillwyn) Bliding,
1963: 79. (= Enteromorpha flexuosa (Wulfen) J. Agardh
subsp. paradoxa; Bliding 1963: 79, figs 42a–g, 45a–f;
372
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Egerod 1974: 132, figs 1–3; N’Yeurt 1996: 365, fig. 16;
Kraft 2000: 525, fig. 7F; N’Yeurt 2001: 697, fig. 46).
Vouchers: WSS 228, 229 St. 3
Comments: this subspecies is characterised by its longer
uniseriate laterals (up to 2 mm long) and regularly
transversely arranged cell rows.
Cladophorales
Cladophoraceae
Chaetomorpha crassa (C. Agardh) Kützing, 1845: 204.
Sartoni 1992: 299, fig. 4E; Littler and Littler 1997: 93,
fig. 125; N’Yeurt 2001: 701, fig. 15. Vouchers: UPF 1336,
1337, 1338 St. 8
Chaetomorpha linum (O.F. Müller) Kützing, 1845: 204.
Egerod 1974: 135, fig. 9; Littler and Littler 1997: 93,
fig. 126; 2003: 198. Vouchers: UPF 1583; WSS 112 St. 8
188, figs 30–33; Kraft 2000: 578, fig. 27A–B; Payri et al.
2000: 76; Littler and Littler 2003: 202; Abbott and Huisman
2004: 89, fig. 29A. Vouchers: UPF 1554, 1657, WSS 166
St. 1, 1286 St. 4, 1330 St. 9, 1305 St. 12, 1297 St. 13
Dictyosphaeria versluysii Weber-van Bosse, 1905: 144.
Egerod 1952: 351, figs 1a, 2h–k; Sartoni 1992: 319,
figs 13B, 14A; Leliaert et al. 1998: 190, figs 37–39;
Payri et al. 2000: 76; Payri et al. 2002: 44, pl. 1
fig. 11; Skelton and South 2002a: 162, fig. 23C, D;
Littler and Littler 2003: 204; Abbott and Huisman
2004: 89, fig. 29B. Vouchers: UPF 1289, 1595 St. 4,
1218 St. 7, 1339 St. 8, 1319 St. 10, 1335 St. 9
Phyllodictyon anastomosans (Harvey) Kraft et Wynne,
1996: 139, figs 16–25. Leliaert et al. 1998: 186, figs 23–24;
Payri et al. 2000: 74; Wynne 2001: 368, fig. 34; Littler and
Littler 2003: 200, Abbott and Huisman 2004: 63, fig. 16A, B.
Vouchers: UPF 1220, 1551, 1555 St. 1, 1476 St. 5; WSS 62
St. 31
Cladophora vagabunda (Linnaeus) van den Hoek, 1963:
144. Sartoni 1992: 304, fig. 6C–E; Littler and Littler 1997:
95, fig. 130; Kraft 2000: 562, fig. 21A–G; Leliaert and
Coppejans 2003: 70, figs 14A–B, 15; Abbott and Huisman
2004: 79, fig. 24A–D. Vouchers: UPF 1221 St. 1, 1373 St. 8,
1562 St. 9
Siphonocladus tropicus (P. Crouan et H. Crouan) J. Agardh,
1887: 105. Sartoni 1992: 315, fig. 12A; Abbott and Huisman
2004: 90, fig. 30. Voucher: UPF 1633, St. 7
Siphonocladaceae
Valoniaceae
This family was previously listed under the Siphonocladales,
but recent morphological and molecular evidence suggests
that the latter order and the Cladophorales are monophyletic
(Leliaert et al. 1998: 178, 2003).
Valonia aegagropila C. Agardh, 1823: 429. N’Yeurt 1996:
372, fig. 15a, b; Leliaert et al. 1998: 192, figs 40, 41; Payri
et al. 2000: 78; Littler and Littler 2003: 206; Abbott and
Huisman 2004: 92, fig. 31A. Vouchers: UPF 1230 St. 7,
1351 St. 8
Boergesenia forbesii (Harvey) J. Feldmann, 1938: 1503,
figs 3–5. Sartoni 1992: 306, fig. 7B; N’Yeurt 1996: 369,
fig. 14; Leliaert et al. 1998: 184, fig. 13; Payri et al. 2002:
44, pl. 1 fig. 10; Littler and Littler 2003: 202. Voucher: UPF
1354 St. 8
Boodlea composita (Harvey) Brand, 1904: 187–190. Sartoni
1992: 306, fig. 7C; N’Yeurt 1996: 368, fig. 19 (as Boodlea
coacta (Dickie) Murray); Leliart et al. 1998: 184, figs 16–19
(non figs 14–15, 20, F. Leliaert pers. com.); Kraft 2000: 569,
fig. 24A–C; Payri et al. 2000: 70 (as Boodlea kaeneana
Brand); Littler and Littler 2003: 200; Abbott and Huisman
2004: 85, fig. 26A–B. Vouchers: UPF 1219, 1546 St. 1, 1202
St. 2, 1189 St. 3, 1231 St. 7, 1308 St. 11, 1426, 1731 St. 23
Comments: this species forms spongy masses amidst spaces
between dead coral. This genus (along with Phyllodictyon
and several others) form a close cluster with Cladophoropsis
in the family Cladophorophyceae based on recent
phylogenetic studies (Kooistra et al. 1993; Wysor 2002;
Leliaert et al. 2003) and a single, highly variable species may
need to be recognised (F. Leliaert, pers. comm.).
Dictyosphaeria cavernosa (Forsskål) Børgesen, 1932: 2.
Sartoni 1992: 319, fig. 13A; N’Yeurt 1996: 371, fig. 12;
Littler and Littler 1997: 87, fig. 117; Leliaert et al. 1998:
Comments: see remarks above for Boodlea composita.
Valonia fastigiata Harvey ex. J. Agardh, 1887: 101, pl. I
fig. 5. Sartoni 1992: 321, fig. 14C; Coppejans et al. 1995:
98, fig. 39; Leliaert et al. 1998: 192, figs 42–44; Payri et al.
2000: 78; Payri et al. 2002: 44, pl. 1 fig. 7; Skelton and South
2002a: 162, fig. 24A; Littler and Littler 2003: 206.
Vouchers: UPF 1217 St. 1, 1187, 1684 St. 3, 1281, 1596
St. 4, 1249, 1250 St. 7, 1340 St. 8, 1332 St. 9, 1309 St. 12
Comments: this species is quite common in Wallis,
especially on the inner reef slopes where it forms abundant
tightly adhering masses nested in the infractuosities of dead
coral. It is also found in the seagrass beds, where it assumes
an unusual, ball-like unattached form up to 10 cm in
diameter.
Valonia macrophysa Kützing, 1843: 307. Littler and Littler
1997: 91, fig. 121; Payri et al. 2000: 80; Littler and Littler
2003: 206. Vouchers: UPF 1631, 1632 St. 7, 1703 St. 20
Ventricaria ventricosa (J. Agardh) Olsen et West, 1988: 104.
Sartoni 1992: 323, fig. 14E; N’Yeurt 1996: 372, fig. 23;
Littler and Littler 1997: 89, fig. 119; Payri et al. 2000: 80;
Payri et al. 2002: 44, pl. 1 fig. 12; Littler and Littler 2003:
204; Abbott and Huisman 2004: 94, fig. 32B. Vouchers: UPF
1226 St. 3, 1471 St. 5, 1329 St. 9, 1306 St. 12, 1377 St. 16
Marine plants from Wallis
Bryopsidales
Bryopsidaceae
Bryopsis pennata Lamouroux var. secunda (Harvey) Collins
et Hervey, 1917: 62. Mitchell et al. 1979: 109, pl. 1 figs 6–8;
Littler and Littler 1997: 97, fig. 134; Payri et al. 2000: 82;
Skelton and South 2002a: 163, fig. 24E; Littler and Littler
2003: 208; Abbott and Huisman 2004: 98, fig. 33C.
Vouchers: UPF 1656; WSS 178 St. 1; UPF 1481 St. 2, 1566
St. 16
Australian Systematic Botany
373
and South and Skelton (2003: 540) reported C. biserrulata
fronds to be thin and delicate, but Wallis plants were found
to be quite coriaceous with thick assimilators, while still
possessing conspicuous double rows of marginal teeth (cf.
Payri et al. 2002, pl. 1 fig. 1). Joly and Semir (1973)
described a further variety of C. brachypus from Brazil, var.
nordestina, which has undulate margins and lacks marginal
proliferations. The latter authors appropriately noted that it
could represent a separate species, and it may be quite
distinct from the Pacific material.
Codium bulbopilum Setchell, 1924: 173, fig. 38; 1926: 84,
pl. 11 fig. 1, pl. 12 fig. 2. Jones and Kraft 1984: 26, figs 4,
5B–F; N’Yeurt 1996: 384, figs 42, 45; Kraft 2000: 591,
fig. 31F; Skelton and South 2002a: 165, fig. 25F; Littler and
Littler 2003: 212. Voucher: UPF 1317, St. 10
Caulerpa cupressoides (Vahl) C. Agardh var. mamillosa
(Montagne) Weber-van Bosse, 1898: 332, pl. 28, fig. 6.
Sartoni 1978: 402, fig. 2b; Coppejans and Prud’homme van
Reine 1992: 679, figs 3A, 8B; N’Yeurt 1996: 377, figs 25,
35; Kraft 2000: 597, fig. 32C, D; Payri et al. 2002: 44, pl. 1
fig. 6; Littler and Littler 2003: 216. Vouchers: UPF 1399,
1400 St. 19, 1456 St 26, 1242, 1243, 1245 St. 6
Comments: the Wallis material agrees with the
hummock-like habit of Lord Howe plants in Kraft (2000).
The relationship of Pacific records of C. geppiorum (e.g.
Payri et al. 2000: 102) to C. bulbopilum remain unclear
pending molecular studies (Skelton and South 2002).
Caulerpa nummularia Harvey ex J. Agardh, 1873: 38.
Littler and Littler 1997: 105, fig. 150; Littler and Littler
2003: 214; South and Skelton 2003: 542, fig. 3; Abbott and
Huisman 2004: 121, fig. 44A. Vouchers: UPF 1363 St. 8,
1390 St. 17; WSS 234 St. 35
Codium mamillosum Harvey, 1855: 565. Womersley 1984:
230, figs 77A, 78A; Abbott 1986: 162, fig. 1; N’Yeurt 2001:
720, figs 48, 50, 69; Littler and Littler 2003: 212; Abbott and
Huisman 2004: 109, fig. 39A–B. Voucher: UPF 1650, WSS
162 St. 1
Comments: this species is distinguished from superficially
similar C. peltata by its crenulated disc-like assimilators
which are tiered, as opposed to smooth, single assimilators in
the latter species (Kraft 2000; Abbott and Huisman 2004).
Codiaceae
Comments: a common species at depths below 20 m on the
outer reef slope, it can form large dominant covers at depths
of up to 100 m in many parts of the South Pacific such as Fiji
and McDonald Seamount (ADR N’Yeurt unpubl. obs).
Caulerpa peltata Lamouroux, 1809b: 332. Weber-van Bosse
1898: 373, pl. 31 fig. 9; Kraft 2000: 601, fig. 33E; N’Yeurt
2001: 714; Skelton and South 2002a: 163, fig. 25B.
Vouchers: UPF 1550 St. 1, 1195 St. 3, 1362 St. 8, 1708
St. 17, 1429 St. 23, 1513 St. 31, 1272 St. 5, 1533 St. 12
Caulerpaceae
Comments: Caulerpa peltata is distinguished from
C. racemosa var. peltata (Lamouroux) Eubank by having
thin stolons with simple, never compound, disk-like
assimilators which never bear clavate ramuli. The two
entities are best kept separate until genetic and molecular
studies provide a definitive answer on their status (Kraft
2000).
Caulerpa biserrulata Sonder, 1871: 64, pl. 2 figs 10–12.
Coppejans and al. 1995a: 74, figs 3, 4; Kraft 2000: 595,
fig. 32A, B; Littler and Littler 2003: 214 (= Caulerpa
brachypus Harvey forma parvifolia (Harvey) Cribb sensu
Cribb 1958: 209, figs 1–7; Coppejans and Meinesz 1988:
184, figs 35–38; Coppejans and Prud’homme van Reine
1992: 673, figs 1A–C, 7A–B; N’Yeurt 2001: 711, fig. 40;
Payri et al. 2002: 44, pl. 1 fig. 1). Vouchers: UPF 1360 St. 8,
1295 St. 13, 1402 St.14, 1453 St. 31
Comments: the Wallis plants have both simple, smooth and
divided, toothed uprights, forming a continuum which
circumscribes Caulerpa brachypus forma parvifolia (Cribb
1958b) and conforms with plants described from Papua New
Guinea by Coppejans et al. (1995a). Caulerpa brachypus
forma parvifolia was put in synonymy with C. biserrulata
(Coppejans 1992: 388), and is distinguished from
superficially similar C. brachypus by its double or triple rows
of spines along the assimilator margins. Kraft (2000: 597)
Caulerpa racemosa (Forsskål) J. Agardh, 1873: 35. Sartoni
1978: 406, fig. 4c; Coppejans and Meinesz 1988: 191,
figs 22, 23; Coppejans 1992: 397, fig. 4C; Coppejans and
Prud’homme van Reine 1992: 698, fig. 18A–B; Coppejans
and al. 1995a: 78, fig. 7; N’Yeurt 1996: 378, figs 28, 36;
Kraft 2000: 602, fig. 34A–D; Payri et al. 2000: 94; N’Yeurt
2001: 714, fig. 43; Payri et al. 2002: 44, pl. 1 fig. 2; Littler
and Littler 2003: 226; Abbott and Huisman 2004: 122,
fig. 44B, C. Vouchers: UPF 1251 St. 7, 1298 St. 13, 1403,
1404 St. 19, 1451 St. 31
Comments: includes C. racemosa vars peltata and uvifera;
see Silva et al. (1996).
374
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Caulerpa serrulata (Forsskål) J. Agardh, 1837: 174. Sartoni
1978: 408, fig. 5a; Meñez and Calumpong 1982: 9, pl. 2E;
Coppejans and Meinesz 1988: 192, figs 27, 28; Coppejans
1992: 403, fig. 7; Coppejans and al. 1992: 701, fig. 20B;
1995a: 78, figs 9, 10; N’Yeurt 1996: 382, figs 30, 40; Payri
et al. 2000: 94; Leliaert et al. 2001: 452, fig. 1; N’Yeurt
2001: 715, fig. 44; Payri et al. 2002: 44, pl. 1 fig. 3; Skelton
and South 2002a: 164, figs 24B–D; Littler and Littler 2003:
230; Abbott and Huisman 2004: 123, fig. 45A. Vouchers:
UPF 1216, 1654 St. 1, 1232, 1233, 1234, 1235, 1236 St. 6,
1320 St. 10, 1296 St. 13, 1405 St. 20
Caulerpa sertularioides (S. G. Gmelin) Howe, 1905: 576.
Sartoni 1978: 410, fig. 5b; Coppejans 1992: 404; Coppejans
and Prud’homme van Reine 1992: 704, fig. 21A; Coppejans
and al. 1995a: 80, fig. 8; Littler and Littler 1997: 107,
fig. 154; Payri et al. 2000: 96; N’Yeurt 2001: 716, fig. 42;
Payri et al. 2002: pl. 1 fig. 4; Littler and Littler 2003: 232;
Abbott and Huisman 2004: 124, fig. 45B, C. Vouchers: UPF
1365 St. 8, 1237, 1238, 1239, 1240, 1241, 1244 St. 6, 1392
St. 19
Caulerpa taxifolia (Vahl) C. Agardh, 1817: 22. Sartoni
1978: 410, fig. 5c; Coppejans 1992: 406, fig. 8A–G;
Coppejans and Prud’homme van Reine 1992: 706, figs 6B,
22B; South and N’Yeurt 1993: 122, fig. 17; Coppejans and
al. 1995: 80, fig. 11; Littler and Littler 1997: 109, fig. 155;
Kraft 2000: 604, figs 34E, 35A; Payri et al. 2000: 98;
Leliaert et al. 2001: 452, fig. 2; Payri et al. 2002: 44, pl. 1
fig. 5; Littler and Littler 2003: 234; Abbott and Huisman
2004: 124, fig. 46A-B. Vouchers: UPF 1196, 1692 St. 3,
1710 St. 17, 1393, 1394, 1395, 1396, 1397, 1398 St. 19
Comments: Coppejans and Prud’homme van Reine (1992:
706) reported an ecad ‘mexicana’ of C. taxifolia from
Indonesia, with partly compressed rachi, and basally
constricted compressed pinnae. Later, Littler and Littler
(2003: 222) reported a plant from Fiji with similar
characteristics under Caulerpa mexicana Sonder ex Kützing.
Caulerpa taxifolia and C. mexicana are distinct species
(Olsen et al. 1998). Caulerpa mexicana from the Caribbean
is characterised by a flattened main axis bearing
non-constricted pinnae; the fronds also have a darker green
colour with a lustrous sheen not found in C. taxifolia.
(Meinesz et al. 1994; Littler and Littler 2000: 364, fig. on
365, upper). A main difference between the two species
would be in the disposition of the rhizoids: sparse on the
stolon and very thick and dense in C. taxifolia; close together
and with few rhizoids per shoot in C. mexicana. Taylor
(1977) cited C. mexicana to be present in the Hawaiian flora,
but after re-examination of the herbarium sheets in question
by Abbott and Huisman (2004: 124) they were found to be
not distinct from C. taxifolia. True C. mexicana has yet to be
confirmed from the South Pacific (A. Meinesz, pers. com.)
and present records of this species from Fiji and other
regional localities would best be ascribed to C. taxifolia. It
would appear however to be present on the Andaman Sea
coast, Thailand (Egerod 1975: 5, fig. 23).
Caulerpa taxifolia (Vahl) C. Agardh ecad. tristichophylla
Svedelius 1906: 112. fig. 5. Littler and Littler 2003: 222
(‘Caulerpa mexicana var. pluriseriata’) (= Caulerpa
taxifolia var. asplenioides Harvey 1863: pl. 178). Voucher:
UPF 1711 St. 17
Comments: this ecad is distinguished from the typical form
of C. taxifolia by its three-dimentional, often radial
arrangement of pinnulae on erect assimilators. Quite
commonly, both two-dimentionally and three-dimentionally
beset assimilators are found on the same plant. Also reported
from Fiji (ADR N’Yeurt, unpubl. data), and Mangareva
Island, French Polynesia (Weber-van Bosse 1910 (as
C. taxifolia f. asplenioides); ADR N’Yeurt and CE Payri
unpubl. data). This appears to be a shallow-water ecotype of
C. taxifolia, rather than an infraspecific taxon (A Meinesz,
pers. comm.). Plants from Fiji reported by Littler and Littler
(2003) as Caulerpa mexicana var. pluriseriata and
collections in SUVA-A examined by the first author conform
to the Wallis material as C. taxifolia ecad. tristichophylla as
understood here (see comments above on C. taxifolia).
Weber-van Bosse (1898: 292; 1910: 2) reported this ecad
under C. taxifolia f. asplenioides, based on C. asplenioides
Greville (1853: figs 1, 2, pl. I) and C. taxifolia var.
asplenioides (Greville) Harvey (1863: pl. 178). The latter
forms and varieties were invalidated by Meinesz et al.
(1984): 108) since the original description of C. asplenioides
was found to represent C. mexicana.
Halimedaceae
The systematics of the Halimedaceae has recently been
revised in molecular studies (Kooistra et al. 2002;
Verbruggen and Kooistra 2004).
Halimeda borneensis W.R. Taylor 1975: 81, figs 1, 2.
(Misapplied name: Halimeda simulans Howe: Valet 1968:
48, pl. 9(4) fig. 3; Hillis 1980: 103, fig. 26; N’Yeurt 1996:
390, figs 76, 85; Littler and Littler 1997: 113, fig. 163;
N’Yeurt 2001: 274); Hillis 1980: 105, fig. 27; Coppejans
et al. 2001b: 411; Littler and Littler 2003: 240. Vouchers:
UPF 1779 St. 3, 1781 St. 4, 1179, 1784, 1785, 1786 St. 5,
1782, 1783 St. 9, 1790 St. 14
Comments: previous Pacific records of Halimeda simulans
are referable to H. borneensis (H. Verbruggen pers. com.) as
H. simulans is strictly an Atlantic species (Kooistra et al.
2002: 134).
Halimeda distorta (Yamada) Hillis 1968 : 33, figs 4, 6(2).
(Misapplied name: Halimeda copiosa Goreau et Graham:
Abbott and Huisman 2004: 131, fig. 49A). Hillis 1980: 120,
fig. 34; 1985: 11; Verheij and Prud’homme van Reine 1993:
135, pl. 5 fig. 4; Payri et al. 2000: 108. Vouchers: UPF 2733
Marine plants from Wallis
St. 1, 1172 St. 10, 1181, 1819 St. 11, 1823 St. 24, 1822
St. 31, 1818 St. 38
Comments: new molecular data suggests that previous
Pacific records of H. copiosa are referable to H. distorta, as
the former is exclusively an Atlantic Ocean species (H.
Verbruggen, pers. comm.). H. distorta is characterised by
having a single, basal holdfast and heavily calcified,
contorted segments, which distinguishes it from
superficially similar H. hederacea which has multiple
rhizoidal holdfasts and keeled, uncontorted segments. The
latest molecular evidence points to a merger of H. distorta
and H. hederacea, but it is felt that further research might
separate the two species again (H. Verbruggen, pers. comm.).
Halimeda gracilis Harvey ex J. Agardh, 1887: 82.
Coppejans et al. 1995: 86, fig. 24; Leliaert et al. 2001: 453,
figs 3, 9–10; Littler and Littler 2003: 246; Abbott and
Huisman 2004: 133, fig. 49F. Vouchers: UPF 1167, 1820,
1821 St. 30
Halimeda hederacea (Barton) Hillis 1968: 30, figs 3, 6:1,
6:4-8. (Misapplied name: Halimeda copiosa Goreau et
Graham: Coppejans et al. 1995: 84, fig. 21). Payri et al. 2002:
46, pl. 2 fig. 1; Littler and Littler 2003: 242. Vouchers: UPF
1823 St. 29, 1822 St. 36 (= Halimeda opuntia var. hederacea
(Barton) Hillis 1959: 360, pl. 2 fig. 7, pl. 5 fig. 4; Taylor 1950:
81, pl. 40 fig. 1; N’Yeurt 1996: 389, figs 70c, 84)
Comments: This species is often listed as a variety or
synonym of H. opuntia, but we prefer to maintain its
specietal status in light of new molecular evidence (Kooistra
et al. 2002; Verbruggen and Kooistra 2004). Additionally, it
has a sprawling habit with smooth, shiny segments, as
opposed to the clumped habit with dull, rough segments of
H. opuntia.
Halimeda incrassata (Ellis) Lamouroux 1816: 307; Hillis
1959: 365, pl. 4 figs 1, 2, pl. fig. 21, pl. 6 figs 21–24, pl. 12;
Egerod 1974: 147, figs 62–64; Littler and Littler 1997: 111,
fig. 159; Payri et al. 2000: 110; N’Yeurt 2001: 721, figs 26,
37; Payri et al. 2002: 46, pl. 2 fig. 3; Littler and Littler 2003:
246; Abbott and Huisman 2004: 133, fig. 50A. Vouchers:
UPF 1779 St. 3, 1185, 1781 St. 6, 1179, 1784, 1785, 1786 St.
7, 1782, 1783 St 9
Halimeda macroloba Decaisne, 1841: 118. Hillis 1959: 375,
pl. 3 fig. 3, pl. 5 figs 19, 20, pl. 6 fig. 17, pl. 12; 1980: 108,
fig. 28; Egerod 1974: 148, figs 65–68; 1975: 61, fig. 33;
Coppejans et al. 1995: 86, fig. 25; Payri et al. 2000: 112;
N’Yeurt 2001: 722, fig. 38; Payri et al. 2002: 46, pl. 2 fig. 4;
Littler and Littler 2003: 248; Abbott and Huisman 2004:
135, fig. 50B. Vouchers: UPF 1178, 1802 St. 6, 1803 St. 7,
1176, 1800, 1801 St. 8, 1799 St. 20
Comments: this species is the largest Halimeda in Wallis,
with characteristic bulbous holdfast and segments to 2 cm in
diameter.
Australian Systematic Botany
375
Halimeda micronesica Yamada, 1941: 121, fig. 15. Hillis
1959: 364, pl. 3 fig. 1, pl. 5 figs 13, 14, pl. 6 fig. 2, pl. 9;
Wynne 1993: 22, fig. 10; N’Yeurt 1996: 387, figs 69, 81–82;
Payri et al. 2000: 112; Payri et al. 2002: 46, pl. 2 fig. 5;
Littler and Littler 2003: 250. Vouchers: UPF 1832, 1833,
1834 St. 10, 1832 St. 35
Comments: the fan-shaped, fused basal segment of this
species is distinctive in the field. Common in neighbouring
Rotuma Island (N’Yeurt 1996), it was recently also found in
Fiji (Littler and Littler 2003).
Halimeda minima (W. R. Taylor) Hillis, 1968: 32. Hillis
1980: 113, fig. 30; Payri et al. 2000: 114; N’Yeurt 2001: 723,
figs 25, 36; Littler and Littler 2003: 250. Vouchers: UPF
1169 St. 1, 1685 St. 3, 1177, 1792 St. 10, 1793 St. 9, 1794
St. 12, 1389 St. 17, 1797 St. 22, 1795 St. 23, 1173, 1787
St. 24, 1790 St. 25, 1796, 1798 St. 29, 1168 St. 30, 1788
St. 35, 1171 St. 36, 1791 St. 38
Halimeda opuntia (Linnaeus) Lamouroux, 1816: 308. Hillis
1959: 359, pl. 2 figs 7, 8, pl. 5 figs 3, 4, pl. 6 fig. 6, pl. 7
fig. 3, pl. 10; Egerod 1974: 147, figs 59–61; Coppejans et al.
1995: 86, fig. 27; N’Yeurt 1996: 388, figs 70a–c, 83; Littler
and Littler 1997: 111; Payri et al. 2000: 114; N’Yeurt 2001:
723; Payri et al. 2002: 46, pl. 2 fig. 2; Skelton and South
2002a: 165, fig. 25C–D; Littler and Littler 2003: 252;
Abbott and Huisman 2004: 135, fig. 50C. Vouchers: UPF
1170, 1805, 1827, St. 1, 1804 St. 3, 1807 St. 4, 1809 St. 5,
1811 St. 6, 1180, 1810 St. 7, 1812 St 6, 1184, 1808, 1829,
1830, 1831 St. 10, 1182 St. 11, 1813 St. 13, 1385 St. 16,
1814 St. 17, 1824 St. 23, 1174, 1815, 1825, 1826 St. 24,
1828 St. 35
Halimeda taenicola W. R. Taylor, 1950: 86, 207, pl. 46
fig. 1. Hillis 1959: 354, pl. 2 fig. 6, pl. 5 fig. 12, pl. 6 fig. 14,
pls 11, 14; Hillis 1980: 139, fig. 42; N’Yeurt 1996: 390,
figs 72, 74, 86; Payri et al. 2000: 116; Payri et al. 2002: 46,
pl. 2 fig. 6; Littler and Littler 2003: 252. Vouchers: UPF
1817 St. 10, 1175, 1816 St. 9
Comments: this normally widely distributed tropical species
was in Wallis restricted to the barrier reef on the eastern part
of the island.
Udoteaceae
Chlorodesmis fastigiata (C. Agardh) Ducker, 1969: 17,
fig. 1. Coppejans and Prud’homme van Reine 1989: 127, pl.
3 figs 1–4, 12; Payri et al. 2000: 120; Coppejans et al.
2001b: 421, figs 22–31; N’Yeurt 2001: 726, fig. 73; Payri
et al. 2002: 44, pl. 1 fig. 8; Skelton and South 2002a: 165,
fig. 26B; Littler and Littler 2003: 238. Vouchers: UPF 1299
St. 11, 1307 St. 12, 1457 St. 31
Chlorodesmis hildebrandtii A et E. S. Gepp, 1911: 16, pl. 8
figs 74–75. Egerod 1974: 143, figs 44–49; Coppejans and
Prud’homme van Reine 1989: 129, pl. 3 figs 5–11; N’Yeurt
1996: 393, figs 54, 63; Coppejans et al. 2001b: p, 422,
376
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
figs 15–21; Littler and Littler 2003: 240. Vouchers: UPF
1267 St. 7; WSS 67 St. 10, 1545 St. 12
Comments: crystals were not seen in Wallis plants.
Chlorodesmis hildebrandtii is distinguished from C.
fastigiata by its truncated, equally constricted dichotomies;
the latter species has unequal constrictions above
dichotomies. Abbott and Huisman (2004: 138) follow the
opinion of Ducker (1967) in ascribing previous Pacific
records of C. hildebrandtii to C. caespitosa J. Agardh, but
other authors (Coppejans et al. 2001; Littler and Littler
2003) retain the name for Pacific material.
Rhipidosiphon javensis Montagne, 1842a: 15. Littler and
Littler 1990: 35; Verheij and Prud’homme van Reine 1993:
140, pl. 7 fig. 6; Wynne 1993: 23, fig. 14; N’Yeurt 1996:
394, fig. 61; Payri et al. 2000: 120; Coppejans et al. 2001b:
422, figs 32–34; N’Yeurt 2001: 726, fig. 33; Littler and
Littler 2003: 254; Abbott and Huisman 2004: 140,
fig. 52C–D (= Udotea javensis (Montagne) A. et E. S. Gepp
1904: 363; Dawson 1954: 395, fig. 13b–c; Egerod 1975: 58,
figs 28–29; Coppejans and Prud’homme van Reine 1989:
139, pl. 10 figs 3–9). Vouchers: UPF 1280, 1604 St. 4, 1275
St. 5; WSS 120 St. 8, 35 St. 11, 72 St. 12
Rhipilia tomentosa Kützing, 1858: 12, pl. 28I. Børgesen
1913: 92, fig. 76a–d; Bucher and Norris 1995: 2, figs 1–2;
Millar and Kraft 2001: 25, figs 17, 18, 21–29 (= Rhipilia
tenaculosa A et E. S. Gepp 1911: 56, figs 130–133; N’Yeurt
2001: 729, figs 51a–d, 70; Payri et al. 2002: 44, pl. 1 fig. 9).
Vouchers: UPF 1200, 1491, WSS 31 St. 2; UPF 1530 St. 10,
1333, 1486 St. 9, 1376 St. 16, 1414, 1628 St. 22, 1442, 1506
St. 29, 1720 St. 30
Comments: after examining the relevant type material, Millar
and Kraft (2001: 26) put R. tenaculosa in synonymy with
R. tomentosa. This species is also found in nearby Fiji
(N’Yeurt 2001). According to Millar and Kraft (2001), Fiji
(178–180°E) appears to be the distributional limit eastward
for the genus Rhipilia in the Pacific, but since Wallis Island
(176° W) is slightly to the north-east of Fiji it would seem
that the limit has been reset to the latter locality. However
that distinction now belongs to the Phoenix Islands, (part of
Kiribati), whose eastern limit is 170°30′ W. Indeed, South
et al. (2001: 565) reported three species of Rhipilia from the
Phoenix Islands (R. diaphana, R. geppii, and R. orientalis),
but interestingly R. tomentosa is not listed from that locality,
nor are any of the Phoenix species found in Wallis. Recent
collections from Samoa (Skelton and South 2002) do not
include any Rhipilia species; however other important
nearby localities such as Tonga remain largely unknown
phycologically.
Rhipiliopsis howensis G. T. Kraft, 1986: 55, figs 22–30;
2000: 620, fig. 38F–G. N’Yeurt 2001: 729, fig. 77a–b.
Voucher: WSS 50, St. 29
Comments: this is the second report of this species outside of
its type locality of Lord Howe Island (Australia); it was also
reported from the Suva Lagoon, Fiji (N’Yeurt 2001).
Ochrophyta (= Heterokontophyta)
The notion of a division of golden-brown algae has been the
subject of much controversy since its inception by Luther
(1899, ‘Heterokontae’, or having unequal flagella on
gametes), as pointed out in de Reviers (2003: 148). van den
Hoek et al. (1995) proposed to restrict the term
‘Heterokontophyta’ for brown algae with tripartite hairs on
the longer flagella (Class Phaeophyceae), a definition
contested by Cavalier-Smith (1995) who argued for the
replacement term ‘Ochrophyta’ on the grounds that van den
Hoek et al. (1995) did not apply the rules of the International
Code of Botanical Nomenclature in their diagnosis. We use
Ochrophyta here as adopted by Wynne (2001) and de Reviers
(2003).
Phaeophyceae
Sphacelariales
Sphacelariaceae
Sphacelaria tribuloides Meneghini, 1840: 2. Littler and
Littler 1997: 71, fig. 89; Draisma et al. 1998: 189, fig. 22;
Abbott and Huisman 2004: 190, fig. 72D–E. Voucher: WSS
183, St. 40
Dictyotales
Dictyotaceae
Dictyopteris repens (Okamura) Børgesen, 1924: 265,
fig. 13. Allender and Kraft 1983: 107, figs 19A–B;
Coppejans et al. 1995b: 178, figs 3–4; N’Yeurt 1996: 399,
figs 91–92; Payri et al. 2000: 128; Littler and Littler 2003:
166; Abbott and Huisman 2004: 197, fig. 75E–F. Vouchers:
WSS 176 St. 1; UPF 1473 St. 5, 1625 St. 22, 1430, 1431
St. 23, 1526 St. 41
Dictyota bartayresiana Lamouroux, 1809a: 43. Allender
and Kraft 1983: 112, figs 21E–F, 22D, 23A
(‘D. bartayresii’); Ajisaka and Enomoto 1985: 38,
fig. 1K–L; Coppejans et al. 1995b: 180, fig. 8; De Clerck
and Coppejans 1997: 416, fig. 9; Payri et al. 2000: 132; De
Clerck et al. 2001; Payri et al. 2002: 48, pl. 3 fig. 1; Abbott
and Huisman 2004: 202, fig. 77A. Vouchers: UPF 1253,
1254, 1640, 1643 St. 7, 1345 St. 8, 1435 St. 25
Comments: this species forms ball-like hemispherical tufts, a
distinctive feature of this species (Abbott and Huisman
2004).
Dictyota divaricata Lamouroux, 1809a: 43. Allender and
Kraft 1983: 112, figs 21C–D, 22B–C; Ajisaka and Enomoto
1985: 37, fig. 1H–I; Coppejans et al. 1995b: 182, fig. 19;
Payri et al. 2000: 134. Vouchers: UPF 1212 St. 3, 1327, 1578
St. 9, 1614 St. 16
Marine plants from Wallis
Dictyota friabilis Setchell, 1926: 91, pl. 13 figs 4–7. pl. 20
fig. 1; Dawson 1954: 401, fig. 16a–b; Ajisaka and Enomoto
1985: 37, fig. 1J; N’Yeurt 1996: 400, figs 93–94, 100–101;
Payri et al. 2000: 132; N’Yeurt 2001: 742; Littler and Littler
2003: 168; Abbott and Huisman 2004: 205, fig. 77E.
Vouchers: UPF 1548, 1660 St. 1, 1681, 1698 St. 3; WSS 159
St. 7
Lobophora variegata (Lamouroux) Womersley ex Oliveira,
1977: 217. Allender and Kraft 1983: 81, figs 4G–H, 5A–B;
N’Yeurt 1996: 401, figs 105–106; Littler and Littler 1997:
79, fig. 99a–b; Payri et al. 2000: 136; Payri et al. 2002: 48,
pl. 3 fig. 2; Skelton and South 2002a: 158, fig. 18C–D, G;
Littler and Littler 2003: 172; Abbott and Huisman 2004:
209, fig. 80A–C. Vouchers: UPF 1549, 1651 St. 1, 1192,
1688 St. 3, 1597 St. 4, 1318 St. 10, 1318, 1528 St. 10, 1326
St. 9, 1447 St. 30, 1516 St. 41
Padina boryana Thivy in W. R. Taylor, 1966: 355, fig. 2.
Payri et al. 2000: 138; Abbott and Huisman 2004: 213,
fig. 81D–F (= Padina tenuis Bory 1827: 590; Egerod 1974:
150, fig. 84; Allender and Kraft 1983: 83, figs 5D–E, 6A;
N’Yeurt 1996: 402, fig. 108f). Vouchers: UPF 1255, 1256,
1644, WSS 151 St. 7; UPF 1341, 1342, 1343, 1344 St. 8,
1409 St. 37, 1672 St. 43
Padina melemele Abbott et Magruder in Abbott, 1996: 143,
figs 1–3. Coppejans et al. 1995b: 184, fig. 24 (‘Padina sp.’);
Wynne 1998: 287; Coppejans et al. 2001: 29; Payri et al.
2002: 48, pl. 3 fig. 3; Littler and Littler 2003: 174; Abbott
and Huisman 2004: 217, fig. 83A. Voucher: UPF 1384 St. 16
Comments: this deepwater species of Padina is characterised
by a bright yellow to orange superior surface (surface
towards which margin is curled; see Wynne 1998: 273) and
a bright white calcified inferior surface. Originally described
from Hawaii, it is now reported from Papua New Guinea
(Coppejans et al. 2001), Fiji (N’Yeurt unpubl.; Littler and
Littler 2003), New Caledonia (Payri unpubl.), Rapa Island,
Austral Group, French Polynesia (ADR N’Yeurt and
CE Payri, unpubl. data) and Wallis Island (this study).
Padina melemele may be quite widespread in the tropical
Pacific, but undercollected because of its inconspicuous
subtidal habitat.
Sargassum polycystum C. Agardh, 1824: 304. Chiang et al.
1992: 36, figs 1–12; N’Yeurt 1996: 403, figs 97, 108a–c;
N’Yeurt 2001: 745; Payri et al. 2002: 48, pl. 3 fig. 4; Skelton
and South 2002a: 160, fig. 21; Littler and Littler 2003: 184.
Vouchers: UPF 1346, 1347 St. 8, 1460 St. 34
Comments: the characteristic knob-like proliferous
outgrowths (elevated cryptostomata) on the distinct main
axis, and large number of small vesicles on mature plants, are
diagnostic for this species (Chiang et al. 1992). The only
species of Sargassum and the largest algae found in Wallis,
plants growing in the deep waters of Mata-Utu harbour
attained up to 2 m in length. Its abundance at a port of entry
Australian Systematic Botany
377
raises questions as to whether this represents an introduced
species. From the literature, it is also present (around Wallis)
in Fiji, New Caledonia, Samoa and Tonga.
Fucales
Sargassaceae
Turbinaria conoides (J. Agardh) Kützing, 1860: 24, pl. 66
figs IIe, f. Barton 1891: 217, pl. LIV figs 1, 7–10, pl. LV
figs 1–6, 8–10; Coppejans et al. 1995b: 192, fig. 36; Payri
et al. 2002: 48, pl. 3 fig. 5; Littler and Littler 2003: 184.
Voucher: UPF 1461, St. 34
Comments: this species is mainly distinguished from the
usually less rare T. ornata by the absence of inward-directed
set of teeth on the apical crown of branchlets. Plants are also
somewhat less coarse, with thinner crown margins, than
T. ornata. The Wallis material, however, has a few
inward-directed set of teeth on some thalli, but otherwise
conforms to the current species concept of T. conoides. This
was also observed for material of T. conoides from New
Caledonia (CE Payri, unpubl. data). On-going molecular
analyses of the genera in the Fucales should reveal more
information on the phylogeny of this group, and whether
T. conoides is a valid species, or simply an ecomorph of
T. ornata.
Turbinaria ornata (Turner) J. Agardh, 1848: 266. Barton
1891: 219, pl. LIV figs 11–12, pl. LV fig. 7; Dawson 1954:
405, fig. 21; Coppejans et al. 1995b: 192, fig. 38; N’Yeurt
1996: 405; Payri et al. 2000: 148; N’Yeurt 2001: 745; Wynne
2001: 365, fig. 32; Payri et al. 2002: 48, pl. 3 fig. 6; Skelton
and South 2002a: 160, fig. 19E; Littler and Littler 2003: 186;
Abbott and Huisman 2004: 242, fig. 95. Vouchers: UPF 1222
St. 1, 1252 St. 7, 1316 St. 10
Comments: this species was not very common in the stations
studied. The apical crown of branches has a distinct row of
inward-directed teeth, and the branches are generally thicker
than T. conoides (see comments above for latter species).
Rhodophyta
Porphyridiales
Porphyridiaceae
Chroodactylon ornatum (C. Agardh) Basson, 1979: 67, pl.
IX fig. 52. Abbott 1999: 42, fig. 1A. Voucher: WSS 140 St. 4
Stylonema alsidii (Zanardini) K. Drew, 1956: 72. Abbott
1999: 44, fig. 1B–C. Voucher: WSS 08 St. 5
Erythropeltidales
Erythrotrichiaceae
Erythrotrichia carnea (Dillwyn) J. Agardh, 1883: 15, pl. I
figs 8–10. Abbott 1999: 45, fig. 1E; N’Yeurt 2001: 746,
fig. 106. Voucher: WSS 10 St. 5
378
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Acrochaetiales
Acrochaetiaceae
Acrochaetium barbadense (Vickers) Børgesen, 1915: 45.
Abbott 1999: 52, fig. 3B. Voucher: WSS 99 St. 16
Acrochaetium kurogii (Lee et Lindstrom) Lee et Lee, 1988:
128. Lee and Yoshida 1997: 186, fig. 10A–E. Voucher: WSS
52 St. 29
Comments: this species is epiphytic on the brown alga
Dictyopteris repens. This minute, elegant plant has opposite
branching from every axial cell, with stellate chloroplasts
and a central pyrenoid. The visible parts of the thallus grow
parallel to the host surface, in a characteristic brittle-star
pattern. The Wallis material is in good agreement with the
description of the type species, and illustrations in Lee and
Yoshida (1997). This appears to be the first report of the
species outside of Japan.
Pterocladiella caerulescens (Kützing) Santelices et
Hommersand, 1997: 118. Abbott 1999: 197, fig. 51E; Payri
et al. 2000: 168. Vouchers: UPF 1645, WSS 155 St. 7
Pterocladiella caloglossoides (Howe) Santelices, 1998: 244.
Abbott 1999: 198, fig. 52A–B; Littler and Littler 2003: 64.
Voucher: UPF 1664 St. 1; WSS 133 St. 8; UPF 1574, WSS
101 St. 16; UPF 1704 St. 20, 1624 St. 22; WSS 232 St. 36
Gelidiellaceae
Gelidiella acerosa (Forsskål) Feldmann et Hamel, 1934:
533. Price and Scott 1992: 25, fig. 4A–E; Abbott 1999: 202,
fig. 53A–C; Payri et al. 2000: 170; N’Yeurt 2001: 760;
Littler and Littler 2003: 56. Vouchers: UPF 1225, 1655 St. 1,
1472 St. 5, 1263, 1264, 1635, 1636 St. 7, 1370 St. 8, 1499,
WSS 36 St. 11; UPF 1539, 1544 St. 12, 1391 St. 17, 1705
St. 20, 1514 St. 31
Nemaliales
Gracilariales
Galaxauraceae
Gracilariaceae
Actinotrichia fragilis (Forsskål) Børgesen, 1932: 6. Abbott
1999: 64, fig. 7A–C; Payri et al. 2000: 160; N’Yeurt 2001:
749, figs 109, 117; Payri et al. 2002: 50, pl. 4 fig. 4; Littler
and Littler 2003: 62. Vouchers: UPF 1283, 1598 St. 4, 1276
St. 5, 1257, 1634 St. 7, 1302 St. 12, 1379 St. 16, 1620 St. 22,
1433 St. 25
Gracilaria textorii (Suringar) De Toni, 1895: 27. Yamamoto
1978: 123, pls 12–14; pl. 42 figs 5–7; pl. 43 figs 1–4;
N’Yeurt 2001: 772, figs 156, 168; Payri et al. 2002: 50, pl. 4
fig. 11. Vouchers: UPF 1357, 1361 St. 8
Galaxaura fasciculata Kjellman, 1900: 53, pl. figs 1–9; pl.
20 fig. 14. Abbott 1999: 66, fig. 7D–E; Payri et al. 2000:
162; N’Yeurt 2001: 749, fig. 116; Payri et al. 2002: 50, pl. 4
fig. 10. Vouchers: UPF 1215, 1224 St. 1, 1282, 1285 St. 4,
1303 St. 12, 1378 St. 16, 1408 St. 20, 1432, 1744 St. 25
Galaxaura filamentosa Chou in W.R. Taylor, 1945: 139.
Abbott 1999: 67, fig. 7F; Payri et al. 2000: 164; Littler and
Littler 2003: 64. Vouchers: UPF 1717 St. 1, 1600 St. 4; WSS
123 St. 8
Comments: this species was growing amidst seagrass beds,
in a situation similar to that reported for this flattened species
on the neighbouring island of Rotuma (N’Yeurt 1996: 415).
Gracilaria sp. Vouchers: UPF 1696, St. 23
Comments: this unusual, flattened species of Gracilaria
could represent a new species; its definite identification
awaits the discovery of fertile spermatangial plants.
Bonnemaisoniales
Bonnemaisoniaceae
Galaxaura marginata (Ellis et Solander) Lamouroux, 1816:
264. Abbott 1999: 67, fig. 7G–H; Payri et al. 2000: 164;
N’Yeurt 2001: 752, figs 118–122; Littler and Littler 2003:
64. Vouchers: UPF 1284, 1592 St. 4
Asparagopsis taxiformis (Delile) Trevisan, 1845: 45. Abbott
1999: 174, fig. 43A–D; Payri et al. 2000: 160; N’Yeurt 2001:
775, figs 141, 142; Skelton and South 2002a: 141, fig. 6G;
Littler and Littler 2003: 68. Voucher: UPF 1415 St. 22
Galaxaura subverticillata Kjellman, 1900: 48, pl. 3 figs 12;
pl. 20 fig. 17. Abbott 1999: 69, fig. 8F–H. Vouchers: WSS
89 St. 1, UPF 1561 St. 2, 1593 St. 4, 1564 St. 16
Comments: the record represents the minute Falkenbergia
stage in the life cycle of the species; larger thalli of the other
stage presumably occur at other seasons in Wallis.
Comments: we follow the opinion of Abbott (1999) in
retaining the species distinct rather than being synonomyzed
with Galaxaura rugosa, since the Wallisian plants also have
abundant assimilatory filaments throughout the thallus.
Gelidiales
Gelidiaceae
Gelidium pusillum (Stackhouse) Le Jolis, 1863: 139. Abbott
1999: 194, fig. 50A; Payri et al. 2000: 168; N’Yeurt 2001:
760. Vouchers: UPF 1552 St. 1, 1477, WSS 14 St. 5
Cryptonemiales
Halymeniaceae
Cryptonemia umbraticola Dawson, 1959: 43, figs 21F, 22A.
Abbott 1999: 136, fig. 32B; Payri et al. 2000: 174. Vouchers:
UPF 1475, WSS 13 St. 5, UPF 1323, 1325, 1488 St. 9, 1311,
1312, 1313, 1314, 1536, 1537, WSS 74, 75 St. 12; UPF 1387
St. 16, 1463 St. 36
Comments: this species forms loose rosettes at the base of
coral heads, 12 m depth in the lagoon.
Marine plants from Wallis
Cryptonemia yendoi Weber-van Bosse, 1921: 249, fig. 77.
Abbott 1999: 138, fig. 32C–E; Masuda et al. 1999: 449,
figs 1–7. Vouchers: UPF 1579, WSS 108 St. 9; UPF 1535,
1538, WSS 73, 76 St. 12, 42 St. 23; UPF 1508 St. 29
Comments: this species can be distinguished from
C. umbraticola by its erect habit, with a prominent stipe.
Kallymeniaceae
Kallymenia sp. Voucher: UPF 1207 St. 2
Comments: the lack of fertile material and fragmentary
nature of the specimen precludes specietal assignment of this
plant at this stage.
Peyssonneliaceae
Peyssonnelia bornetii Boudouresque et Denizot, 1973;
1975: 27, figs 24–47. Marcot et al. 1976: 237, fig. V; Payri
et al. 2000: 172; N’Yeurt 2001: 777, figs 138, 146.
Vouchers: UPF 1570, WSS 94, 98 St. 16
Peyssonnelia inamoena Pilger, 1911: 311, figs 24, 25.
Boudouresque et Denizot 1975: 58, figs 107–115; Abbott
1999: 156, fig. 38B; Payri et al. 2000: 172; N’Yeurt 2001:
778, figs 139, 140a–e, 143–144; Littler and Littler 2003: 82.
Vouchers: UPF 1204, WSS 221 St. 2; UPF 1304 St. 12,
1627, WSS 146 St 17; UPF 1502 St. 29
Peyssonnelia sp. Vouchers: WSS 180; UPF 1701, WSS 200
St. 3; UPF 1334 St. 9, 1669, 1729 St. 23, 1745 St. 25, 1721
St. 30
Comments: this species is closest to Peyssonnelia conchicola
Piccone et Grunow sensu Marcot et al. 1988: 266; however,
in the species from Madagascar the unicellular rhizoids are
not laterally inserted, and the thallus is only 3–6 layers thick
compared to 7–8 in the Wallis material. Both plants have
terminal cystoliths. Peyssonnelia sp. also has been reported
from the Cook Islands (ADR N’Yeurt unpubl. data).
Gelidiella maschrisiana Dawson, 1957a: 17, fig. 4B. Abbott
1999: 204, fig. 53G–H; Payri et al. 2000: 170. Vouchers:
WSS 127 St. 8; UPF 1605 St. 4, 1741, WSS 218 St. 10
Corallinales
Corallinaceae
Amphiroa foliacea Lamouroux in Quoy et Gaimard, 1824:
628, pl. 93 figs 2, 3. Abbott 1999: 178, figs 44C, 45B; Payri
et al. 2002: 52, pl. 5 fig. 1 (as Amphiroa crassa); Skelton and
South 2002a: 140, fig. 4D; Littler and Littler 2003: 26.
Voucher: UPF 1201 St. 2, 1380 St. 16
Comments: Abbott (1999: 178) states that there exists a high
degree of variation in intergenicula shape and size in
Amphiroa foliacea, which may encompass the species
circumsciption of A. tribulus, A. misakiensis and other taxa.
Amphiroa tribulus would be the earlier name, but a critical
study of its type is needed before making taxonomic
Australian Systematic Botany
379
decisions. In the field however, A. foliacea can be readily
distinguished by its flat, coarse habit with axes 3–5 mm wide,
as opposed to the fragile, thinner branches of A. tribulus.
Amphiroa fragilissima (Linnaeus) Lamouroux, 1816: 298.
Price and Scott 1992: 45, fig. 11A–B; Payri et al. 2000: 182;
Skelton and South 2002a: 140, fig. 4E; Littler and Littler
2003: 26. Vouchers: UPF 1661 St. 1, 1324, 1608 St. 9, 1445,
1512 St. 29
Amphiroa tribulus (Ellis et Solander) Lamouroux ,1816:
302. Payri et al. 2000: 182; N’Yeurt 2001: 761, figs 131,
134–135; Payri et al. 2002: 52, pl. 5 fig. 2; Littler and Littler
2003: 26. Voucher: UPF 1436 St. 25
Hydrolithon farinosum (Lamouroux) Penrose et
Chamberlain, 1993: 295–303, figs 1–19. Gordon et al. 1976:
255, pl. III figs 1–4 (as Fosliella farinosa (Lamouroux)
Howe); Penrose in Womersley 1996: 260, fig. 118A–D; Payri
et al. 2000: 188; Ringeltaube and Harvey 2000: 437,
figs 12–13; Littler and Littler 2003: 30. Voucher: WSS 161
St. 1
Hydrolithon murakoshii Iryu et Matsuda, 1996: 528. Payri
et al. 2000: 190; Littler and Littler 2003: 32. Vouchers: WS
65 St. 1, 284 St. 17
Hydrolithon onkodes (Heydrich) Penrose et Woelkerling,
1992: 83. Gordon et al. 1976: 266, pl. IX figs 1–4;
Ballesteros and Afonso-Carrillo 1995: 209, fig. 12;
Womersley 1996: 261, fig. 119A–D; Payri et al. 2000: 192;
Ringeltaube and Harvey 2000: 438, figs 14–16; Payri et al.
2002: 52, pl. 5 fig. 7. Vouchers: WS 75 St. 1, 1 St. 3, 120
St. 5, 283 St. 17, 399 St. 29, 403 St. 30
Hydrolithon reinboldii (Weber-van Bosse et Foslie in Foslie)
Foslie, 1909: 55. Gordon et al. 1976: 255, pl. III figs 5–6, pl.
IV figs 1–3; Payri et al. 2000: 192; Ringeltaube and Harvey
2000: 439, figs 17–20; Payri et al. 2002: 52, pl. 5 fig. 10;
Littler and Littler 2003: 32. Vouchers: WS 67, 70 St. 1, 161
St. 11, 269 St. 16, 288 St. 17, 297 St. 18
Hydrolithon rupestris (Foslie) Penrose, 1996: 265, fig. 121.
Penrose in Womersley 1996: 265, fig. 121A–C; Payri et al.
2000: 194. Voucher: WS 119 St. 5
Jania adhaerens Lamouroux, 1816: 270. Price and Scott
1992: 48, fig. 12A–C; Abbott 1999: 187, fig. 48A; N’Yeurt
2001: 765; Skelton and South 2002a: 141, fig. 6D–F; Littler
and Littler 2003: 32. Vouchers: UPF 1547, 1727 St. 1, 1682
St. 3, 1716 St. 15, 1520 St. 41
Lithophyllum flavescens Keats, 1997: 357, figs 23–39.
Payri et al. 2000: 204. Voucher: WS 333 St. 23
Lithophyllum insipidum Adey, Townsend et Boykins, 1982:
44, fig. 29A–F. Keats 1997: 352, figs 1–22; Payri et al. 2000:
206. Vouchers: WS 3 St. 3, 282 St. 17
Lithophyllum kotschyanum Unger, 1858: 22, pl V fig. 15.
Gordon et al. 1976: 267, pl. IX figs 5, 6, pl. X figs 1, 2; Adey
380
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
et al. 1982: 37, fig. 24A–F; Ballesteros and Afonso-Carrillo
1995: 207, fig. 2; Payri et al. 2000: 206; Littler and Littler
2003: 36. Vouchers: WS 124 St. 5, 264 St. 16, 294 St. 18
Lithophyllum pygmaeum (Heydrich) Heydrich, 1897b: 412.
Ballesteros and Afonso-Carrillo 1995: 207, fig. 3 (as
Lithophyllum tamiense); N’Yeurt 1996: 414, fig. 115 (as
Lithophyllum tamiense); Ringeltaube and Harvey 2000: 436,
figs 7–11 (as Lithophyllum tamiense); Littler and Littler
2003: 38. Vouchers: WS 118 St. 5, 444 St. 37, 452 St. 38
Comments: forming unattached balls on the lagoon floor, or
attached to coral boulders in the lagoon. According to Silva
et al. (1996: 251), L. pygmaeum includes Lithophyllum
tamiense (Heydrich) (Verheij 1994: 103).
Lithothamnion proliferum Foslie in Weber-van Bosse and
Foslie 1904: 18, fig. 8, pl. I figs 17–20. Keats et al. 1996:
188, figs 1–36 (‘L. prolifer’); Ringeltaube and Harvey 2000:
445, figs 32, 33 (‘L. prolifer’); Payri et al. 2002: 52, pl. 5
fig. 11; Skelton and South 2002a: 141, fig. 5B; Littler and
Littler 2003: 38. Voucher: WS 262 St. 16
Mastophora pacifica (Heydrich) Foslie, 1903: 25.
Woelkerling in Womersley 1996: 248, figs 111A–E,
112A–E, 113A–B; Payri et al. 2000: 196; Ringeltaube and
Harvey 2000: 441, figs 21, 22; Payri et al. 2002: 52, pl. 5
fig. 4; Littler and Littler 2003: 40. Voucher: WS 136 St. 4,
361 St. 24, 396 St. 29
Mesophyllum erubescens (Foslie) Lemoine, 1928: 252.
Gordon et al. 1976: 252, pl. I figs 5–8; Ballesteros and
Afonso-Carrillo 1995: 209, fig. 11; Payri et al. 2000: 208;
Ringeltaube and Harvey 2000: 445, figs 34–37; Payri et al.
2002: 52, pl. 5 fig. 8; Littler and Littler 2003: 42. Vouchers:
WS 158 St. 11, 397 St. 29
Mesophyllum mesomorphum (Foslie) Adey, 1970: 25.
Gordon et al. 1976: 252, pl. II figs 1–3. Voucher: WS 295
St. 18
Neogoniolithon brassica-florida (Harvey) Setchell et
Mason, 1943: 91. Penrose in Womersley 1996: 281,
fig. 129A–D; Ringeltaube and Harvey 2000: 441,
figs 23–27; Payri et al. 2002: 52, pl. 5 fig. 5 (as
Neogoniolithon frutescens). Vouchers: WS 67 St. 1, 4 St. 3,
291, St. 17, 360 St. 24, 376 St. 25, 450 St. 38
Pneophyllum conicum (Dawson) Keats, Chamberlain et
Baba, 1997: 264. Gordon et al. 1976: 259, pl. IV figs 6–8, pl.
VI fig. 1 (as Neogoniolithon conicum (Dawson) Gordon,
Masaki et Akioka); Adey et al. 1982: 13, figs 6–7 (as
Paragoniolithon conicum (Dawson) Adey, Townsend et
Boykins); Payri et al. 2000: 202; Payri et al. 2002: 52, pl. 5
fig. 6; Littler and Littler 2003: 48. Voucher: WS 6 St. 3
Comments: this common Indo-Pacific species with a
tortuous taxonomic history is reported to overgrow and kill
live corals (Keats et al. 1997).
Sporolithon ptychoides Heydrich, 1897: 67–69. Keats and
Chamberlain 1993: 542, figs 1–13; Ballesteros and
Afonso-Carrillo 1995: 209, fig. 13; Payri et al. 2000: 211;
Littler and Littler 2003: 52. Vouchers: WS 66 St. 1106 St. 7,
247 St. 8
Gigartinales
Caulacanthaceae
Caulacanthus ustulatus (Turner) Kützing, 1843: 395.
Wynne 1995: 277, figs 13–15; Abbott 1999: 104,
fig. 20D–F; Masuda et al. 1999: 451, figs 8–11. Vouchers:
UPF 1694, WSS 195 St. 1, 26, 27 St. 4, 67 St. 10; UPF 1715
St. 15
Comments: this species is superficially similar to Hypnea
spinella, Caulacanthus ustulatus is distinguished by its more
flattened axes, and its polygonal to hexagonal cortical cells
in surface view.
Corynocystaceae
Corynocystis prostrata Kraft, in Kraft et al. 1999: 26, figs 6,
45–60. N’Yeurt 2001: 797, figs 107, 108, 217; Skelton and
South 2002a: 142, fig. 7C; Littler and Littler 2003: 70.
Voucher: UPF 1210, 1211, 1606 St. 2, 1322 St. 9, 1382
St. 16, 1623, WSS 144 St. 22; UPF 1446 St. 29
Comments: now found throughout the South Pacific
(including French Polynesia, pers. obs. and In Herb. UPF),
this characteristic alga is in most case associated with an
ascidian which forms a thick white layer on the inferior
surface. The genus, previously listed in the Acrotylaceae, has
recently been transferred to a family of its own based on
molecular studies (see Saunders et al. 2004).
Dumontiaceae
Gibsmithia hawaiiensis Doty, 1963: 458, figs 1–17. Abbott
1999: 108, fig. 22D–G; Payri et al. 2000: 212; Payri et al.
2002: 50, pl. 4 fig. 8. Vouchers: UPF 1205 St. 2; WSS 24
St. 4; UPF 1487 St. 9, 1416 St. 22, 1438, 1439 St. 29, 1462
St. 36
Hypneaceae
Hypnea cervicornis J. Agardh, 1851: 451. Abbott 1999: 113,
fig. 24A. Voucher: UPF 1587, St. 8
Comments: Haroun and Prud’homme van Reine (1993)
reduced H. cervicornis in synonymy with H. spinella (C.
Agardh) Kützing, but we prefer to follow the conservative
opinion of Abbott (1999) to distinguish entities that look
relatively different in the field.
Hypnea charoides Lamouroux, 1813: 132, pl. 10 figs 1–3.
Weber-van Bosse 1928: 449, figs 188, 189; Yamagishi and
Masuda 1997: 136, figs 1–9; 2000: 31, figs 10–15; South
2004: 136, fig. 2. Vouchers: UPF 1261, 1262 St. 7, 1366
St. 8
Marine plants from Wallis
Hypnea pannosa J. Agardh, 1847: 14. Price and Scott 1992:
38, figs 8A–D, 9A; Payri et al. 2000: 222; non N’Yeurt 2001:
780; Skelton and South 2002a: 143, fig. 7D; Littler and
Littler 2003: 76. Vouchers: WSS 182 St. 1; UPF 1676 St. 3,
1279 St. 4, 1268, 1269 St. 5, 1259, 1260 St. 7, 1367, 1586
St. 8, 1424, 1425, 1428, 1671 St. 23, 1434 St. 25
Comments: records of Hypnea pannosa from the Suva
Lagoon, Fiji in N’Yeurt (2001) were relegated to the status of
an undescribed species close to the Japanese Hypnea
charoides-valentiae complex by South (2004: 135). The
Wallis material is in good agreement with H. pannosa from
the Great Barrier Reef (Price and Scott 1992).
Hypnea saidana Holmes, 1896: 256, pl. 11 figs 3a, b.
Wynne 1995: 276, fig. 16; Payri et al. 2000: 224. Vouchers:
UPF 1658 St. 1, 1193, 1194, 1646, 1699 St. 3, 1478, WSS
15, 16 St. 5; UPF 1543 St. 12, 1565 St. 16, 1523 St. 41
Hypnea spinella (C. Agardh) Kützing, 1847a: 23. Price and
Scott 1992: 40, figs 9B, 10A–F; Abbott 1999: 117,
fig. 25B–E; Payri et al. 2000: 224; Littler and Littler 2003:
76. Vouchers: UPF 1647, WSS 194 St. 3; UPF 1274, 1468,
WSS 05, 06 St. 5; UPF 1368, 1588 St. 8, 1573, WSS 99
St. 16; UPF 1732, WSS 39 St. 23; UPF 1484 St. 26; WSS 53
St. 29; UPF 1521 St. 41
Comments: this is the most common minute Hypnea species
in the collections, ubiquitous in most habitats. Based on
culture and molecular studies, Yamagishi and Masuda
(2000) relegated the entity previously known as Hypnea
cervicornis/spinella from Japan to the rank of a new species,
H. flexicaulis. The Wallis material agrees with the concept of
H. spinella in Price and Scott (1992) and Abbott (1999).
Hypnea valentiae (Turner) Montagne, 1841: 161. Weber-van
Bosse 1928: 452, text fig. 190; Børgesen 1934: 17; Littler
and Littler 2003: 76. Vouchers: UPF 1637, WSS 149, 152
St. 7; UPF 1673 St. 43
Comments: Yamagishi and Masuda (1997: 140; 2000)
consider H. valentiae from Japan and Vietnam (Dawson
1954: 436, fig. 47) as belonging to the H. charoidesvalentiae complex. The Wallis material agrees with the
descriptions of H. valentiae in Weber-van Bosse (1928) and
Børgesen (1934), and the two names are retained separate as
a conservative measure until future studies on their
taxonomic status.
Lomentariaceae
Lomentaria corallicola Børgesen, 1939: 113, figs 30–32.
Price and Scott 1992: 63, fig. 18A–E; Payri et al. 2000: 236;
N’Yeurt 2001: 811, figs 223, 224; Littler and Littler 2003:
106. Vouchers: UPF 1663, WSS 171 St. 1
Nemastomataceae
Platoma cyclocolpum (Montagne) Schmitz, 1894: 627.
Masuda and Guiry 1994: 194–201, figs 1–34; Huisman
Australian Systematic Botany
381
1999; N’Yeurt 2001: 781, figs 169–173; Payri et al. 2002:
50, pl. 4 fig. 6. Vouchers: UPF 1440, WSS 45, 46, 47 St. 29
Comments: the largest and most spectacular fleshy red alga
collected from Wallis, the material agrees well with the
description of material from the Canary Islands by by
Masuda and Guiry (1994) and from Australia by Huisman
(1999).
Predaea laciniosa Kraft, 1984: 11, figs 25–35. Payri et al.
2000: 214; Payri et al. 2002: 50, pl. 4 fig. 5; Littler and
Littler 2003: 78. Vouchers: WSS 19 St. 5; UPF 1626, WSS
145 St. 22; UPF 1419, 1482 St. 23
Predaea weldii Kraft et Abbott, 1971: 194, figs 1–15. Kraft
1984: 15, figs 36–42; Abbott 1999: 153, fig. 37D–F;
N’Yeurt 2001: 783, figs 183, 228; Littler and Littler 2003:
78. Vouchers: UPF 1417, 1418, WSS 198 St. 22
Solieriaceae
Meristotheca procumbens Gabrielson et Kraft, 1984: 241,
fig. 14A–D. N’Yeurt 1995: 248, figs 3–10; 1996: 416,
figs 137, 142–147, 203–205; Payri et al. 2000: 220; N’Yeurt
2001: 793, figs 208, 209, 231; Littler and Littler 2003: 92.
Vouchers: UPF 1223, 1719, WSS 204 St. 1, 102 St. 2; UPF
1203 St. 3; WSS 136 St. 4; UPF 1575 St. 9, 1613 St. 16
Comments: this edible species is relatively infrequent and
cryptic in Wallis, presumably the reason why it is not
consumed traditionally as in neighbouring islands such as
Rotuma (N’Yeurt 1995, 1996) where it is locally abundant in
intertidal habitats.
Sarconema filiforme (Sonder) Kylin, 1932: 22. Børgesen
1932: 11, fig. 7; Papenfuss and Edelstein 1974; 32, figs 1–3,
13, 20–25; Payri et al. 2000: 220. Vouchers: UPF 1348,
1349, 1350, 1591, WSS 117 St. 8
Comments: this species was recorded growing amidst
seagrass beds, with other algae. This distinctive species can
be recognised by its lubricous, terete thalli with bifurcate
tips. This is the fourth report for this predominantly Indian
Ocean genus in the South Pacific. Papenfuss and Edelstein
(1974: 41) mention having examined herbarium material
from New South Wales, Australia, and Upolu, Western
Samoa (the latter as Dicranema setaceum var. upolensis
Grunow). It is also found seasonally in Tahiti, French
Polynesia (Payri et al. 2000).
Halymeniales
Halymeniaceae
Halymenia actinophysa Howe, 1911: 509, pl. 34. Abbott
1999: 143, fig. 34A. Vouchers: WSS 134 St. 8; UPF 1611
St. 16
Comments: the Wallis material is in good agreement with the
description of the Type of the species by Howe (1911),
especially the macelike subcortical cells with several blunt
arms.
382
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Prionitis sp. Voucher: UPF 1413, St. 22.
Comments: The material being sterile,
indentification is not possible at this stage.
specific
Sebdeniaceae
Sebdenia sp. Vouchers: WSS 177, 179 St. 1 (tetrasporic);
UPF 1501, 1670 St. 23
Comments: the fragmentary nature of the material precludes
the definite identification of this taxon at the present time,
but it could represent a new species. It is close to the
flattened Sebdenia sp. mentioned from Fiji in N’Yeurt (2001:
806).
Rhodymeniales
Champiaceae
Champia compressa Harvey, 1838: 402. Millar 1990: 371,
figs 30A–D; Masuda et al. 2001: 84, figs 14–17; Skelton and
South 2002a: 143, fig. 7E, F; Littler and Littler 2003: 102.
Vouchers: WSS 84, 181 St. 1, UPF 1691 St. 3; WSS 153
St. 7, 121, 130 St. 8; UPF 1531 St. 10, 1577, WSS 105 St. 9,
222, 239 St. 16; UPF 1723 St. 30
Champia parvula (C. Agardh) Harvey, 1853: 76. Millar
1990: 371, figs 29G, H; Price and Scott 1992: 55,
fig. 14A–E; Abbott 1999: 218, fig. 60A–C; Payri et al. 2000:
238; N’Yeurt 2001: 808, fig. 238; Skelton and South 2002a:
143, fig. 8A–C. Vouchers: WSS 170 St. 1, 156 St. 7
Rhodymeniaceae
Chamaebotrys boergesenii (Weber-van Bosse) Huisman,
1996: 106, figs 35–38, 40–42. Abbott 1999: 228,
fig. 63E–G; Payri et al. 2000: 230; Masuda et al. 2001: 81,
figs 1–6; N’Yeurt 2001: 812, figs 218, 235, 236; Littler and
Littler 2003: 110. Voucher: UPF 1695, WSS 196 St. 3; UPF
1465, WSS 04 St. 5; UPF 1702 St. 9, 1496 St. 11, 1737,
1500, WSS 37 St. 23
Chrysymenia okamurae Yamada et Segawa, 1953: 110,
fig. 3. Abbott 1999: 230, fig. 64C–D. Vouchers: UPF 1572,
WSS 96 St. 16
Coelothrix irregularis (Harvey) Børgesen, 1920: 389. Price
and Scott 1992: 60, fig. 17A–D; Abbott 1999: 233,
fig. 65A–D; Payri et al. 2000: 230. Vouchers: UPF 1638,
1639, WSS 150 St. 7
Gelidiopsis intricata (C. Agardh) Vickers, 1905: 61. Price
and Scott 1992: 51, fig. 13A–F; Abbott 1999: 221, fig. 61A;
Payri et al. 2000: 232; Skelton and South 2002a: 144,
fig. 8D–G. Vouchers: UPF 1493 St. 2, 1641, 1642, WSS 154
St. 7, UPF 1328, 1331, 1580 St. 9, 1310 St. 12, 1374, 1375,
1714 St. 15, 1388 St. 16, 1494 St. 40; WSS 34 St. 37
Gelidiopsis scoparia (Montagne et Millardet) De Toni,
1900: 410. Abbott 1999: 221, fig. 61B; Littler and Littler
2003: 106. Vouchers: UPF 1206 St. 2, 1690 St. 3, 1470 St. 5,
1497 St. 11, 1563, 1618 St. 16, 1739 St. 36, 1495 St. 40
Halichrysis coalescens (Farlow) R.E. Norris et Millar in
R. Norris, 1991: 583, 585, 587, figs 7–13. Abbott 1999: 235,
fig. 66A–C; N’Yeurt 2001: 812, figs 220, 221, 234; Littler
and Littler 2003: 116. Vouchers: UPF 1469, 1480, WSS 18
St. 5; UPF 1738, WSS 211 St. 23
Leptofauchea anastomosans (Weber-van Bosse) R. Norris
et Aken, 1985: 58, figs 10–12. Dawson 1956: 52, fig 49
(‘Rhodymenia anastomosans’). Vouchers: UPF 1571, WSS
95 St. 16; WSH 277, WSS 43, 212 St. 23; UPF 1443, 1509
St. 29
Comments: growing intricately associated with articulated
coralline algae (Amphiroa spp.) to which the blades form
numerous secondary attachments, this species also occurs in
the Cook islands in similar habitats (ADR N’Yeurt, unpubl.
data).
Ceramiales
Ceramiaceae
Anotrichum tenue (C. Agardh) Nägeli, 1862: 399. Abbott
1999: 247, fig. 68D; Masuda et al. 1999: 453, figs 15–23;
Payri et al. 2000: 240; Skelton and South 2002a: 145,
fig. 9A–F. Voucher: WSS 67 St. 10
Comments: Womersley (1998: 344) discusses the range of
characteristics in specimens of the A. tenue complex, which
support the concept of a single species with several varieties
sometimes described as separate species.
Antithamnion decipiens (J. Agardh) Athanasiadis, 1996:
151. Abbott 1999: 250, fig. 69C–D. Voucher: WSS 40,
St. 23
Balliella repens Huisman et Kraft, 1984: 77, figs 16–26.
Price and Scott 1992: 72, figs 20D, 21A–B; Abbott 1999:
257, fig. 71H. Vouchers: WSS 140 St. 4, 69 St. 12, 49 St. 29
Centroceras minutum Yamada, 1944: 42. Dawson 1956: 54,
fig. 54; Ardré 1987: 285, figs 30–37; Wynne 1993: 12,
fig. 7; 1995: 290, fig. 27; Abbott 1999: 262, fig. 73H–I;
N’Yeurt 2001: 817, figs 249, 250. Vouchers: WSS 228 St. 3,
68 St. 10
Comments: reasons to maintain Centroceras minutum as a
separate species from C. clavulatum are given in Wynne
(1993: 12). N’Yeurt (2001: 817) examined slide-mounted
type material of C. minutum in SAP and found it also to
differ from C. clavulatum.
Ceramium codii (Richards) Mazoyer, 1938: 324. Millar
1990: 393, fig. 41D–F; Abbott 1999: 270, fig. 75D–F; Payri
et al. 2000: 246; South and Skelton 2000: 56, figs 11–14.
Vouchers: WSS 08 St. 5, 126 St. 8, 218 St. 10, 81, 82, St. 12
Ceramium flaccidum (Kützing) Ardissone, 1871: 40.
Womersley 1978: 234, figs 4A–D, 14E–H; Price and Scott
1992: 89, fig. 27A–E; Womersley 1998: 410, figs 188E, H,
190A–D; Abbott 1999: 274, fig. 76D–H; Payri et al. 2000:
248; South and Skelton 2000: 65, figs 32–39, 41–44;
Marine plants from Wallis
Australian Systematic Botany
383
N’Yeurt 2001: 817, fig. 253. Vouchers: UPF 1683, 1697,
WSS 188, 199, 228 St. 3; UPF 1585, 1713, WSS 114, 202
St. 8; UPF 1742, WSS 67, 217 St. 10, 48 St. 29
Diplothamnion jolyi van den Hoek 1978: 47; Huisman
1991; Stegenga and Vroman 1987: 411, figs 35, 36; Abbott
1999: 297, fig. 83C. Voucher: WSS 25, St. 4
Comments: the broad rows of basipetal cortical cells and
forcipate apices are characterisctic for this species. This is
the largest and most common Ceramium species in Wallis,
and is locally abundant especially forming reddish, silky
tufts on branches of dead coral.
Euptilota articulata (J. Agardh) Schmitz, 1896: 7.
Womersley 1998: 355, pl. 2 fig. 2, figs 141E, 164, 165;
N’Yeurt 2001: 820, figs 254–263; Payri et al. 2002: 50, pl. 4
fig. 3; Littler and Littler 2003: 122. Vouchers: UPF 1214
St. 3, WSS 139 St. 4, 129 St. 8; UPF 1421, 1422, 1423,
1609, 1621 St. 22, 1449, 1722 St. 30
Ceramium kramerii South et Skelton, 2000: 69, figs 45–51.
Voucher: WSS 126, St. 8
Comments: a minute, distinctive species, epiphytic on larger
algae growing on coral debris.
Ceramium macilentum J. Agardh, 1894: 15. Womersley
1978: 232, figs 3E, 14A–D; Price and Scott 1992: 100,
fig. 31; South and Skelton 2000: 71, figs 52–62. Vouchers:
WSS 194 St. 3, 08 St. 5, 208 St. 23, 48 St. 29
Ceramium sp. aff. C. marshallense Dawson, 1957b: 120,
figs 27a, b. Price and Scott 1992: 102, fig. 32; Wynne 1993:
14, figs 8–9; 1995: 294, figs 38, 39 South and Skelton 2000:
75, figs 63–71, 73. Vouchers: WSS 25, 139 St. 4
Ceramium subdichotomum Weber-van Bosse, 1923: 333,
fig. 125. Wynne 1995: 296, figs 44, 45; 1999: 195, figs 8, 9;
Payri et al. 2000: 250 (‘subdichotum’); South and Skelton
2000: 78, figs 74–79. Voucher: WSS 208, St. 23
Ceramium upolense South et Skelton, 2000: 81, figs 80–88.
Vouchers: WSS 07, 08 St. 5, 203 St. 9, 99 St. 16, 60, 64
St. 31
Corallophila apiculata (Yamada) R. Norris, 1993: 395.
N’Yeurt 1996: 420, figs 148–151, 163 (‘Centroceras
apiculatum’); Abbott 1999: 288, fig. 81A–C; Payri et al.
2000: 242. Vouchers: WSS 08 St. 5, 67 St. 10
Corallophila huysmansii (Weber-van Bosse) R. Norris,
1993: 396. Abbott 1999: 290, fig. 81D–E; South and Skelton
2002: 148, fig. 11A. Voucher: WSS 127 St. 8
Crouania minutissima Yamada, 1944: 40. Dawson 1956: 55,
fig. 56; Abbott 1999: 294, fig. 82E–G. Vouchers: WSS 33
St. 2, 195 St. 3; UPF 1529, WSS 65 St. 10, 104 St. 9
Comments: the Wallis material has straight to slightly
upcurved whorl branches with spherical to slightly elongate
terminal cells. Cribb (1983) suggested that previous records
of C. minutissima with three whorl-branchlets actually
represent C. capricornica Saenger et Wollaston, as discussed
in Skelton and South (2002: 149). On the other hand, Abbot
(1999: 294) reported having examined authentic
C. minutissima from Ant atoll in SAP, as well as recent
material from that locality, and found three whorl branchlets
in both cases. There hence would seem to be an error in
Yamada’s original diagnosis, and C. minutissima is retained
here as a conservative measure.
Comments: this elegant, plumose material agrees well with
that described by Itono (1977: 139) from Southern Japan.
Haloplegma duperreyi Montagne, 1842b: 258, pl. 7 fig. 1.
Price and Scott 1992: 127, fig. 43A–D; Womersley 1998:
282, fig. 133A–E; Abbott 1999: 305, fig. 86E; Littler and
Littler 2003: 122. Vouchers: UPF 1492, WSS 32 St. 2; UPF
1213 St. 3, WSS 141 St. 4; UPF 1315 St. 10, 1412 St. 22,
1448, 1724 St. 30
Ptilothamnion sp. Voucher: WSS 128 St. 8.
Comments: The material is sterile and specific identification
is not possible.
Spyridia filamentosa (Wulfen) Harvey in W. Hooker, 1833:
337. Womersley and Cartledge 1975: 222, figs 1, 3A–B;
Price and Scott 1992: 131, fig. 45A–E; Womersley 1998:
372, figs 171, 173A–B; Abbott 1999: 313, fig. 88A–B; Payri
et al. 2000: 256; Littler and Littler 2003: 124. Vouchers: UPF
1358, 1359, 1584, WSS 113 St. 8
Comments: frowing as unattached clumps within seagrass
beds.
Wrangelia argus (Montagne) Montagne, 1856: 444. Price
and Scott 1992: 134, fig. 46A–E; N’Yeurt 1996: 424,
figs 167, 169–172. Vouchers: WSS 86 St. 1, 192 St. 3
Wrangelia dumontii (Dawson) Abbott, 1979: 222,
figs 21–26, 29–30; 1999: 316, fig. 89A–B. Littler and Littler
2003: 126. Voucher: WSS 139 St. 4
Comments: Wrangelia dumontii is readily distinguished
from W. argus by its obtuse branch tips, contrasted to the
needle-like tips of the former species.
Dasyaceae
Dasya anastomosans (Weber-van Bosse) M.J. Wynne, 2002:
539. Millar 1990: 433, fig. 59D–G (‘Dasya pilosa’); Abbott
1999: 325, fig. 93A–E (‘Dasya pilosa’); Payri et al. 2000:
264 (‘Dasya pilosa’); Littler and Littler 2003: 128 (‘Dasya
pilosa’). Vouchers: UPF 1665, WSS 173 St. 1
Comments: the tortuous taxonomy of this species is
discussed in Wynne (2002). The Wallis plants have the
typical furry habit with long assimilatory filaments.
Weber-van Bosse (1921: 309, pl. 7 fig. 10) only gave a Latin
diagnosis for her new species, with a fuller description given
in Weber-van Bosse (1923: 376).
384
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Dasya baillouviana (S. Gmelin) Montagne, 1841: 165.
Littler and Littler 1997: 55, fig. 62; 2000: 170, fig. middle
171; Payri et al. 2000: 260. Vouchers: UPF 1534, WSS 71
St. 12
Eupogodon geppii (Weber-van Bosse) P. Silva in P. Silva,
Meñez et Moe, 1987: 130 (misapplied name: Exophyllum
wentii Weber-van Bosse (1928: 478, pl. VI figs 5, 6):
Hollenberg 1968c: 81, figs 6, 7; Abbott 1999: 363,
fig. 105D-H; Payri et al. 2000: 278; Skelton and South
2002a: 155, fig. 16C). Vouchers: UPF 1652, WSS 163, 164,
165 St. 1
Comments: Dasyopsis geppii was first described from the
Chagos Archipelago and Seychelles by Weber-van Bosse
(1913: 130, pl. 13 figs 18–20; pl. 14 fig. 33), together with
Dasyopsis palmatifida Weber-van Bosse (1913: 130, pl. 13
fig. 21). Later, Weber-van Bosse (1923: 379) was of the
opinion that both species could be conspecific, a view
echoed by Dawson (1957b: 124) and confirmed by de De
Jong et al. (1997: 445). Both entities were earlier transferred
to the genus Eupogodon Kützing (1845: 312) by Silva et al.
(1987: 130). More recently, Millar (1996: 156) and De Jong
et al. (1997: 445) alluded to the possibility that Eupogodon
geppi (Weber-van Bosse) P. C. Silva and E. palmatifidus
(Weber-van Bosse) PC Silva formed a genus of their own.
The first author has examined type material of Dasyopsis
palmatifida (L 938.7–422) and it matches well the Wallis
plants, but its affinities to the genus Eupogodon is not
immediately apparent. Some Pacific records were
misidentified in the past as Exophyllum wentii Weber-van
Bosse; the first author has examined the dried Lectotype
(L 0061135) and the liquid-preserved Syntype (L 0109993)
and Isotype (L 0109994) of the latter species, and they do not
match at all the Wallis, Hawaiian, Fijian or Samoan plants in
vegetative or reproductive anatomy. Ongoing molecular
analyses (N’Yeurt and PW Gabrielson, unpubl. data) should
shed more light on the taxonomy and generic affinities of
this unusual plant in the near future.
Heterosiphonia crispella (C. Agardh) Wynne var. laxa
(Børgesen) Wynne, 1985: 87. Price and Scott 1992: 161,
fig. 56A–D; Lluch and Garreta 1993: 468, fig. 4; N’Yeurt
1996: 425, figs 175, 181; Payri et al. 2000: 266; Littler and
Littler 2003: 130; Yamagishi et al. 2003: 540, figs 14–23.
Vouchers: WSS 176 St. 1, 214 St. 23
Delesseriaceae
Hypoglossum simulans Wynne, I. Price et Ballantine, 1989:
31, figs 12–26. Wynne 1989: 515, figs 1H, 2A–E; Abbott
1999: 340, fig. 98A–B; Payri et al. 2000: 258; N’Yeurt 2001:
832, figs 281, 282, 286; Skelton and South 2002a: 152,
fig. 12C–D; Littler and Littler 2003: 134. Voucher: WSS 233
St. 35
Martensia fragilis Harvey, 1854: 145. Millar 1990: 418,
figs 53C–E; Yoshida and Mikami 1996: 101, figs 1, 4–18;
Abbott 1999: 344, fig. 99C–E; Payri et al. 2002: 50, pl. 4
fig. 12; Skelton and South 2002a: 153, fig. 14A–D; Littler
and Littler 2003: 36. Vouchers: WSS 177 St 1; UPF 1209
St. 3; WSS 11, 12 St. 4; UPF 1383, 1569, 1607, WSS 93
St. 16; UPF 1725 St. 23, 1441, 1505 St. 29; WSS 63 St. 31;
UPF 1525 St. 41
Comments: Martensia fragilis can be distinguished from
superficially similar M. elegans by the alternate production
of membranous and reticulate portions of the thallus,
whereas in M. elegans reticulate portions never produce
further membranous parts (Millar 1990). This elegant alga is
quite common in subtidal habitats surveyed in Wallis.
Nitophyllum adhaerens Wynne, 1997: 215, figs 1–14.
Abbott 1999: 347, fig. 100E. Vouchers: WSS 177 St. 1, 191
St. 3; UPF 1576, WSS 103 St. 9; UPF 1668 St 18; WSS 51
St. 29
Rhodomelaceae
Acanthophora pacifica (Setchell) G.T. Kraft, 1979: 128,
figs 1–6, 18–24. Abbott 1999: 353, fig. 102A–C; De Jong et
al. 1999: 229, fig. 43; Payri et al. 2000: 268; Payri et al.
2002: 50, pl. 4 fig. 1; N’Yeurt 2001: 33, figs 293–299; Littler
and Littler 2003: 142. Vouchers: UPF 1420, 1622 St. 22,
1503 St. 29
Acanthophora spicifera (Vahl) Børgesen, 1910: 201,
figs 18A–C; 19A-E. Abbott 1999: 355, fig. 102D–E; De
Jong et al. 1999: 31, figs 3, 33–39, 46; Payri et al. 2000: 270;
Skelton and South 2002a: 153, figs 16A, 17A; Littler and
Littler 2003: 142. Vouchers: UPF 1352, 1353 St. 8
Comments: an often overlooked edible species, growing on
the reef crest and seagrass beds.
Bostrychia tenella (Lamouroux) J. Agardh, 1863 (1851–
1863): 869. Falkenberg 1901: 515, pl. 12 figs 10–13; King
and Puttock 1989: 34, fig. 15A–D; N’Yeurt 1996: 429,
figs 129, 186; Littler and Littler 1997: 59, fig. 68; Payri et al.
2002: 50, pl. 4 fig. 2. Voucher: UPF 1410 St. 38
Comments: found as spongy, sediment-infiltrated mats at the
base of mangrove roots and on beach rock at a marina on the
south coast of the island. This species grows together with
Laurencia brachyclados. It is also found growing on
mangrove roots on nearby Rotuma Island (N’Yeurt 1996).
Chondria arcuata Hollenberg, 1945: 447, figs 2–4.
Gordon-Mills and Womersley 1987: 551, fig 24D–E, 27;
Abbott 1999: 57, fig. 103A; Womersley 2003: 427,
fig. 186A–E. Voucher: WSS 236, St. 35
Chondria armata (Kützing) Okamura, 1907: 69, pl. XVI
figs 9–19. Millar 1990: 459, figs 71A–B; Ballesteros 1994:
538, fig. 1a–c; Payri et al. 2002: 50, pl. 4 fig. 7; Littler and
Marine plants from Wallis
Littler 2003: 144. Vouchers: UPF 1265, 1266, 1301 St. 7,
1542, WSS 79, 80, 81 St. 12
Comments: this distinctive species of Chondria is also found
in neighbouring Fiji (Garbary et al. 1991).
Chondria dangeardii Dawson, 1954: 60, fig. 62f–g. Price
and Scott 1992: 165, fig. 57A–D; Ballesteros 1994: 538
fig. 2a–c; Abbott 1999: 359, fig. 103B–C; Payri et al. 2000:
273; Skelton and South 2002a: 155, fig. 17B. Vouchers: UPF
1718, 1659, WSS 167, 168, 169 St. 1
Chondria dasyphylla (Woodward) C. Agardh, 1817: XVIII.
Gordon-Mills and Womersley 1987: 246, figs 30–57;
N’Yeurt 1996: 430, figs 134, 188; Payri et al. 2000: 74.
Vouchers: UPF 1364, 1589 St. 8
Chondria leptacremon (Melvill) De Toni, 1903: 848. Littler
and Littler 2000: 202. Vouchers: UPF 1188, 1677, 1693,
1700, WSS 184, 185, 193 St. 3; UPF 1649, WSS 160, 161
St. 7; UPF 1406, WSS 225 St. 19
Chondria simpliciuscula Weber-van Bosse, 1913: 125, pl.
12 figs 9, 10. Price and Scott 1992: 169, fig. 59A–E; N’Yeurt
1996: 431, figs 130, 133, 190; Abbott 1999: 361
fig. 104A–F; Payri et al. 2000: 276. Vouchers: WSS 06, 10
St. 5, 60 St. 31
Herposiphonia delicatula Hollenberg, 1968b: 540,
figs 1A,B, 2H, 3. Wynne 1995: 303, fig. 56; Abbott 1999:
371, fig. 107F-G. Voucher: WSS 170 St. 1
Herposiphonia obscura Hollenberg, 1968b: 549, fig. 25.
Abbott 1999: 73, fig. 108B. Voucher: WSS 67 St. 10
Australian Systematic Botany
385
842, figs 309–312, 318); Littler and Littler 2003: 148.
Vouchers: UPF 1662, WSS 172 St. 1, 135 St. 8; UPF 1612
St. 16
Comments: this species can be readily distinguished by its
markedly projecting outer cortical cells.
Chondrophycus succica (Cribb) Nam, 1999: 463. (=
Laurencia succica Cribb 1958a: 163, pl. 1 figs 1–3; Price
and Scott 1992: 192, fig. 69A–D; N’Yeurt 2001: 845,
figs 314, 316, 317; Payri et al. 2000: 280). Vouchers: WSS
175 St. 1; UPF 1678 St. 3, 1594, WSS 118 St. 4; UPF 1467,
WSS 03 St. 5, 119 St. 8; UPF 1386, WSS 239 St. 16; UPF
1667, 1733, 1734, WSS 41 St. 23; UPF 1444, 1507 St. 29
Laurencia brachyclados Pilger, 1920: 6, figs 9, 10. Dawson
1954: 458, fig. 61; McDermid 1988: 232, figs 1, 2; Abbott
1999: 382, fig. 111A–C Vouchers: UPF 1411, WSS 207
St. 38
Comments: growing mixed with Bostrychia tenella at the
base of mangrove roots.
Laurencia cervicornis Harvey, 1853: 73, pl. 18C. Littler and
Littler 2000: 212. Vouchers: UPF 1735, WSS 209 St. 23
Laurencia flexilis Setchell, 1926: 101, pl. 19. Masuda et al.
1999: 455, figs 29–34; Payri et al. 2000: 280; Littler and
Littler 2003: 52. Vouchers: UPF 1629, 1630, WSS 148 St. 7;
UPF 1483 St. 26
Chondrophycus cartilaginea (Yamada) Nam, 2004: 267. (=
Laurencia cartilaginea Yamada 1931: 230, pl. 19 figs a, o;
McDermid 1988: 232, fig. 3; Nam and Saito 1990: 08,
figs 1–7; Abbott 1999: 384, fig. 111D–E). Vouchers: WSS
174, St. 1; UPF 1675, WSS 186 St. 3; UPF 1355, 1356,
1590, WSS 115, 116 St. 8; UPF 1666, 1736, WSS 210 St. 23
Comments: originally described from Tahiti (Setchell 1926),
this species has since been widely reported in the Pacific and
Indian Ocean (Silva et al. 1996). It could be confused with
Laurencia venusta Yamada, but the latter has both
longitudinally oriented secondary pit connection between
superficial cortical cells and lenticular thickenings in
medullary cells, which are not found in L. flexilis (Masuda
et al. 1999).
Comments: Nam and Saito (1999) and Nam (2004) discuss
the Laurencia complex and its relationship to
Chondrophycus.
Laurencia implicata J. Agardh, 1852: 745. Price and Scott
1992: 185, figs 65A–E, 66A–C. Vouchers: UPF 1712, WSS
201 St. 17; UPF 1458, 1515, WSS 56 St. 31
Chondrophycus dotyi (Saito) Nam, 1999: 463. (= Laurencia
dotyi Saito 1969: 154, figs 9, 10; McDermid 1988: 233,
figs 7, 8; Abbott 1999: 385, fig. 112A–B; Payri et al. 2000:
284; Payri et al. 2002: 50, pl. 4 fig. 9). Vouchers: UPF 1190,
1208 St. 3; WSS 143 St. 4; UPF 1270, 1271, 1466, WSS 02
St. 5, 131, 132 St. 8; UPF 1381, 1610, 1619 St. 16, 1427
St. 23, 1504 St. 29, 1524 St. 41
Laurencia mariannensis Yamada, 1931: 200, pl. 5 fig. b,
text figs f, g. Abbott 1999: 388, fig. 113A–B. Vouchers: UPF
1485, WSS 23 St. 4
Comments: Garbary and Harper (1998) delineated the
characteristics that separate the genus Chondrophycus from
Laurencia and Osmundea. Further new combinations were
given by Nam (1999: 463).
Chondrophycus parvipapillata (C. K. Tseng) Garbary et
Harper, 1998: 195. (= Laurencia parvipapillata Tseng 1943:
204, pl. IV; Abbott 1999: 391, fig. 114A–B; N’Yeurt 2001:
Laurencia cf. L. sp. in Price and Scott, 1992: 194,
fig. 70A–C. Vouchers: UPF 1567, WSS 91 St. 16
Comments: this species is similar to the minute Laurencia
reported from the Great Barrier Reef by Price and Scott
(1992), and could well represent a new species. Further
studies are required to elucidate its affinities.
Polysiphonia apiculata Hollenberg, 1968a: 61, figs 1D, 8, 9.
Abbott 1999: 411, fig. 120C. Vouchers: WSS 22 St. 2; UPF
1474, WSS 09 St. 5, 127, 128 St. 8, UPF 1581, WSS 109
St. 9
386
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
Comments: this species is readily distinguished by its
abruptly tapering apices with a prominent apical cell.
Polysiphonia delicatula Hollenberg, 1968a: 62, fig. 1F.
Abbott 1999: 412, fig. 120E. Vouchers: UPF 1687, WSS 190
St. 3, 25 St. 4, 07, 08 St. 5, 60, 64 St. 31
Polysiphonia homoia Setchell et Gardner, 1930: 162. Abbott
1999: 417, fig. 122F–G. Voucher: WSS 67 St. 10
Polysiphonia poko Hollenberg, 1968a: 70, figs 3A, 15. Price
and Scott 1992: 209, fig. 76A–C; Abbott 1999: 421,
fig. 123D-G. Voucher: WSS 233, 237 St. 35
Polysiphonia scopulorum Harvey, 1855: 540. Stegenga and
Vroman 1988: 307, figs 21–26, 29–31; Millar 1990: 445,
fig. 65E–G; Price and Scott 1992: 210, fig. 77A–D; Payri
et al. 2000: 286; N’Yeurt 2001: 37, figs 304, 308; Payri et al.
2000: 286; Womersley 2003: 175, fig. 78A–E. Voucher:
WSS 205 St. 30
Polysiphonia sphaerocarpa Børgesen, 1918: 271,
figs 267–271. Stegenga and Vroman 1988: 307, figs 27, 28;
Millar 1990: 446, fig. 66A–E; Price and Scott 1992: 212,
fig. 78A–D; Schneider and Searles 1997: 204, fig. 19; Abbott
1999: 28, fig. 127A–G. Voucher: WSS 219, 220 St. 10
Comments: this species is easily recognised by its small, erect
tufted habit and large, adaxially curved globular cystocarps
regularly placed on the main axes. Kim and Lee (1999: 280)
transferred P. sphaerocarpa to their new genus Neosiphonia
based on reproductive and vegetative differences with
Polysiphonia, but we concur with Skelton and South (2002:
156) in temporarily maintaining the South Pacific material
under Polysiphonia pending a better understanding and
verification of relevant type material of regional species.
Tolypiocladia glomerulata (C. Agardh) Schmitz in Schmitz
and Falkenberg, 1897: 441. Falkenberg 1901: 177, pl. 21
figs 27–29; Dawson 1954: 452, fig. 59b–c; Price and Scott
1992: 219, fig. 81A–D; Wynne 1995: 321, fig. 80; Abbott
1999: 442, fig. 132F–H; N’Yeurt 2001: 837, figs 302, 303;
Littler and Littler 2003: 158. Vouchers: WSS 157, St. 7; UPF
1730, WSS 208 St. 23
Incertae sedis
Wurdemannia miniata (Sprengel) Feldmann et Hamel,
1934: 544. Hatta and Prud’homme van Reine 1991: 375,
fig. 14a–f; Price and Scott 1992: 31, fig. 6A–E; Abbott
1999: 238. Vouchers: UPF 1556 St. 1, 1599 St. 4; WSS 122
St. 8, UPF 1540, WSS 77 St. 12; UPF 1706 St. 20; WSS 44
St. 23; UPF 1510 St. 29
Magnoliophyta (seagrasses)
Hydrocharitales
Hydrocharitaceae
Halophila ovalis (R. Brown) J.D. Hooker, 1858: 45. Dawson
1954: 376, fig. 1d–f; Meñez et al. 1983: 30, figs 21A–C, 22;
Lanyon 1986: 27, fig. 8, pl. 1 fig. f, pl. 3 fig. f; Payri et al.
2000: 96; Payri et al. 2002: 53, pl. 6 fig. 2; Littler and Littler
2003: 186. Vouchers: UPF 1197, 1198, 1199 St. 3, 1227
St. 6, 1246 St. 7, 1369 St. 8, 1459 St. 34
Comments: the smallest seagrass
characteristic translucent oval leaves.
in
Wallis,
with
Potamogetonales
Cymodoceaceae
Halodule pinifolia (Miki) den Hartog, 1964: 309. Meñez
et al. 1983: 13, figs 7A–B, 8; Lanyon 1986: 20, pl. 1 fig. d,
pl. 3 fig. d; Payri et al. 2002: 53, pl. 6 fig. 3; Littler and
Littler 2003: 288. Voucher: UPF 1371, WSS 245 St. 8
Comments: the Wallis material has irregularly serrate
margins and furcate midribs at the apices of leaves, which are
diagnostic features of the species. The blades are grass-like,
flattened.
Syringodium isoetifolium (Ascherson) Dandy in Dandy and
Tandy, 1939: 116. Meñez et al. 1983: 18, figs 11A–D, 12;
Lanyon 1986: 35, fig. 12, pl. 2 fig. c, pl. 4 fig. c; Payri et al.
2002: 53, pl. 6 fig. 4; Littler and Littler 2003: 288. Vouchers:
UPF 1228, 1229, 1247, 1248 St. 7, 1372 St. 8
Comments: the fleshy, terete blades of this species easily
distinguishes it in the field from Halodule pinifolia.
Discussion
Taxonomy and algal distribution patterns
This study has yielded a total of 194 species of macrophytes,
including 14 Cyanobacteria, 11 Heterokontophyta, 41
Chlorophyta, 128 Rhodophyta and three species of
seagrasses. The non-geniculate Corallinaceae are not fully
treated, and numerous species remain to be added in future.
This study represents the first published records of seagrasses
and marine algae for the Wallis islands, increasing our
knowledge of the tropical marine flora of the western Pacific.
Even though the outer part of the reef has not been sampled
in the eastern part of the island, most of the habitats have been
examined in the other areas from the surface to a depth of
25 m, both inside and outside of the lagoon.
With respect to the distribution of species among the 42
different stations (Table 2), the highest species richness (130
species) is in the area of the fringing reef, owing to the
presence of the large seagrass beds in the sheltered, shallow
and soft-bottomed areas which provide microhabitats for
various algae. The canopy of the seagrass beds comprises the
large Syringodium isoetifolium, and the dominant algae are
common green species such as Halimeda macroloba,
H. incrassata, H. opuntia, Caulerpa spp. as well as the
diminutive seagrass Halophila ovalis. A third species of
seagrass Halodule pinifolia was less common and found at
only one site. The fringing shallow reef flats were dominated
by brown algae such as Padina boryana, Dictyota
Marine plants from Wallis
Australian Systematic Botany
387
Table 3. Taxa comparison between Wallis and several tropical Pacific areas (excluding Cyanobacteria and coralline algae)
Fiji and Rotuma are treated here as a single entity, since all species from Rotuma in common with Wallis also occur in Fiji
Locality
Wallis (this study)
Fiji and Rotuma
(N’Yeurt et al. 1996b; Littler and Littler 2003; South and Skelton 2003)
French Polynesia
(Payri and N’Yeurt 1997; Payri et al. 2000; Littler and Littler 2003)
Solomon Islands
(Womersley and Bailey 1970; Littler and Littler 2003)
Rarotonga, Cook Islands
(Chapman 1977; ADR N’Yeurt unpubl. data)
Samoa (including American Samoa)
(Skelton and South 1999, 2002a, 2002b; Littler and Littler 2003)
bartayresiana, Turbinaria conoides and Sargassum
polycystum. Moroever, 30% of the species sampled in the
fringing reefs were only found in this particular area (e.g.
Acanthophora spicifera, Coelothrix irregularis, Gracilaria
textorii, Hypnea cervicornis, Sarconema filiforme,
Pterocladia caerulescens, Spyridia filamentosa, Hypnea
charoides, Anadyomene tropicus, Siphonocladus tropicus,
Borgesenia forbesii, Chaetomorpha linum, Valonia
aegagropila, Sargassum polycystum and Turbinaria
conoides). The least species rich of all sites are the pinnacles
with only 24 species, none of which is restricted to this zone.
Seventy-one per cent of the pinnacle flora also occur on the
fringing reef, which is undoubtedly due to the very similar
geomorphological features of the reef flats between the two
areas, and the influence of the eroding land mass
(terrigenous inputs leading to sedimentation, water
turbidity). Unlike the barrier reefs, most of the pinnacles are
not under direct oceanic water influence.
The flora of the barrier reef and lagoon reef patches is rich
and similar in terms of species number (115 v. 106 sp.), and
most of the species in these areas have a wide distribution
pattern (18 and 14%, respectively, are restricted to these
areas). However, barrier reefs and lagoon components cannot
be considered as one single area, because the species
composition differs from one area to the next, and only five
species were found in both areas, (e.g. Chlorodesmis
fastigiata, Hydrolithon murakoshii and Sebdenia sp.). The
barrier reef is dominated by non-geniculate coralline algae,
especially mastophoroids (Hydrolithon spp. and
Neogoniolithon spp.), and Lithophylloids (Lithophyllum
insipidium, L. flavescens and the branching L. kotschyanum)
which form large crusts on hard substratum and play an
important role in consolidating the rubbles and large pieces
of corals resulting from destructive dynamite fishing
practices. The shallow and soft-bottom areas of the lagoon
were locally dominated by patches of green algae such as
Halimeda incrassata and Caulerpa spp. (C. brachypus,
C. taxifolia and C. sertularioides), and Cyanobacteria
assemblages, which appear to be seasonal based on
Chlorophyta
Ochrophyta
Rhodophyta
Total
041
136
11
46
111
226
163
408
097
42
146
285
076
30
116
222
044
23
099
166
075
43
155
273
examination of satellite imagery data over several years. The
flora of the coral reef patches in the lagoon area is dominated
by abundant clumps of the common green algae, Halimeda
opuntia and H. minima, as well as turf communities of algae,
mostly red (e.g. Polysiphonia spp., Ceramium spp.). Few
species (five) were restricted to this area: Codium
mamillosum and Dasya spp. Finally, the outer reef slopes
which are less rich than the adjacent barrier reef (90 species),
contain 23% of the species that are not found in other areas;
they are mostly Rhodophyta, such as Acanthophora pacifica,
Chrysymenia okamurae, Platoma cyclocolpum, Predaea
weldii, Prionitis sp., and a few Chlorophyta such as
Rhipiliopsis howensis, Halimeda gracilis and H. hederacea.
Few species are found in both the outer slope and barrier reef
areas; they are mostly Halimeda spp. (H. distorta,
H. taenicola and H. micronesica) and coralline algae.
Overall, the most represented genera are Laurencia (10
species), followed by Halimeda (nine species), Ceramium
(seven species) and Hydrolithon (five species).
The general pattern observed from the distribution of the
Wallis marine flora is characterised by very few species (4%:
e.g. Halimeda opuntia, H. minima, Hydrolithon onkodes,
Valonia fastigiata) shared between the different sites, while
most of them are area-restricted. This agrees with what is
described in French Polynesia by Payri et al. (2000) for the
algae, and also by Galzin and Legendre (1988) for fishes as
a result of biotic (inter specific competition, and grazing)
and abiotic factors (microhabitat, nutrient levels and water
movement, Littler and Littler 1994).
Biogeographic comparisons
Establishing biogeographic comparisons of an area’s flora
requires accurate species taxonomy and also a similar
sampling effort from both the study site and the other regions
selected for the comparison. Thus, in respect of this
assumption, the comparison has been restricted to the
neighbouring regions and those located at the same latitude
range (Table 3). Coralline algae and Cyanobacteria have not
388
A. D. R. N’Yeurt and C. E. Payri
Australian Systematic Botany
been included as they are generally understudied; forms and
varieties of species have been counted as distinct taxa.
The number of taxa (crustose corallines and
Cyanobacteria excluded) of Wallis at first sight appear to be
generally low compared with neighbouring islands such as
Fiji and Samoa. However, this may be a reflection of limited
sampling rather than a true biogeographical difference, as
this study was based on a single 16-day survey in which
some habitats (such as the turbulent eastern region of Wallis)
were excluded. Wallis has no endemic or notable records;
neighbouring Rotuma, on the other hand, has one endemic
taxon (Avrainvillea rotumensis; N’Yeurt et al. 1996a) but this
species has since been reported north of Rotuma, in Tuvalu
(J. Orempuller, pers. comm.).
It is interesting that several very common tropical genera,
such as Hydroclathrus, Rosenvingea, Colpomenia,
Chnoospora, Asteronema, Neomeris and Cheilosporum, are
absent and to note that (excepted for Asteronema,
Chnoospora, Neomeris and Cheilosporum) they do not occur
either north of Fiji such as in Rotuma (N’Yeurt 1996), the
Solomon Islands (Womesley and Bailey 1970), Nauru
(South and Yen 1992) or Kiribati (South et al. 2001). It could
be a seasonal factor, as these species are known to be mainly
present during the warm season (Payri 1987) and collecting
was done in Wallis at the onset of the cooler season. In the
Solomon Islands, Womersley and Bailey (1970: 294) report
another seasonal brown alga, Colpomenia sp., as rare during
the cooler month of August. On the other hand, in isolated
oceanic islands close to equatorial regions such as Rotuma,
Wallis, Nauru, Kiribati there is no marked variation in air and
sea temperatures throughout the year, and these conditions
could exclude the presence of seasonal species whose
distribution range does not reach these higher latitudes. In
the case of habitat-specific species such as the widespread
brown algae Chnoospora minima and usually associated
Asteronema breviarticulatum, their absence from Wallis (but
presence in Rotuma) could be explained by the lack of
exposed, rocky habitas in the former island. However,
seasonality cannot explain the puzzling absence of the green
alga Halimeda discoidea, which is usually ubiquitous in the
tropical Pacific throughout the year, or Cheilosporum spp.
which is usually common in overhangs on the reef crest (both
of these species, incidentally, do occur in neighbouring
Rotuma). An instance of apparently disjunct north–south
biogeographical distribution is the presence of the distinctive
Japanese species Acrochaetium kurogii in Wallis, and also in
the Cook Islands (ADR N’Yeurt, unpubl. data). However,
hasty conclusions should not be made as such minute
epiphytic species are easily overlooked in floristic surveys,
and sampling effort is far from normalised in the region, with
many areas remaining unexplored. It is becoming clear that
an increasing number of species previously considered
endemic or restricted to certain areas are in fact much more
widespread or pantropical, as shown in the recent discovery
of the deepwater red algae Predaea weldii and Gibsmithia
hawaiiensis (type localities in the central Pacific) in South
Africa (De Clerck et al. 2002) and Amansia rhodantha (type
locality in the Indian Ocean) in the South Pacific (N’Yeurt
2002).
Doty (1954) discussed the mutual exclusiveness of the
genera Rhipilia and Sargassum in equatorial localities, such
as Kiribati and Bikini atoll. Wallis island (13°18' S) could be
considered close to the equator, and the presence of Rhipilia
tenaculosa would, following this logic, prevent the
occurrence of Sargassum. Interestingly, a single species of
Sargassum, S. polycystum, occurs at Wallis, albeit only at
one site which is a port of entry. It could be hypothesised that
Sargassum does not occur in the natural state in Wallis, and
its presence is the result of introduction via shipping.
Whatever the case might be, other species are surely to be
discovered in Wallis, especially in the turbulent eastern
region, which is yet to be explored. The present study
provides our first knowledge of the benthic marine
macrophytes associated with the ecosystem diversity of
Wallis. It is the first, but important, step towards the
conservation and management of the habitats of this island’s
marine environment.
Acknowledgments
The authors acknowledge financial support for this study
from the French Coral Reef Reasearch Institute, IFRECOR.
Mr Heroen Verbruggen (Ghent University, Belgium) is
thanked for identifying Halimeda species, as well as for
helping us with collections and sorting of material in the
field, and for being a valuable dive buddy. We thank the
following individuals: Mr Paino Vanai, Head of the
Environmental Service and all his devoted staff, for logistical
support in the field and on land, without which this mission
would have been impossible; the Prefect and Chief
Administrator of Wallis and Futuna; Lavelua, King of
Wallis; the members of the Traditional Council of Chiefs of
Wallis, and Professor C. Chauvet, Université de la
Nouvelle-Calédonie. Dr Serge Andrefouët is thanked for
providing the base map of Wallis, as is Professor Philippe
Morat for kindly providing information on seagrass
taxonomy. An anonymous reviewer is thanked for
constructive criticism and comments on an earlier draft of
this paper that substantially improved its content.
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Yamada Y (1931) Notes on Laurencia, with special reference to the
Japanese species. University of California Publications in Botany
16, 185–310.
Yamada Y (1941) Species of the genus Halimeda from the South Seas.
Kagaku Nanyõ 4, 108–121.
Yamada Y (1944) A list of marine algae from the Atoll of Ant. Scientific
Papers of the Institute of Algological Research, Faculty of Science,
Hokkaido Imperial University 3, 31–45.
Yamada Y, Segawa S (1953) On some new or noteworthy algae from
Hachijo Island. Records of Oceanographic Works in Japan, series 2,
1, 109–114.
Yamagishi Y, Masuda M (1997) Species of Hypnea from Japan. In
‘Taxonomy of economic seaweeds, with reference to some Pacific
species. Vol. 6’. (Ed. A Abbott) pp. 135–162. (California Sea Grant
College System: La Jolla)
Yamagishi Y, Masuda M (2000) A taxonomic revision of a Hypnea
charoides-valentiae complex (Rhodophyta, Gigartinales) in Japan,
with a description of Hypnea flexicaulis sp. nov. Phycological
Research 48, 27–35. doi:10.1046/J.1440-1835.2000.00181.X
Yamagishi Y, Masuda M, Abe T, Uwai S, Kogame K, Kawaguchi S,
Phang SM (2003) Taxonomic notes on marine algae from Malaysia.
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Yamamoto H (1978) Systematic and anatomical study of the genus
Gracilaria in Japan. Memoirs of the Faculty of Fisheries, Hokkaido
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genus Martensia (Delesseriaceae, Rhodophyta), with the
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101–106.
Manuscript received 17 September 2003, accepted 24 June 2004
Taxonomic index
Acanthophora pacifica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acanthophora spicifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acrochaetium barbadense. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acrochaetium kurogii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Actinotrichia fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Amphiroa foliacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Amphiroa fragilissima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Amphiroa tribulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Anotrichium tenue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Antithamnion decipiens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Asparagopsis taxiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Balliella repens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Boergesenia forbesii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Boodlea composita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Bostrychia tenella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Bryopsis pennata var. secunda . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Calothrix confervicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulacanthus ustulatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa biserrulata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa cupressoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa nummularia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa peltata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa racemosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa serrulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa sertularioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa taxifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caulerpa taxifolia ecad. tristichophylla . . . . . . . . . . . . . . . . . . . . .
Centroceras minutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium codii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium flaccidum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium kramerii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium macilentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium sp. aff. C. marshallense . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium subdichotomum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ceramium upolense. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
384
384
378
378
378
379
379
379
382
382
378
382
372
372
384
373
370
380
373
373
373
373
373
374
374
374
374
382
382
382
383
383
383
383
383
Chaetomorpha crassa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chaetomorpha linum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chamaebotrys boergesenii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Champia compressa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Champia parvula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chlorodesmis fastigiata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chlorodesmis hildebrandtii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria arcuata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria armata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria dangeardii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria dasyphylla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria leptacremon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chondria simpliciuscula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chroodactylon ornatum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chronodrophycus cartilaginea . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chrondrophycus dotyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chrondrophycus parvipapillata . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chrondrophycus succica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chrysymenia okamurae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cladophora vagabunda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Codium bulbopilum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Codium mamillosum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Coelothrix irregularis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Corallophila apiculata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Corallophila huysmansii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Corynocystis prostrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Crouania minutissima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cryptonemia umbraticola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cryptonemia yendoi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dasya anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dasya baillouviana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dictyopteris repens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dictyosphaeria cavernosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dictyosphaeria versluysii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dictyota bartayresiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
372
372
382
382
382
375
375
384
384
385
385
385
385
377
385
385
385
385
382
372
373
373
382
383
383
380
383
378
379
383
384
376
372
372
376
Marine plants from Wallis
Australian Systematic Botany
Dictyota divaricata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
Dictyota friabilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Diplothamnion jolyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Erythrotrichia carnea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Eupogodon geppii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Euptilota articulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Galaxaura fasciculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Galaxaura filamentosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Galaxaura marginata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Galaxaura subverticillata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Gelidiella acerosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Gelidiella maschrisiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Gelidiopsis intricata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Gelidiopsis scoparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Gelidium pusillum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Gibsmithia hawaiiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Gracilaria textorii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Gracilaria sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Halichrysis coalescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Halimeda borneensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374
Halimeda distorta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374
Halimeda gracilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda hederacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda incrassata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda macroloba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda micronesica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda minima. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda opuntia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halimeda taenicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
Halodule pinifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Halophila ovalis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Haloplegma duperreyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Halymenia actinophysa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Herposiphonia delicatula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Herposiphonia obscura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Heterosiphonia crispella var. laxa . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Hydrolithon farinosum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Hydrolithon murakoshii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Hydrolithon onkodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Hydrolithon reinboldii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Hydrolithon rupestris. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Hypnea cervicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Hypnea charoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Hypnea pannosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Hypnea saidana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Hypnea spinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Hypnea valentiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Hypoglossum simulans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Jania adhaerens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Kallymenia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Laurencia brachyclados . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Laurencia cervicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Laurencia flexilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Laurencia implicata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Laurencia mariannensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Laurencia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Leptofauchea anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Lithophyllum flavescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Lithophyllum insipidum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Lithophyllum kotschyanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Lithothamnion proliferum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Lithophyllum pygmaeum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Lobophora variegata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Lomentaria corallicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
397
Lyngbya majuscula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370
Lyngbya polychroa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Martensia fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Mastophora pacifica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Mesophyllum erubescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Mesophyllum mesomorphum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Meristotheca procumbens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Neogoniolithon brassica-florida. . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Nitophyllum adhaerens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Nostoc calcicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370
Oscillatoria bonnemaisonii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Oscillatoria margaritifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Padina boryana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Padina melemele. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Peyssonnelia bornetii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Peyssonnelia inamoena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Peyssonnelia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
Phormidium nigroviride . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Phormidium papyraceum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Phyllodictyon anastomosans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Platoma cyclocolpum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Pneophyllum conicum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Polysiphonia apiculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Polysiphonia delicatula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Polysiphonia homoia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Polysiphonia poko . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Polysiphonia scopulorum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Polysiphonia sphaerocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Predaea laciniosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Predaea weldii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Prionitis sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Pterocladiella caerulescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Pterocladiella caloglossoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Ptilothamnion sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Rhipidosiphon javensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
Rhipilia tormentosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
Rhipiliopsis howensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
Sarconema filiforme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Sargassum polycystum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Scytonema polycystum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370
Sebdenia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Siphonocladus tropicus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Sphacelaria tribuloides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
Sphaenosiphon olivaceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
Spirulina subsalsa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Sporolithon ptychoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380
Spyridia filamentosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Stylonema alsidii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Symploca atlantica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Symploca hydnoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Symploca laeteviridis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Syringodium isoetifolium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Tolypiocladia glomerulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Turbinaria conoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Turbinaria ornata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Ulva clathrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Ulva flexuosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Valonia aegagropila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Valonia fastigiata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Valonia macrophysa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Ventricaria ventricosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
Wrangelia argus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Wrangelia dumontii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Wurdemannia miniata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
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