Benthic algal and seagrass communities in Baa Atoll, Maldives

Abstract

The present survey was undertaken to provide the first census of the marine flora (macroalgae and seagrasses) of Baa Atoll, one of the 26 Maldivian atolls, and to serve along with the macro-fauna biodiversity inventories for conservation purposes. Species collection and inventories have been conducted at 27 sites covering the widest selection of habitats recognized based on satellite images including islands shorelines, reef flats, faros, patch reefs, passes as well as shallow and deep outer reef slopes. A total of 405 specimens were collected and 176 species representing 10 Phaeophyceae, 58 Chlorophyta, 108 Rhodophyta and two seagrasses were identified. The lagoon patch reefs and the oceanic reef slopes were the most diverse geomorphological habitat types and displayed the highest species richness with 38 spp. All lagoon sites shown a similar richness compared to each other with an average species number of 26 spp, while the deep lagoon floor and the seagrass beds in oceanic-exposed reef flats were the less species-rich habitats. The most common species, occurring at all visited sites, were Tydemania expeditionis and Halimeda minima and the most species rich genera appeared to be Halimeda and Caulerpa. No community structure nor strongly supported species assemblages associated to geomorphological habitat types was found.
Previous lists available for other Maldivian atolls listed 208 algal species. Sixty of these records were found in Baa Atoll while 113 of the species recorded in the present study represent new records for the Maldives bringing the total number of algal species to 321. The resulting species list shows that the Maldivian algal flora is typically tropical and most of the species belong to the Indo-Pacific biogeographic province. In this paper, we give a general description of the representative macrophyte communities of Baa Atoll in relation to the geomorphology of reefs.

Full text

ATOLL RESEARCH BULLETIN NO. 590
Issued by
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
JANUARY 2012
Edited by
Serge Andréfouët
BIODIVERSITY, RESOURCES, AND CONSERVATION
OF
BAA ATOLL (REPUBLIC OF MALDIVES):
A UNESCO MAN AND BIOSPHERE RESERVE

________________________________________________________
1!"#$%$&$'()'*)+,)-+,)'./&-'0)'123)0/..)4)"$5'67895':/&42;5':)<'=;0)(/"%;>'?4;%0@''
claude.payri@ird.fr
2'7;+%A+'=)"$-)'B/-'?"3%-/"4)"$'C'D&#$;%";E0)'1)3)0/.4)"$'F78=?GD1H5'I,)'J"%3)-#%$K'/B'$,)'D/&$,'
''7;+%A+5'7>L>'6/M'NNOP'5'D&3;5'Q%R%
3'7-)#)"$';((-)##@'J"%3)-#%$K'/B'=;.)'I/<"5'6/$;"K'1).;-$4)"$';"('S;-%")'*)#);-+,'!"#$%$&$)5'=;.)'
Town, South Africa
BENTHIC ALGAL AND SEAGRASS COMMUNITIES IN BAA ATOLL,
MALDIVES
BY
CLAUDE E. PAYRI,1 ANTOINE D.R. N’YEURT,1,2 AND LYDIANE MATTIO1,3
ABSTRACT
I,)'.-)#)"$'#&-3)K'<;#'&"()-$;T)"'$/'.-/3%()'$,)'A-#$'+)"#&#'/B'$,)'4;-%")'
U/-;'F4;+-/;0V;)';"('#);V-;##)#H'/B'6;;'8$/005'/")'/B'$,)'WO'S;0(%3%;"';$/00#5';"('$/'
serve along with the macro-fauna biodiversity inventories for conservation purposes.
Species collection and inventories have been conducted at 27 sites covering the widest
selection of habitats recognized based on satellite images including islands shorelines,
-))B'U;$#5'B;-/#5'.;$+,'-))B#5'.;##)#';#'<)00';#'#,;00/<';"('()).'/&$)-'-))B'#0/.)#>'8'
total of 405 specimens were collected and 176 species representing 10 Phaeophyceae,
9P'=,0/-/.,K$;5'NXP'*,/(/.,K$;';"('$</'#);V-;##)#'<)-)'%()"$%A)(>'I,)'0;V//"'.;$+,'
reefs and the oceanic reef slopes were the most diverse geomorphological habitat types
and displayed the highest species richness with 38 spp. All lagoon sites shown a similar
richness compared to each other with an average species number of 26 spp, while the
()).'0;V//"'U//-';"('$,)'#);V-;##'E)(#'%"'/+);"%+G)M./#)('-))B'U;$#'<)-)'$,)'0)##'
species-rich habitats. The most common species, occurring at all visited sites, were
Tydemania expeditionis and Halimeda minima and the most species rich genera appeared
to be Halimeda and Caulerpa. No community structure nor strongly supported species
assemblages associated to geomorphological habitat types was found.
Previous lists available for other Maldivian atolls listed 208 algal species. Sixty
of these records were found in Baa Atoll while 113 of the species recorded in the present
study represent new records for the Maldives bringing the total number of algal species
$/'YWN>'I,)'-)#&0$%"V'#.)+%)#'0%#$'#,/<#'$,;$'$,)'S;0(%3%;"';0V;0'U/-;'%#'$K.%+;00K'$-/.%+;0'
;"('4/#$'/B'$,)'#.)+%)#'E)0/"V'$/'$,)'!"(/G7;+%A+'E%/V)/V-;.,%+'.-/3%"+)>'!"'$,%#'.;.)-5'
we give a general description of the representative macrophyte communities of Baa Atoll
in relation to the geomorphology of reefs.

32
INTRODUCTION
Z)-K'0%4%$)('%"B/-4;$%/"'%#';3;%0;E0)'/"'S;0(%3)#';-+,%.)0;V/'4;-%")'U/-;'
despite a large number of oceanographic expeditions carried out in the region. Most
/B'$,)'-)+/-(#';-)'E;#)('/"'#./-;(%+'#$&(%)#>'I,)'A-#$'.,K+/0/V%+;0'$;M/"/4%+'-)+/-(#'
were published from the limited biological material collected during the expeditions of
[>D>'\;-(%")-'%"'$,)');-0K'N]XX#'F\;-(%")-5'N]XYH';"('#$&(%)('EK'6;-$/"'FN]XYH5'Q/#0%)'
FN]XY5'N]X^H5';"('_)E)-';"('Q/#0%)'FN]X`H>'I,)'D);0;-T'?M.)(%$%/"'%"'N]X9'-)#&0$)('%"'
\)..';"('\)..a#'FN]X`H';"('_)E)-G3;"'6/##)a#'FN]N`H'-)+/-(#'/B'#/4)'=,0/-/.,K$;'
FbV-))"';0V;)cH';"('*,/(/.,K$;'Fb-)(';0V;)cH>':)<$/"'FN]9YH'.&E0%#,)('/"0K'/")'
#);<))('-)+/-('B-/4'$,)'[>'S&--;K')M.)(%$%/"'N]YYGN]Y`>'8$'$,%#'$%4)'$,)'T"/<0)(V)'
/B'$,)'S;0(%3%;"';0V;0'U/-;'<;#'-)#$-%+$)('$/'W`'#.)+%)#'%"+0&(%"V'N^'*,/(/.,K$;5'A3)'
Chlorophyta and two Pheaophyceae (“brown algae”). During the expedition led by D.R.
D$/((;-$'B-/4'=;4E-%(V)'%"'N]O`5'D%V))'FN]OOH'#;4.0)('0;"(';"('4;-%")'3)V)$;$%/"'
at Addu Atoll. The preliminary results of the Addu Atoll expedition were published
in Atoll Research Bulletin'EK'D$/((;-$'FN]OOH>'I,)'D%V))'+/00)+$%/"'<;#'#$&(%)('EK'
I#&(;';"(':)<,/&#)'FN]OOH'<,/'.&E0%#,)(';'$;M/"/4%+'0%#$5';((%"V'#%V"%A+;"$0K'$/'
the total species number of macroalgae with 37 Rhodophyta, 30 Chlorophyta , nine
Phaeophyceae and seven cyanobacteria (“bluegreen algae”). Extensive collections of
benthic macroalgae were made at nine Maldivian atolls during Cruise B and Cruise 5
of the R/V Te Vega'?M.)(%$%/"#'%"'N]O`'EK'd>?>'d;+T)$$';"('S>[>'_K"")'-)#.)+$%3)0K>'
d;+T)$$'FN]O]H'#$&(%)(',%#'/<"'+/00)+$%/"'B/-',%#'7,1'(%##)-$;$%/"';#'<)00';#'4;$)-%;0'
+/00)+$)('EK'=>'*,K")'%"'N]O^';$'8((&'8$/00'(&-%"V'$,)'J>D>':;3K'6%/0/V%+;0'?M.)(%$%/"'
to the Chagos Archipelago. Based on these collections additional records were published
%"'d/00)"E)-Va#'FN]OP;5'EH'4/"/V-;.,#';"('8-)V//(';"('d;+T)$$'FN]^NH'()#+-%E)(';'
new species of Rhodophyta (Dictyurus maldiviensis'd;+T)$$'C'8-)V//(H>'e;$)-5'd;+T)$$'
FN]^^H'.&E0%#,)('$,)'4/#$'+/4.-),)"#%3)'+;$;0/V&)'/B'$,)'S;0(%3%;"'4;-%")';0V;)'
with 248 records including 136 Rhodophyta, 74 Chlorophyta, 17 Phaeophyceae and 21
=K;"/E;+$)-%;>'8'"&4E)-'/B'$,)#)'-)+/-(#',/<)3)-'<)-)'%()"$%A)('/"0K';$'$,)'V)"&#'
0)3)0>'8'K);-'0;$)-5'I%$0K;"/3;';"('6&$/-%"'FN]^PH'.&E0%#,)(';'#,/-$'0%#$'/B'4;+-/;0V;)'FNP'
$;M;H'B-/4'$</';$/00#'/B'$,)'S;0(%3)#>'Q%";00K'_K"")'FN]]YH'.&E0%#,)(';'0%#$'/B'9X'#.)+%)#'
based on his own collections including the description of Bangia halymeniae Wynne
from Malé Atoll.
!"'WXX]5'E)B/-)'$,)'6;;')M.)(%$%/"5'$,)'4/#$'&.(;$)('+/4.%0;$%/"'/B'$,)'4;-%")'
U/-;'/B'$,)'S;0(%3)#'<;#';3;%0;E0)'B-/4'b;0V;)E;#)>+/4c';"('0%#$)('NWX'*,/(/.,K$;'5'^X'
Chlorophyta, 18 Phaeophyceae and 21 Cyanobacteria records (Guiry and Guiry, 2011).
I,)'.-)#)"$'#&-3)K'<;#'&"()-$;T)"'$/'.-/3%()'$,)'A-#$'+)"#&#'/B'$,)'4;-%")'
U/-;'/B'6;;'8$/00';"('$/'#)-3)';0/"V'<%$,'$,)'4;+-/GB;&";'E%/(%3)-#%$K'%"3)"$/-%)#'B/-'
+/"#)-3;$%/"';"('%()"$%A+;$%/"'/B'E%/(%3)-#%$K',/$G#./$#'Fd;4)0';"('8"(-2B/&f$5'$,%#'
issue). We also provide here a general description of the representative macrophyte
communities of Baa Atoll in association with the geomorphology of reefs.

33
SAMPLING SITES AND METHODS
Baa Atoll is situated in the Northern Indian Ocean at latitude 5°11'N and longitude
^Wg9]h?>'6;;'%#'/")'/B'$,)'WO'S;0(%3%;"';$/00#'#$-)$+,%"V'%"';'"/-$,G#/&$,'(%-)+$%/"'/BB'
India’s Lakshadweep islands. It stands in the Laccadive Sea, about 700 km south-west
of Sri Lanka and 400 km south-west of India. Baa Atoll is 42 km long and 32 km wide.
I,)'$-/.%+;0'+0%4;$)'%#'+/4./#)('/B'$</'4;%"'#);#/"#@'$,)'(-K'#);#/"';##/+%;$)('<%$,'$,)'
winter north-eastern monsoon and the rainy season with strong winds and storms.
I,)'.-)#)"$';0V;0'U/-;';"('#);V-;##'%"3)#$%V;$%/"'/B'6;;'8$/00'<;#';+,%)3)('
(&-%"V'S;K';"('[&")'WXX]'R&#$';B$)-'$,)'4/%#$'#/&$,G<)#$'4/"#//">'D&-3)K#'<)-)'
conducted at 27 sites (Fig. 1) covering the widest selection of habitats recognized based
/"'#;$)00%$)'%4;V)#'%"+0&(%"V'%#0;"(#'#,/-)0%")#5'-))B'U;$#5'B;-/#5'.;$+,'-))B#5'.;##)#'
as well as shallow and deep outer reef slopes. Most of the sites were prospected by
D=J68'B-/4'9X'4'$/'$,)'#&-B;+)>'I,)'#,;00/<';-);#'%"+0&(%"V'B-%"V%"V'-))B'U;$#5'.;$+,'
reefs and shorelines were sampled by snorkelling or reef walk. The sampling effort was
standardized and inventory duration at each site was set to 80 min.
All specimens collected were sorted, pressed and air-dried as herbarium vouchers.
Photographs of collected specimens were taken in-situ and referenced according to
herbarium accessions. Samples of selected specimens were pickled in a solution of
buffered formalin in seawater (5%) for further anatomical studies. Samples from a
selection of taxa were preserved in silicagel or ethanol for further DNA analyses. Since
all herbarium specimens were air dried (no formalin), DNA extraction is feasible for
further studies if necessary. DNA samples of Dictyotales (Dictyota J.V. Lamouroux and
Padina Adanson) and Halimeda J.V. Lamouroux have already been processed and will be
included in regional phylogenetic studies.
Overall, specimens were collected to represent a baseline taxonomical collection
for the area and the species inventory was compiled in order to reach the more
comprehensive species list for Baa Atoll. In agreement with the Maldive Research Center
(MRC), the collection was deposited in the phycological herbarium of IRD (Institut de
Recherche pour le Développement) in Nouméa (IRD-NOU), New Caledonia.
RESULTS
Representative Algal Communities and Associated Habitats
D)3)"'+0;##',;E%$;$#',;3)'E))"'()A")('E;#)('/"'V)/4/-.,/0/VK';"('4/#$'/B'$,)4'
were prospected. All habitats could not be sampled with the same effort and some of
$,)45'#&+,';#'#);V-;##'E)(#'/-'/+);"%+'-))B'U;$#5'<)-)'/"0K'3%#%$)('/"+)'FI;E0)'NH>'
During the present investigation 405 specimens were collected from 27 sites
FQ%V>'NH>'8'$/$;0'/B'N^O'#.)+%)#'<)-)'%()"$%A)(';"('-).-)#)"$)('NX'7,;)/.,K+);)5'9P'
=,0/-/.,K$;5'NXP'*,/(/.,K$;'i:6@'/"0K'$,)'4/#$'+/44/"'-)('+/-;00%")#';0V;)'<)-)'
+/"#%()-)(j';"('W'#);V-;##)#>'I,)'$;M/"/4%+'+0;##%A+;$%/"'&#)('(&-%"V'$,%#'</-T'
followed
The catalogue of the benthic marine algae of the Indian Ocean by Silva and
+/G;&$,/-#'FN]]OH>
The species list is given in Appendix 1. Records belong to 17 orders,
Y9'B;4%0%)#';"(']`'V)")-;'FI;E0)'WH>

34
Class
Habitat
Oceanic
reef flat
(seagrass)
Lagoon
reef flat
and slope
Lagoon
patch reef
Lagoon
reef flat
Deep
lagoon
Oceanic
reef flat
Oceanic
reef slope
number
of sites
(1)
(10)
(5)
(3)
(1)
(2)
(5)
Site
label
2
1, 5, 6, 8,
12, 13, 21,
22, 24, 28
10, 11,
N^5'N]5'W9
3, 16, 20
23
N95']
4, 7, 14,
18, 27
Figure 1. Location of the sampling sites in Baa Atoll.
I;E0)'N>'D;4.0%"V'#%$)#'(%#$-%E&$%/"'%"'$,)'#)3)"',;E%$;$'+0;##)#'()A")('E;#)('/"'
geomorphology.

35
S/#$'/B'$,)'#.)+%4)"#',;3)'E))"'%()"$%A)('$/'#.)+%)#'0)3)0'FP9'k'/B'$,)'
+/00)+$%/"H5'E&$'#/4)'/B'$,)'#.)+%4)"#'FN9'k'/B'$,)'+/00)+$%/"H'-)4;%"'&"%()"$%A)('B/-'
lack of reproductive parts or poor sampling. In addition the two seagrasses Syringodium
isoetifolium (Ascherson) Dandy and Thalassia hemprechii (Ehrenberg) Ascherson have
been observed forming beds in only a single location (Baa2, Fig.1).
The most species rich genera appeared to be the green algae Halimeda and
Caulerpa J.V. Lamouroux, however surprisingly, no bulbose Halimeda spp. were
recorded during this survey nor the large Fucales such as Sargassum C. Agardh. All the
#.)+%)#'#;4.0)('(&-%"V'$,)')M.)(%$%/"'<)-)';##/+%;$)('$/',;-('#&E#$-;$&4')M+).$'/")@'
Boodleopsis pusilla FQ>D>'=/00%"#H'_>*>'I;K0/-5'8>6>'[/0K'C'6)-";$/<%+l'<,%+,'/++&--)('
on faros’ sandy bottoms. (Appendix 2).
Species Richness Distribution in Baa Atoll.
Species richness per site. Species richness per site (Fig. 2) ranged from two at
Baa 23 (deep lagoon) to 38 species at Baa 11 (lagoon patch reef) and Baa 7 (oceanic reef
slope) (Fig. 1). These two sites, coral built and exposed to strong water movements, were
the most diverse and displayed the highest species richness with 38 spp.
I;E0)'W>':&4E)-'/B'L-()-#5'Q;4%0%)#5'\)")-;';"('D.)+%)#'/B'4;+-/.,K$)#'%()"$%A)('B-/4'
Baa Atoll
Orders
Families
Genera
Species
Rhodophyta
10
22
64
108
Macroalgae
Chlorophyta
5
11
25
58
Phaeophyceae
2
2
5
10
TOTAL
17
35
]`
176
Seagrasses
Magnoliophyta
1
2
2
2
Figure 2. Distribution of species richness per sites.

36
Because only two species have been collected from the sandy bottom of the
()).)#$'0;V//"'#%$)'F6;;'WY5'9X'4'()).H';"('$,;$'6;;'WX'F0;V//"'-))B'U;$H'<;#'"/$'
properly prospected (no SCUBA on the deeper part), we believe that the data are not
reliable enough to report on the species richness of corresponding habitats. The seagrass
bed (Baa 2) housed few algal species (four spp, including two coralline rhodoliths).
I,)'#;"(K',;E%$;$#'F6;;']5'NY5'NOH';"('$,)'-))B'U;$#'F6;;'P5'NW5'N9H'<%$,'0/<)-'E%/$/.)'
(%3)-#%$K'/-')M./#)('$/'3)-K'#$-/"V'+&--)"$'F6;;'N^5'N]H'#,/<)('4/()-;$)'#.)+%)#'
richness (from 11to 24 spp). For all the other sites algal richness varied between 25 and
35 species (Fig. 3).
Species richness per geographic areas. The main features of Maldivian reef
+/4.0)M'+;"'E)'+0;##%A)('%"$/'$</'4;R/-'+0;##)#@'F%H'$,)'A-#$'+0;##'%"+0&()#'$,)';$/00'-%4'
4;()'/B'B;-/#'<%$,')"+0/#)('0;V//"#'()0%4%$)('EK'0;-V)'#);<;-('-))B'U;$#5'()).'/&$)-'
reef slopes, forereefs, inner slopes and channels/passes; (ii) the second class includes
$,)'0;V//"'#$-&+$&-)#'<%$,'4/#$0K'B;-/#';"(')"+0/#)('0;V//"#5'-))B'U;$#5'-))B'.;$+,)#5'
pinnacles and deep lagoon sections.
No contrasting spatial variation could be observed between the different sections
of the atoll rim (Table 3). The sampling effort in each geographic area was not strictly
similar and could partially affect the resulting values. The average species richness did
not contrast strongly from North (21 spp.) to South (28 spp.) and from East (26.2 spp.)
to West (27 spp.). Nevertheless, the lowest species richness was observed in the northern
rim section. No contrasting difference could be found between the average species
richness of the global atoll rim (25.2 spp.) and the lagoonal faros (24.7 spp.).
Figure 3.Spatial distribution of the species richness in Baa Atoll.

37
Species richness per geomorphological habitat type. Average species richness
varied within each geomorphological habitat type (Fig. 4). Figure 4 shows that the deep
0;V//"';"('$,)'#);V-;##'E)(#'%"'/+);"%+'-))B'U;$#'<)-)'$,)'0);#$'#.)+%)#'-%+,',;E%$;$#5'
while these sites were also the least sampled (n=1 site for each of them). All lagoon sites
showed a similar richness compared to each other with an average species number of 26
spp. but different sampling efforts (n= 3'#%$)#'B/-'$,)'m0;V//"'-))B'U;$a5'n=10 sites for the
m0;V//"'-))B'U;$';"('#0/.)a';"('n= 5 sites for the ‘lagoon patch reef’). The oceanic slope
showed an average of 30 spp. for the 5 sites.
The spatial distribution of the species varied with bathymetry and some
species were observed at a wide range of depths. This is true for instance for the green
algae Caulerpa diligulata, Halimeda discoidea, Rhipidosiphon javensis, Tydemania
expeditionis and the red Botryocladia which were found from 5 m down to 40 m,
independently of geographical location. Other species were found to be restricted to deep
l/")#5'#&+,';#@'Padina okinawensis, Cladophora feredayoides, Microdictyon okamurae,
Caulerpa sedoides, or cryptic and only in shaded areas such as Cryptonemia umbraticola
or Corynocystis prostrata. Many other species were restricted to shallow waters such as
Valonia aegagropila, Dictyurus purpurascens, Halymenia actinophysa, Hypnea spp. and
Turbinaria ornata.
Table 3. Species richness per geographic area (sites 2 and 20 were excluded because not
properly prospected)
Geographic Area
Number of sites
Northern
Rim
(4)
Southern
Rim
(3)
Eastern
Rim
(5)
Western
Rim
(2)
Atoll Rim
(14)
Atoll
Lagoon
(8)
Site label (BaaX)
14, 15,
16, 17
1, 3, 4
^5'P5']5
NP5'N]
27, 28
1, 3, 4, 7,8,
]5'N`5'N95
16, 17, 18,
N]5 27, 28
5, 6,10,
11,12,13,
21, 22, 23,
24, 25
Average species
21
28
26.2
27
25.2
24.7
Figure 4. Variation of species richness per habitat type.

38
Species rarity. Patterns of the algal vegetation in the Baa Atoll were characterized
by the dominance of species with very low occurrences at all the prospected sites (Fig. 5).
Half of the species (n=87) were found in less than 8 % (2/27) of the sites thus revealing
;',%V,'E)$;G(%3)-#%$K>':/'#.)+%)#'<)-)'.-)#)"$';$');+,'/B'$,)'W^'#%$)#>'I,)'4/#$'B-)n&)"$'
species were the Chlorophyta Tydemania expeditionis Weber-van Bosse occurring in
20/27 sites and Halimeda minima F_>*>I;K0/-H'd%00%#G=/0%"3;&MH'.-)#)"$'%"'N]oW^'#%$)#>'
Ninety percent of the species occurred in only 13 sites which represents less than half of
the prospected sites. Less than 10% of the species were represented by a single specimen.
Main Algal Assemblages.
The most common species are illustrated in Appendix 3.
!"#$"%&'#'($()*+%*+%"$#**+%,''-%.$(/%$+0%/"*1'/2%The algal communities of the
e;V//"'-))B'U;$#';"('#0/.)#';++/&"$'B/-';'0;-V)'"&4E)-'/B')"+-&#$%"V'+/-;00%")';0V;)>'
They are mostly represented either by Hydrolithon onkodes which develops thick crusts
and a candle-like Hydrolithon sp., or branched clumps of Lithophyllum kotschyanum
4%M)('<%$,'#)3)-;0'U)#,K'#.)+%)#'V-/<%"V'%"'+/-;0'+-)3%+)#>'L"'$,)'-))B'U;$#5'4;"K'
species (mostly red algae) grow under the branches of corals. They include the large
spreading mats of Dictyurus purpurascens, Hypnea pannosa and Hypnea spinella,
isolated clumps of 3$"$4$5,$%6"$7'+(*/$, Actinotrichia fragilis, or the delicate and
frondose Halymenia durvillei. The green algae were well represented with Tydemania
expeditionis and several Halimeda spp. including the very common and abundant
H. minima, H. opuntia and H. gracilis (with tiny segments) while the larger species
H. distorta and H. discoidea were common features of the reef slope’s deeper parts.
Figure 5. Histogram of species occurrences at the 27 sites.

Y]
Some branched Codium geppiorum were also observed. Dead corals were colonized
by turfs of Gelidiopsis intricata, Champia vieillardii and Caulerpa nummularia. The
vegetation on the slopes was scarce, less abundant and dominated by calcareous species
such as Lithothamnium proliferum and Halimeda gracilis. Fleshy algae were less
abundant and mostly represented by Gibsmitha hawaiiensis, G. dotyii, and Botryocladia
skottsbergii,Chamaebotrys boergesenii and Portieria hornemanii. Various thin and
small fronds of the dark green algae Rhipiliella spp. and Rhipiliopsis spp. formed small
associations in the shady areas with Corynocystis prostrata Kaft and Cryptonemia
umbraticola. The Caulerpa spp. were poorly represented in these environments and
Phaeophyceae were mainly represented by small Dictyotales such as Dictyota friabilis
and Dictyopteris repens. The large Fucales Turbinaria ornata was very rarely found and
represented only by juveniles while no Sargassum species were observed.
!"#$"%&'#'($()*+%*+%"$#**+%,''-%.$(/2'I,)'#,;00/<'-))B'U;$#'%"'$,)'0;V//"';..);-)('
heterogeneous and some of them in the south-west of the atoll (Baa 3, Fig. 1) showed
much more species richness and biomass than those located in the north-eastern section
of the atoll. H. micronesica and H. taenicola were observed only in the north (Baa 20).
I,)'3)V)$;$%/"';##)4E0;V)'<;#'#%4%0;-'$/'$,;$'/B'$,)'0;V//"'-))B'U;$#'<%$,'"&4)-/&#'
encrusted corallines including Hydrolithon onkodes, the candle-like Hydrolithon sp. and
numerous rhodoliths of Hydrolithon reinboldii. Various articulated coralline species such
as Amphiroa spp. formed clumps on the reef top.
Thin and delicate Rhodophyta such as Hypoglossum spp., Nitophyllum spp. and
several Laurencia spp., were observed in the crevices of hard substratum along with
the very abundant green fan-like Rhipidosiphon javensis, the bright green Anadyomene
wrigthii and Rhipiliella verticillata. The typical fan-like Lobophora variegata and several
large Dictyota spp. were found growing on dead corals along with the green sponge-like
Boodlea composita and the plumose dark green Bryopsis pennata.
Algal vegetation in lagoon patch reefs. The species assemblages and richness
observed in lagoon patch reefs were relatively variable from one site to another with an
;3)-;V)'/B'W9'#..>'S/-)'$,;"',;0B'/B'$,)'#.)+%)#'<)-)'.-)#)"$';$'0);#$'$,-))'/&$'/B'A3)'
sites visited in this same geomorphologic habitat type. The assemblage was dominated by
large green Tydemania expeditionis, as well as Halimeda minima and H. opuntia with a
lesser abundance of by H. cuneata and H. gracilis, the bright green pompom-like Chlo-
rodesmis fastigiata, the dark green Avrainvillea lacerata and Asteromenia anastomosans.
Caulerpa diligulata /++&--)(';$';00'#%$)#'+0;##%A)('%"'$,%#'V)/4/-.,/0/V%+;0',;E%$;$'$K.)>'
Most of the investigated sites showed turf assemblages associated with dead coral includ-
ing mostly Gelidiopsis intricata, Dictyota humifusa, and Champia compressa. Several
species such as !8$+(9*19*,$%1$8)68$, Caulerpa diligulata, Cladophoropsis vaucheri-
aeformis, C. herpestica were also common component of the oceanic reef slopes assem-
blages which are described hereafter.
Algal vegetation of the oceanic reef slope. The species assemblages associated
<%$,'$,)'/&$)-'-))B'#0/.)';..);-)('$/'E)'$,)'4/#$'(%3)-#)';"('-%+,5')3)"'%B'$,)'U)#,K'#.)-
+%)#'<)-)'"/$'3)-K'(%3)-#%A)(>'8E/&$'YX'#.)+%)#'<)-)'/E#)-3)('%"'4/#$'/B'$,)'#%$)#>'I,)'

40
vegetation was dominated by coralline species especially on the upper part of the slope.
Some sections of the outer reef slope were very steep or vertical walls with numerous
crevices, overhangs and small caves. Coral walls were encrusted by coralline species and
Peyssonnelia spp. picturing an attractive mosaic of forms and colors. Lithothamnion pro-
liferum was easily recognizable thanks to its pink crust and numerous short knobs. Along
the slope several Rhodymeniales including Leptofauchea spp. and Rhodymenia spp. oc-
curred in caves and crevices as well as Cryptonemia umbraticola, Corynocystis prostrata,
the iridescent Halichrysis irregularis and the star–shaped Asteromenia anastomosans.
Numerous small green species such as Phyllodictyon anastomosans, Rhipidosiphon ja-
vensis and Rhipilia crassa were present in the crevices while Cladophora feredayoides
and Caulerpa sedoides were collected from rubbles. Conversely the large Gibsmithsia
hawaiiensis and G. dotyii as well as the delicate Kallymenia thompsonii, Dasya anasto-
mosans and D. baillouviana remained scarce. The most obvious species were the green
Halimeda spp. (H. gracilis, H. minima and H. cuneata) and Tydemania expeditionis.
Apart from Caulerpa diligulata, which was relatively abundant, the other species of
Caulerpa (:2%6")8*)0'/ and C. sedoides) were very inconspicuous.
!"#$"%&'#'($()*+%*-%(9'%*8'$+)8%,''-%.$(2%L+);"%+'-))B'U;$#'<)-)'"/$'#&-3)K)(';#'
B-)n&)"$0K';#'$,)';E/3)'()#+-%E)(',;E%$;$#';"(';-)'-).-)#)"$)('%"'$,%#'#$&(K'EK'/"0K'$</'
sites. The species richness was similar from one site to the other with an average of 12
species. Species assemblages however differed strongly. Only two species were common
$/'E/$,'#%$)#@'$,)'3)-K'<%()#.-);('Halimeda opuntia ;"('$,)'_)#$'7;+%A+'Padina okinaw-
ensis. Considering the low sampling effort applied to this geomorphological habitat type
(&-%"V'$,)'#&-3)K5'"/'()A"%$%3)'B);$&-)#'+;"'E)'()#+-%E)(',)-)>'
Algal vegetation associated to seagrass beds. D);V-;##)#';-)'U/<)-%"V'.0;"$#'E)-
longing to the Cymodoceaceae and Hydrocharitaceae families which are currently classi-
A)('%"'$,)'/-()-'80%#4;$;0)#'F"/4)"+0;$&-)'E;#)('/"'.,K0/V)")$%+'#$&(%)#'87\!!!5'WXX]H>'
In tropical regions, they are almost permanently immersed in sheltered marine and estua-
rine biotopes which offer a suitable substrate for rooting in mud, sand or coarse rubble. In
some instances they may also develop into large meadows or beds in deeper lagoon parts
down to 40 m deep, or on barrier reefs surrounding lagoon islands. They are remarkable
habitats in tropical shallow waters and they often represent keystone ecosystems on sandy
bottoms and along shorelines between mangroves and coral reefs.
In Baa, only one site showing typical seagrass habitat was surveyed (oceanic
-))B'U;$5'6;;'WH>'I,)'#);V-;##'#.)+%)#'(%3)-#%$K'<;#'n&%$)'0/<'<%$,'/"0K'$</'#.)+%)#@'
Syringodium isoetifolium and Thalassia hemprechii, forming a dense bed in an area
exposed to strong currents. The algal vegetation associated to this meadow was very poor
with only four large species including Halimeda opuntia, Valonia aegagropila and two
-,/(/0%$,GB/-4%"V'+/-;00%")';0V;)@'Neogoniolithon frutescens and N. laccadivicum.
More prospection is needed to assess the status of seagrass beds in Baa Atoll.

41
DISCUSSION
S;-%")'S;+-/.,K$)#'%"'6;;@'\)")-;0'!"#%V,$#>
8'$/$;0'/B'N^`'4;+-/;0V;0'#.)+%)#'<)-)'%()"$%A)('B-/4'$,)'#&-3)K'/B'6;;'
Atoll. This result does not include the full diversity of coralline algae especially for
the encrusting forms which were not fully sampled in the present study. This group
%#'$;M/"/4%+;00K'(%BA+&0$';"(';'4/-)'+/4.-),)"#%3)'%"3)"$/-K'%#'"))()('$/'.-/.)-0K'
describe its diversity in Baa. Similarly, microscopic epiphytes and epilithic species
have not been exhaustively sampled and studied. A more focused study would most
probably reveal a higher diversity. Nevertheless, our results document and acknowledge
$,)')+/0/V%+;0'-;-%$K'$K.%+;0'%"'$-/.%+;0')+/#K#$)4';#'<)00';#'+/"A-4'.-)3%/&#'#$&(%)#'
conducted in coral reef environments on biodiversity of molluscs and crustaceans
(Bouchet et al., 2002). Overall, and in the framework of the Baa expedition, taxonomic
-)#&0$#5'#.)+%)#'(%#$-%E&$%/"';"('/++&--)"+)#'/E$;%")('B/-'$,)'4;-%")'U/-;';-)'#%4%0;-'$/'
those obtained for the other marine groups studied during this expedition (cf. this issue of
Atoll Research Bulletin).
We carried out a multivariate analysis based on species absence/presence within
$,)'W^'#$&(%)('#%$)#'F-)#&0$#';3;%0;E0)'&./"'-)n&)#$'$/'$,)'A-#$';&$,/-H>'*)#&0$#'#,/<)('
"/'+/44&"%$K'#$-&+$&-)'/-'#$-/"V'%"(%+;$%/"'/B'#.)+%A+'#.)+%)#';##)4E0;V)#';##/+%;$)('
to geomorphological habitat type. This relative homogeneity could be explained by
limited habitat diversity. From its geographical location, Baa atoll appears greatly
%"U&)"+)('EK'#,%B$%"V'4/"#//";0'/+);"%+'+/"(%$%/"#>'I,%#'+/&0('V)")-;$)',/4/V)")/&#'
environmental forcing thus limiting habitat diversity and in turn leading to a more or
0)##',/4/V)")/&#0K'(%#$-%E&$)('U/-;';$'$,)';$/00'#+;0)>'8$'-))B'#+;0)')"3%-/"4)"$;0'
factors are not strictly homogeneous and benthic community assemblages may show
spatial heterogeneity (Vroom et al. 2005) which could be the case in Baa. Here, no
#%V"%A+;"$'(%BB)-)"+)'%"'$,)'#.)+%)#'-%+,")##'/B'$,)'4;-%")'U/-;',;#'E))"'#,/<"';4/"V'
the different areas of the atoll, however the number of restricted species was much higher
$,;$'$,)'"&4E)-'/B'#.)+%)#'<%()0K'(%#$-%E&$)(>'I,%#'-)#&0$'n&)#$%/"#'<,)$,)-'4;+-/;0V;0'
communities within a same geomorphological area are ecologically similar.
=K+0/")#';"('E0);+,%"V')3)"$#';-)'#%V"%A+;"$'(%#$&-E;"+)#'-)#&0$%"V'./$)"$%;00K'%"'
;'#,%B$'B-/4'+/-;0'(/4%";$)('$/'4;+-/;0V;)'(/4%";$)('-))B#'FS+=//T5'N]]]p'6)00<//('
et al., 2006). However no evidence of algal dominated communities was observed during
$,)')M.)(%$%/"5'+/"(&+$)('NN'K);-#';B$)-'$,)'4;##%3)'N]]P'E0);+,%"V')3)"$'$,;$'%4.;+$)('
Maldives.
Previous lists available for other Maldivian atolls (Guiry and Guiry, 2011) listed
208 algal species. Sixty three of these records were found in Baa Atoll. Conversely, 113
of the species recorded in the present study represent new records for the Maldives,
E-%"V%"V'$,)'$/$;0'"&4E)-'/B';0V;0'#.)+%)#'$/'YWN'FWXX'*,/(/.,K$;5']^'=,0/-/.,K$;';"('
24 Phaeophyceae). Comparison with previous studies undertaken in the Maldives show a
narrow overlap of the diversity of the species between the different atolls studied.
As observed from the literature and from this study, the Maldivian macroalgal diversity
varies from one atoll to another and several very common tropical species have not been

42
recorded during the present survey. Some of the species are seasonal (e.g. Rosenvingea
intricata) and did not occur in May-June at the time of the survey. Another likely
hypothesis is that Baa Atoll does not offer the suitable habitats that support those
particular species.
Biogeography
' I,)'#.)+%)#'0%#$')#$;E0%#,)('B-/4'$,%#'#&-3)K'#,/<#'$,;$'$,)'S;0(%3%;"';0V;0'U/-;'
%#'$K.%+;00K'$-/.%+;0';"('4/#$'/B'$,)'#.)+%)#'E)0/"V'$/'$,)'!"(/G7;+%A+'E%/V)/V-;.,%+'
.-/3%"+)>'D)3)-;0'#.)+%)#'()#+-%E)('B-/4'$,)'7;+%A+'-)V%/"'<)-)'-)+/-()('B/-'6;;'(&-%"V'
$,%#'#$&(K';"('-).-)#)"$'$,)%-'A-#$'-)+/-('B/-'$,)'!"(%;"'L+);">'I,)K';-)'B/-')M;4.0)'
the Dictyotales Padina okinawensis described from Southern Japan, the Delesseriaceae
Myriogramme heterostroma and M. melanesiensis originally described from the Solomon
!#0;"(#';"('Z;"&;$&'F_)#$)-"'7;+%A+H';"('$,)'d;0%4)(;+);)'Halimeda xishaensis from
China (Gulf of Tonkin). This suggests that the species geographic distribution is broader
$,;"'/-%V%";00K'$,/&V,$';"('&"()-0%")#'$,)'E%/V)/V-;.,%+';BA"%$%)#'/B'$,)'S;0(%3)#'4;-%")'
U/-;'<%$,'$,)'$-/.%+;0'_)#$'7;+%A+>
=/4.;-%#/"'<%$,'U/-;#'B-/4';(R;+)"$'-)V%/"#'%#'0%4%$)('(&)'$/'(%BB)-)"+)'%"'
sampling effort and lack of recently revised species lists. However we compared different
archipelogoes from the West Indian Ocean based on species lists available at algaebase.
com (Table 4). The proportion of species shared by Baa and other atolls/islands of the
Maldives, Laccadives, Chagos, Seychelles and La Reunion was 35.7, 25.5, 17.6, 43.1 and
W]>9k'-)#.)+$%3)0K>'I,)',%V,)#$'.)-+)"$;V)'#%4%0;-%$K';..);-)('$/'E)'<%$,'$,)'D)K+,)00)#'
Islands and the other Maldivian atolls. The lowest similarity was observed with the
Laccadives (10.57° N and 72.62° E) and Chagos (6° S and 72° E). The reason for such a
low similarity despite the geographical location of these Islands (Chagos and Laccadives
are located about 600 km off the south and about 250 km off the north of the Maldives,
respectively), could be explained by low collecting efforts at these localities resulting in
incomplete species lists. A Sorensen’s Similarity Index was calculated between the Baa
4;-%")'U/-;'+/4./#%$%/"';"('$,/#)'/B'$,)'/$,)-'0/+;0%$%)#'FI;E0)'`H>'I,)',%V,)#$'3;0&)#'
were observed for the other Maldivian atolls (0.24). SI values were mostly low and
%00&#$-;$)';'#.)+%)#'(%3)-#%$K'#.)+%A+'$/');+,'/B'$,)'(%BB)-)"$';-);#'+/"#%()-)(>'8'"&4E)-'
of species were not observed in Baa; including Phaeophyceae taxa, among which several
species of Turbinaria and Sargassum. This latter genus was not observed in Baa atoll
during the present study nor has it been reported before. Nevertheless, several Sargassum
species have been mentioned by MRC staff and drift specimens have been collected from
other Maldivian atolls. The reason why species of this widespread genus is missing from
the Baa inventory warrants further investigation. Grazing pressure, seasonality or very
restricted distribution within Baa atoll (i.e. unprospected sites) are plausible hypotheses.
The absence of Sargassum /"'7;+%A+';$/00#'<;#'A-#$'(%#+&##)('EK'1/$K'FN]9`H>'I#&(;'
FN]^OH'()#+-%E)('$,)'.-)#)"+)'/B'S. crassifolium'/"'$</'7;+%A+';$/00#5'J0%$,%'8$/00'
(Yap State) and Kayangel Atoll (Palau); later Hodgson and McDermid (2000) reported
Sargassum sp. on Ant Atoll (Pohnpei State). Sterile plants were found in January at
Kayangel and fertile plants were found in June and July at Ulithi. The interesting fact is

43
that all Sargassum were collected on the northeast (windward side) of the atolls (Tsuda
com.pers.).It is interesting that the relative absence of Sargassum on Indian Ocean atolls
%#'#%4%0;-'$/'+;#)#'%"'$,)'7;+%A+'L+);">
Finally, our results address the issue of representativeness, which is critical in
biodiversity management. The little overlap of the macroalgal assemblages between
the different atolls demonstrates that, even at small biogeographical scales the spatial
,)$)-/V)")%$K'%#'%4./-$;"$>'I,%#'0);(#'$/'n&)#$%/"'$,)'+/"+).$'/B'b-).-)#)"$;$%3)'
protected area” in larger marine ecosystem like the Maldives regions atoll complex.
ACKNOWLEDGMENTS
Thanks are expressed to Serge Andréfouët and Shiham Adam who provided
&#'$,)'/../-$&"%$K'$/'#$&(K'B/-'$,)'A-#$'$%4)'$,)'4;+-/;0V;0'U/-;'/B'6;;>'I,)'.-/R)+$'
was supported by the French Fondation pour la Recherche sur la Biodiversité (FRB)
and by the Atoll Ecosystem Conservation project. This work is part of the COREUS
team Programme (Institut de Recherche pour le Développement). Laury Dijoux and
Nathalie Duong are thanked for their contribution to molecular analysis on Halimeda
and Dictyotales. Mélanie Hamel is thanked for her contribution to the various maps
given in this manuscript. The Marine Research Center of the Maldives is acknowledged
B/-'#+%)"$%A+'+/00;E/-;$%/"5';"('<)';-)')#.)+%;00K'$,;"TB&0'$/'D,;AK;':;))4';"('q//#&B'
Rilwan from MRC for their logistic and sampling assistance as well as their hospitality
;"('<;-4%"V'+/4.;"K'(&-%"V'$,)'A)0('#&-3)K>'_)';0#/'$,;"T'$,)':/;,'$);4'B/-',;3%"V'
made our life onboard easy and enjoyable. Finally, our thanks are addressed to Dr Roy
Tsuda for his helpful comments on the manuscript.
Table 4. Species richness, Sorensen’s similary Index (SI =2x/2x+y+z; where x is
the number of shared species, y'$,)'"&4E)-'/B'$,)'$/$;0'#.)+%)#'/B'$,)'A-#$'%#0;"('
and z is the total species of the second island or group) and % of common species
calculated between species diversity in Baa atoll and other archipelagoes of the
West Indian Ocean.
Baa
Maldives
Laccadives
Chagos
Seychelles
Reunion
Rhodophyta
108
200
71
26
212
122
Chlorophyta
58
]^
41
34
102
58
Phaeophyceae
10
24
20
7
54
36
Total species
176
321
132
68
374
215
Shared species
63
45
31
76
52
% Baa
35.7
25.5
17.6
43.1
W].5
Sorensen index
(SI)
0.24
0.22
0.2
0.21
0.21
!
!

44
REFERENCES
8"V%/#.)-4'7,K0/V)"K'\-/&.'!!!'Fb87\'!!!c@'6-)4)-5'6>5'r>'6-)4)-5'S>_>'=,;#)5'S>Q>'
Fay, J.L. Reveal, D.E. Soltis, P.S. Soltis, and P.F. Stevens, with collaboration of A.A.
Anderberg, M.J. Moore, R.G. Olmstead, P.J. Rudall, K.J. Sytsma, D.C. Tank, K. Wur-
dack, J.Q.-Y. Xiang, and S. Zmarzty)
WXX]>'' 8"'&.(;$)'/B'$,)'8"V%/#.)-4'7,K0/V)"K'\-/&.'+0;##%A+;$%/"'B/-'$,)'/--
()-#';"('B;4%0%)#'/B'U/<)-%"V'.0;"$#@'87\'!!!>Botanical Journal of the
Linnean Society'NON@'NX9GNWN>'
Aregood, C.C., and H.E. Hackett
N]^N>' '8'")<'Dictyurus (Rhodophyceae, dasyaceae) from the Maldive Islands,
Indian Oceanic Journal of the Elisha Mitchell Science Society'P^@']NG]O>
Barton E.S.
N]XY>'' e%#$'/B'4;-%")';0V;)'+/00)+$)(';$'$,)'S;0(%3)';"('e;++;(%3)'!#0;"(#'EK'[>D>'
\;-(%")-5'?#n>5'S>8>' Journal of the Linnean Society, Botany 35 : 475-482,
pl.
Bellwood, D.R., T.P. Hughes, and A.S. Hoey
2006. Sleeping functional group drives coral-reef recovery. Current Biology NO@'
W`Y`GW`Y]>
Bouchet, P., P. Lozouet, P. Maestrati, and V. Heros
2002. Assessing the magnitude of species richness in tropical marine
)"3%-/"4)"$#@')M+).$%/";00K',%V,'"&4E)-#'/B'4/00&#+#';$';':)<'
Caledonia site. Biological Journal of the Linnean Society'^9@'`WNs`YO>
Doty, M. S.
N]9`>'1%#$-%E&$%/"'/B'$,)';0V;0'V)")-;'Rhipilia and Sargassum in the central
' 7;+%A+>';$8)68%<8)'+8''P@'YO^GYOP>
Foslie, M.
N]XY>'' I,)'e%$,/$,;4"%;'/B'$,)'S;0(%3)#';"('e;++;(%3)#>'7;V)#'`OXG`^N'FZ/0>'NH'
in J.S. Gardiner (ed.). The fauna and geography of the Maldives and Lac-
cadive archipelagoes. Being the account of the work carried on and of the
collections made by an expedition during the years 1899 and 1900. Cam-
bridge University Press, Cambridge.
Foslie, M
N]X^>'' I,)'e%$,/$,;4"%;>'!"'\;-(%")-5'[>D>'N]X^GN]YO>'7)-+K'D0;()"'I-&#$'?M.)-
(%$%/"'$/'$,)'!"(%;"'L+);"'%"'N]X9>'NFWH'Transactions of the Linnean Soci-
ety of London, Second Series,Zoology'W@'N^^GN]W>
Gardiner, J. S.
N]XY>'' I,)'B;&";';"('V)/V-;.,K'/'B'$,)'S;0(%3)';"('e;++;(%3)'8-+,%.)0;V/)#>'
Vols. 1-2. Cambridge University Press, Cambridge.
Gepp, A., and E.S. Gepp
N]X`>'' Rhipidosiphon and Callipsygma. Journal of Botany'OW@'YOYGYOO>
Guiry, M.D., and G.M. Guiry
2011. AlgaeBase version 4.2. World-wide electronic publication, National
University of Ireland, Galway5',$$.@oo<<<>;0V;)E;#)>/-Vp'#earched on 15
April 2011.

45
Hackett, H.E.
N]O]>'' S;-%")';0V;)'%"'$,)';$/00')"3%-/"4)"$@'S;0(%3)'!#0;"(#>'Proceedings of the
International Seaweed Symposium O@'NP^GN]N>
Hackett, H.E.
N]^^>'' S;-%")';0V;)'T"/<"'B-/4'$,)'S;0(%3)'!#0;"(#>'Atoll Research Bulletin,
WNX@'NGYX5'.0#>'NC'W>
Hodgson, L. M., and K. J. McDermid.
WXXX>'' S;-%")'.0;"$#'/B'7/,".)%';"('8"$'8$/00@'' =,0/-/.,K$;5'7,;)/.,K$;';"('
Magnoliophyta. Micronesica'YW@'WP]GYX^
Hollenberg, G.J.
N]OP;>''8"';++/&"$'/B'$,)'#.)+%)#'/B'Polysiphonia of the central and western tropi-
+;0'7;+%A+'L+);">'!>'L0%V/#%.,/"%;>';$8)68%<8)'+8''WW@'9OG]P>
Hollenberg, G.J.
N]OPE>'8"';++/&"$'/B'$,)'#.)+%)#'/B'-)(';0V;) Herposiphonia occurring in the
+)"$-;0';"('<)#$)-"'$-/.%+;0'7;+%A+'L+);">';$8)68%<8)'+8''WW@'9YOG99]>
McCook, L.J.
N]]]>'' S;+-/;0V;)5'"&$-%)"$#';"('.,;#)'#,%B$#'/"'+/-;0'-))B#@'#+%)"$%A+'%##&)#';"('
4;";V)4)"$'+/"#)n&)"+)#'B/-'$,)'\-);$'6;--%)-'*))B>'Coral Reefs NP@'
357-367.
Newton, L.M.
N]9Y>'' S;-%")'80V;)>'I,)'[/,"'S&--;K'?M.)(%$%/"'N]YYGY`'D+%>'Reports']F9H@'
Y]9G`WX5'.0#>'!G!Z>'6->'S&#>'F:;$>'d%#$>H5'e/"(/">
Sigee, D.C.
N]OO>' '7-)0%4%";-K';++/&"$'/B'$,)'0;"(';"('4;-%")'3)V)$;$%/"'/B'8((&'8$/00>'..>'
61-74. In D. R. Stoddart (ed.), Reef studies at Addu Atoll, Maldive Islands,
7-)0%4%";-K'-)#&0$#'/B';"')M.)(%$%/"'$/'8((&'8$/00'%"'N]O`>'Atoll Research
Bulletin NNO@'NGNWW>
Silva, P.C., Basson, P.W. and Moe, R.L.
N]]O>' =;$;0/V&)'/B'$,)'E)"$,%+'4;-%")';0V;)'/B'$,)'!"(%;"'L+);">'University of
California Publications in Botany'^]@'NGNW9]>'
Stoddart, D.R. ed.
N]OO>'' *))B'#$&(%)#'%"'8((&'8$/005'S;0(%3)'%#0;"(#>'7-)0%4%";-K'-)#&0$#'/B';"')M-
.)(%$%/"'$/'8((&'8$/00'%"'N]O`>'Atoll Research Bulletin'NNO@'NGNWW>
Titlyanova, T.V., and P.V. Butorin
N]^P>'' 80V;)'/B'$,)'S;0(%3)';"('D)K+,)00)#'!#0;"(#>'!"'Biology of coral reefs:
photosynthesis of Zooxanthellae. Institute of Marine Biology, FarEast Sci-
ence Centre, Academy of Sciences of the USSR, Vladivostok, Transac-
$%/"#':/'NW@'N]GWP>
Tsuda, R. T.
N]^O>'' L++&--)"+)'/B'$,)'V)"&#'Sargassum F7,;)/.,K$;H'/"'$</'7;+%A+'' ;$/00#>'
Micronesica'NW@'W^]GWPW>
Tsuda, R.T., and J. Newhouse.
N]OO>'' S;-%")'E)"$,%+';0V;)'B-/4'8((&'8$/005'S;0(%3)'!#0;"(#>'..>']YtNXW>'
In D.R. Stoddart (ed.), Reef studies at Addu Atoll, Maldive Islands,

46
7-)0%4%";-K'-)#&0$#'/B';"')M.)(%$%/"'$/'8((&'8$/00'%"'N]O`>'Atoll Research
Bulletin NNO@'NtNWW>
Vroom, P. S. K.N. Page, K.A. Peyton and J. K. Kukea-Shultz
2005. Spatial heterogeneity of benthic community assemblages with an emphasis
on reef algae at French Frigate Shoals, Northwestern Hawai‘ian Islands.
Coral reefs'W`@9^`G9PN'>'
Weber Van Bosse, A., and M. Foslie
N]X`>' 'I,)'=/-;00%";+);)'/B'$,)'D%E/V;'?M.)(%$%/">'D%E/V;'?M.)(%$%)'ON>'?>[>'
Brill, Leyden.
Weber-van Bosse, A.
N]N`>'' I,)'7)-+K'D0;()"'I-&#$'?M.)(%$%/"'$/'$,)'!"(%;"'L+);"'%"'N]X95'&"()-'
$,)'0);()-#,%.'/B'S->'[>'D$;"0)K'\;-(%")-5'S>8>'*)./-$#@':/>'u!Z>'S;-%")'
algae, Rhodophyceae. Transactions of the Linnean Society of London, Sec-
ond Series, Zoology NO@'WO]GYXO>
Wynne M.J.
N]]Y>' '6)"$,%+'4;-%")';0V;)'B-/4'$,)'S;0(%3)#5'!"(%;"'L+);"5'+/00)+$)('(&-%"V'
the RV Te Vega Expedition. Contributions of the University of Michigan
Herbarium N]@'9GYX5'N`>

47
Appendix 1. Taxonomic list of the macrophytes recorded for Baa Atoll during the present study
Rhodophyta
Bonnemaisoniales
Bonnemaisoniaceae
Asparagopsis
taxiformis
(Delile) Trevisan
Ceramiales
Callithamniaceae
Crouania
minutissima
Yamada
Seirospora
orientalis
G. T. Kraft
Ceramiaceae
Centroceras
clavulatum
(C. Agardh) Montagne
Centroceras
minutum
Yamada
Ceramium
maryae
Weber-van Bosse
Ceramium
mazatlanense
E.Y. Dawson
Corallophila
apiculata
(Yamada) R.E. Norris
Cryptonemia
umbraticola
E.Y. Dawson
Gayliella
transversalis
FQ>D>'=/00%"#')$'d)-3)KH'I>L>'=,/')$'Q-)()-%+n
Griffithsia
heteromorpha
Kützing
Dasyaceae
Dasya
anastomosans
(Weber-van Bosse) M.J. Wynne
Dasya
baillouviana
(S. G. Gmelin) Montagne
Dasya
palmatifida
(Weber-van Bosse) A.J.K. Millar et E. Coppejans
Dictyurus
purpurascens
Bory de Saint-Vincent
Heterosiphonia
crispella
(C. Agardh) M.J. Wynne
Thuretia
sp
Delesseriaceae
Hypoglossum
simulans
M.J. Wynne, Price et Ballantine
Martensia
fragilis
Harvey
Martensia
sp. 'petit'
Myriogramme
heterostroma
N'Yeurt, M.J. Wynne et Payri
Myriogramme
melanesiensis
N'Yeurt, M.J. Wynne et Payri
Myriogramme
sp.
Nitophyllum
adhaerens
M. J. Wynne
Rhodomelaceae
Acanthophora
pacifica
(Setchell) Kraft
Chondria
arcuata
Hollenberg
Chondria
bullata
N'Yeurt et Payri
Chondria
ryukyuensis
Yamada
Chondria
simpliciuscula
Weber-van Bosse
Chondrophycus
succisus
(A.B. Cribb) K.W. Nam
Coelothrix
irregularis
(Harvey) Børgesen
Dipterosiphonia
dendritica
(C. Agardh) F. Schmitz

48
Herposiphonia
secunda
(C. Agardh) Ambronn f. tenella (C. Agardh) M.J. Wynne
Rhodophyta
Ceramiales
Rhodomelaceae
Laurencia
cf minuta
Vandermeulen, Garbary et Guiry
Laurencia
distichophylla
J. Agardh
Laurencia
sp. 1
Laurencia
sp. 2
Laurencia
sp. 3
Laurencia
sp. 4
Laurencia
sp. 5
Leveillea
jungermannioides
(K. Hering et G. Martens) Harvey
Neosiphonia
apiculata
(Hollenberg) Masuda et Kogame
Neosiphonia
ferulacea
(Suhr ex J. Agardh) S.M. Guimarães et M.T. Fujii
Palisada
parvipapillata
(C. K. Tseng) K. W. Nam
Polysiphonia
delicatula
Hollenberg
Polysiphonia
sertularioides
(Grateloup) J. Agardh
Tolypiocladia
glomerulata
(C. Agardh) F. Schmitz
Wrangeliaceae
Wrangelia
sp. inedit
Corallinales
Corallinaceae
Amphiroa
foliacea
Lamouroux in Quoy et Gaimard
Amphiroa
fragilissima
(Linnaeus) Lamouroux
Amphiroa
rigida
J.V. Lamouroux
Amphiroa
tribulus
(Ellis et Solander) Lamouroux
Amphiroa
sp
Hydrolithon
gardneri
!"#$%&'()*'+,'&-).)/+012,#33')456)7'&6'
Hydrolithon
onkodes
(Heydrich) D. Penrose et Woelkerling
Hydrolithon
reinboldii
(Weber-van Bosse et Foslie) Foslie
Hydrolithon
sp
Jania
adhaerens
Lamouroux
Lithophyllum
bamleri
(Heydrich) Heydrich
Lithophyllum
kotschyanum
Unger
Lithothamnion
proliferum
Foslie
Lithothamnion
sp
Mesophyllum
erubescens
(Foslie) M. Lemoine
Mesophyllum
sp
Neogoniolithon
brassica-florida
(Harvey) Setchell et L.R. Mason
Neogoniolithon
frutescens
!"#$%&'()8'9:,'%%).);<7<=5$#6
Appendix 1 (Con’td)

`]
Neogoniolithon
laccadivicum
!"#$%&'()*'+,-'%%).)/0$#1
Rhodophyta
Gelidiales
Gelidiaceae
Caulacanthus
ustulatus
(Turner) Kützing
Gelidium
isabelae
W.R. Taylor
Gelidium
sp
Pterocladiella
caespitosa
(Kylin) Santelices
Pterocladiella
caloglossoides
(M.A. Howe) Santelices
Gelidiellaceae
Gelidiella
acerosa
(Forsskål) Feldmann et G. Hamel
Gelidiella
myrioclada
(Børgesen) Feldmann et G. Hamel
Gigartinales
Corynocystaceae
Corynocystis
prostrata
G.T. Kraft
Dumontiaceae
Gibsmithia
dotyi
Kraft et Ricker
Gibsmithia
hawaiiensis
Doty
Hypneaceae
Hypnea
nidulans
Setchell
Hypnea
pannosa
J. Agardh
Hypnea
spinella
(C. Agardh) Kützing
Kallymeniaceae
Kallymenia
thompsonii
Abbott et McDermid
Peyssonneliaceaea
Peyssonnelia
cf. boergesenii
Weber-van Bosse
Peyssonnelia
inamoena
Pilger
Rhizophyllidaceae
Portieria
hornemannii
(Lyngbye) P.C. Silva
Halymeniales
Halymeniaceae
Halymenia
actinophysa
M. A. Howe
Halymenia
durvillei
Bory de Saint-Vincent
Halymenia
maculata
J. Agardh
Nemaliales
Galaxauraceae
Actinotrichia
fragilis
(Forsskål) Børgesen
Actinotrichia
sp
Galaxaura
filamentosa
R. Chou
Rhodymeniales
Champiaceae
Champia
compressa
Harvey
Champia
parvula
(C. Agardh) Harvey
Coelothrix
irregularis
(Harvey) Børgesen
Faucheaceae
Gloiocladia
iyoensis
(Okamura) R. Norris
Leptofaucheaceae
Leptofauchea
sp
Rhodymeniaceae
Asteromenia
anastomosans
(Weber-van Bosse) G. W. Saunders, C. E. Lane, C. W.
Schneider et Kraft
Botryocladia
skottsbergii
(Børgesen) Levring
Botryocladia
tenuissima
W.R. Taylor
Appendix 1 (Con’td)

50
Rhodophyta
Rhodymeniales
Rhodymeniaceae
Chamaebotrys
boergesenii
(Weber-van Bosse) Huisman
Gelidiopsis
intricata
(C. Agardh) Vickers
Halichrysis
irregularis
Kützing
Lomentaria
corallicola
Børgesen
Rhodymenia
sp.1
Rhodymenia
sp.2
Rhodymenia
sp.4
Rhodymenia
sp.5
Spirocladia
barodensis
Børgesen
Sporolithales
Sporolithaceae
Sporolithon
ptychoides
Heydrich
Chlorophyta
Bryopsidales
Bryopsidaceae
Bryopsis
pennata
J.V. Lamouroux
Bryopsis
plumosa
(Hudson) C. Agardh
Bryopsis
sp
Caulerpaceae
Caulerpa
cupressoides
(Vahl) C. Agardh
Caulerpa
diligulata
G.T. Kraft et A.J.K. Millar
Caulerpa
filicoides
Yamada
Caulerpa
nummularia
Harvey ex J. Agardh
Caulerpa
racemosa
(Forsskål) J. Agardh var. peltata (Lamouroux) Eubank
Caulerpa
sedoides
C. Agardh
Caulerpa
serrulata
(Forsskål) J. Agardh
Caulerpa
sertularioides
(S. Gmelin) M. Howe
Caulerpa
taxifolia
(Vahl) C. Agardh
Caulerpa
tongaensis
Papenfuss
Codiaceae
Codium
arabicum
Kützing
Codium
geppiorum
O.C. Schmidt
Halimedaceae
Halimeda
cuneata
Hering
Halimeda
discoidea
Decaisne
Chlorophyta
Bryopsidales
Halimedaceae
Halimeda
distorta
(Yamada) Hillis-Colinvaux
Halimeda
fragilis
W.R. Taylor
Halimeda
gracilis
Harvey ex J. Agardh
Halimeda
micronesica
Yamada
Halimeda
minima
(W.R. Taylor) Colinvaux
Halimeda
opuntia
(Linnaeus) Lamouroux
Halimeda
taenicola
W.R. Taylor
Appendix 1 (Con’td)

51
Chlorophyta
Bryopsidales
Halimedaceae
Halimeda
velasquezii
W.R. Taylor
Halimeda
xishaensis
S>e>1/"V'C'=>r>I#)"V
Halimeda
sp
Udoteaceae
Avrainvillea
lacerata
Harvey ex J. Agardh
Boodleopsis
pusilla
(F.S. Collins) W.R. Taylor, A.B. Joly et Bernatowicz
Boodleopsis
sp.
Chlorodesmis
fastigiata
(C. Agardh) Ducker
Chlorodesmis
hildebrandtii
A. Gepp et E.S. Gepp
Rhipidosiphon
javensis
Montagne
Rhipilia
crassa
A.J.K. Millar et Kraft
Rhipiliella
verticillata
G.T. Kraft
Rhipiliopsis
gracilis
Kraft
Tydemania
expeditionis
Weber-van Bosse
Chlorophyta
Cladophorales
Anadyomenaceae
Anadyomene
wrightii
Harvey ex J. Gray
Cladophora
dotyana
Gilbert
Cladophora
feredayoides
Kraft et Millar
Cladophora
goweri
A.H.S. Lucas
Cladophora
prehendens
Kraft et Millar
Cladophora
rupestris
(Linnaeus) Kützing
Cladophora
vagabunda
(Linnaeus) Hoek
Cladophora
sp.
Microdictyon
okamurae
Setchell
Siphonocladacaeae
Boergesenia
forbesii
(Harvey) J. Feldmann
Boodlea
composita
(Harvey) F. Brand
Cladophoropsis
herpestica
(Montagne) M.A. Howe
Cladophoropsis
vaucheriaeformis
(J.E Areschoug) Papenfuss
Dictyosphaeria
cavernosa
(Forsskål) Børgesen
Dictyosphaeria
versluysii
Weber-van Bosse
Phyllodictyon
anastomosans
(Harvey) Kraft et M.J. Wynne
Valoniaceae
Valonia
aegagropila
C. Agardh
Valonia
fastigiata
Harvey ex J. Agardh
Valonia
ventricosa
J. Agardh
Valoniopsis
pachynema
(G. Martens) Børgesen
Chlorophyta
Dasycladales
Dasycladaceae
Neomeris
annulata
Dickie
Appendix 1 (Con’td)

52
Polyphysaceae
Parvocaulis
parvulus
(Solms-Laubach) S. Berger et al.
Ulvales
Ulvaceae
Ulva
flexuosa
(Wulfen) J. Agardh
Phaeophyceae
Dictyotales
Dictyotaceae
Dictyopteris
repens
(Okamura) Børgesen
Dictyota
bartayresiana
Lamouroux
Phaeophyceae
Dictyotales
Dictyotaceae
Dictyota
ceylanica
Kützing
Dictyota
friabilis
Setchell
Dictyota
grossedentata
De Clerck et Coppejans
Dictyota
humifusa
Hörnig, Schnetter et Coppejans
Dictyota
sp.1
Lobophora
variegata
(Lamouroux) Womersley ex Oliveira
Padina
okinawaensis
Ni-NI-_%"5'D>'8-;%'C'd>'r;<;%
Fucales
Sargassaceae
Turbinaria
ornata
(Turner) J. Agardh
Magnolophyta
Alimastales
Cymodoceacea
Syringodium
isoetifolium
(Ascherson) Dandy
Hydrocharitaceae
Thalassia
hemprichii
(Ehrenberg) Ascherson
!
Appendix 1 (Con’td)

53
Appendix 2. Presence /absence of macroalgal species at prospected sites in Baa Atoll
Genus
species
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
27
28
!
!
Anadyomene wrightii
!!
!!
"!
!!
!!
!!
!!
"!
!!
!!
!!
"!
!!
!!
"!
!!
"!
"!
!!
!!
"!
!!
!!
"!
!!
!!
!!
Avrainvillea lacerata
!
!
!
!
!
!
!
!
!
!
"!
!
!
"!
!
!
!
!
"!
!
"!
"!
!
!
"!
!
"!
Boergesenia forbesii
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
Boodlea composita
!
!
!
!
!
!
!
"!
"!
!
"!
!
"!
!
!
"!
!
"!
"!
!
"!
!
!
!
!
!
"!
Boodleopsis pusilla
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
Boodleopsis
#$%!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Bryopsis pennata
!!
!!
"!
"!
!!
!!
"!
"!
!!
"!
"!
"!
!!
"!
!!
"!
!!
!!
"!
!!
"!
"!
!!
"!
"!
!!
!!
Bryopsis plumosa
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Bryopsis
#$%!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
"!
Caulerpa cupressoides
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Caulerpa diligulata
"!
!!
!!
"!
!!
!!
"!
!!
!!
"!
"!
!!
!!
"!
!!
!!
"!
"!
"!
!!
!!
!!
!!
!!
!!
"!
!!
Caulerpa filicoides
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
Caulerpa nummularia
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
"!
!!
!!
Caulerpa racemosa
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
"!
Caulerpa sedoides
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Caulerpa serrulata
!
!
!
!
"!
!
!
!
"!
!
!
!
!
!
!
!
"!
"!
!
!
!
!
!
!
!
!
!
Caulerpa sertularioides
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Caulerpa taxifolia
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
"!
"!
!
!
!
!
!
!
!
!
!
!
Caulerpa tongaensis
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Chlorodesmis fastigiata
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
"!
"!
"!
!
!
!
!
!
!
!
!
Chlorodesmis hildebrandtii
"!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
Cladophora dotyana
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
"!
!
!
!
!
!
!
!
!
!
Cladophora feredayoides
"!
!!
!!
"!
!!
!!
"!
!!
!!
"!
!!
!!
!!
"!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Cladophora goweri
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Cladophora prehendens
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
Cladophora rupestris
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Cladophora vagabunda
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
Cladophora
#$%!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Cladophoropsis herpestica
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
"!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
Cladophoropsis vaucheriaeformis
"!
!
!
"!
!
!
"!
!
!
"!
"!
!
!
"!
"!
!
!
"!
"!
!
!
!
!
!
!
"!
!
Codium arabicum
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
"!

54
Codium geppiorum !"
"
"
"
"
"
"
!"
"
"
"
!"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
Dictyosphaeria cavernosa "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" !" "" ""
Dictyosphaeria versluysii
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
Halimeda cuneata !" "" !" "" !" !" "" "" "" "" "" "" "" !" "" "" !" !" !" "" !" !" "" !" !" !" !"
Halimeda distorta !"
"
"
"
!" !" !"
"
"
"
!"
"
"
!"
"
"
"
"
!"
"
"
"
!"
"
"
!"
"
Halimeda gracilis !" "" "" "" !" !" !" "" "" "" !" "" "" "" "" !" !" !" "" "" !" "" "" !" !" "" !"
Halimeda micronesica
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
!" !"
"
"
"
"
"
"
"
Halimeda minima "" "" !" "" !" !" !" !" "" !" !" !" "" !" "" !" !" !" !" "" !" !" "" !" !" !" !"
Halimeda opuntia !" !"
"
"
"
!"
"
!" !" !" !" !" !" !" !"
"
"
!" !" !"
"
"
"
"
!"
"
!"
Halimeda taenicola
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
!" !"
"
"
"
"
"
"
"
Halimeda velasquezii "" "" "" "" "" "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" ""
Halimeda xishaensis
"
"
"
"
"
"
!"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
"
"
"
!"
"
Halimeda #$%" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Microdictyon okamurae "" "" "" "" "" "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" ""
Neomeris annulata
"
"
"
"
"
"
"
!" !"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
Parvocaulis parvulus "" "" "" "" "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Phyllodictyon anastomosans
"
"
!"
"
"
"
!"
"
"
"
!"
"
"
!"
"
"
"
!"
"
"
!"
"
"
!"
"
!"
"
Rhipidosiphon javensis "" "" !" !" !" !" "" "" "" "" "" !" "" !" "" "" !" !" !" "" "" !" "" !" !" "" !"
Rhipilia crassa
"
"
"
"
"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
"
"
"
"
!"
Rhipiliella verticillata "" "" !" "" "" "" "" "" "" "" "" !" "" "" "" "" !" "" "" "" "" "" "" !" "" "" ""
Rhipiliopsis gracilis
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
Tydemania expeditionis !" "" !" !" "" !" !" "" "" !" !" !" "" !" !" "" !" !" !" !" !" !" "" !" !" !" !"
Ulva flexuosa
"
"
"
"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
Valonia aegagropila "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Valonia fastigiata
"
"
"
!"
"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
"
"
"
"
"
Valonia ventricosa !" "" !" "" !" "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" !" "" !" "" "" ""
Valoniopsis pachynema
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
!"
"
"
"
"
"
"
Dictyopteris repens "" "" !" "" "" "" "" "" "" "" "" !" "" "" !" "" "" "" "" "" !" "" "" "" !" "" ""
Dictyota bartayresiana
"
"
"
"
"
!" !"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
Dictyota ceylanica "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Dictyota friabilis !"
"
"
"
"
!" !" !"
"
!" !"
"
"
"
"
"
!" !" !"
"
!"
"
"
!" !"
"
"
Dictyota grossedentata "" "" "" "" "" "" "" "" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Dictyota humifusa
"
"
"
"
"
"
!"
"
"
!" !"
"
"
"
"
"
"
!"
"
"
"
"
"
"
"
"
!"
Dictyota #$%!" "" "" !" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ""
Lobophora variegata !"
"
!"
"
!" !" !"
"
"
!" !" !"
"
"
"
"
"
!"
"
"
!"
"
"
!"
"
"
"
Padina okinawaensis "" "" "" "" "" "" "" "" !" "" "" "" "" "" !" "" "" !" "" "" "" "" "" "" "" "" ""
Appendix 2 (Con’td)
Genus
species
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
27
28

55
Turbinaria ornata
!
!
!
!
"! "!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Acanthophora pacifica !! !! !! !! !! "! "! !! !! "! "! !! !! "! !! "! !! !! !! !! !! !! !! !! !! "! !!
Actinotrichia fragilis
!
!
"!
!
"!
!
!
!
!
!
"! "!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
Actinotrichia #$%! "! !! "! !! !! "! "! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Amphiroa foliacea
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Amphiroa fragilissima !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !!
Amphiroa rigida
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Amphiroa tribulus
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Amphiroa sp. !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Asparagopsis taxiformis !! !! !! !! !! !! !! "! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !!
Asteromenia anastomosans
!
!
!
"!
!
!
"!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
"!
!
Botryocladia skottsbergii !! !! !! !! !! "! !! !! !! !! "! !! !! !! "! !! !! !! !! !! "! !! !! !! !! "! "!
Botryocladia tenuissima
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Caulacanthus ustulatus !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "!
Centroceras clavulatum
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Centroceras minutum !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Ceramium maryae
!
!
"!
!
"!
!
!
!
!
"! "!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
Ceramium mazatlanense !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !!
Chamaebotrys boergesenii "!
!
!
!
!
!
"!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
Champia compressa !! !! "! "! !! "! "! !! !! !! "! !! !! !! !! !! "! !! !! !! !! "! !! "! "! !! "!
Champia parvula
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
Chondria arcuata !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !!
Chondria bullata
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
Chondria ryukyuensis !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !!
Chondria simpliciuscula
!
!
"!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Chondrophycus succisus !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Coelothrix irregularis
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
Corallophila apiculata !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Corynocystis prostrata
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
"!
!
!
!
!
!
!
Crouania minutissima !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !!
Cryptonemia umbraticola "!
!
!
!
"!
!
!
!
!
!
!
!
!
"!
!
!
"! "!
!
!
!
!
!
!
!
!
!
Dasya anastomosans !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !!
Dasya baillouviana
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
"!
!
!
!
!
!
!
!
!
!
Dasya palmatifida !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Dictyurus purpurascens
!
!
"!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
"!
Dipterosiphonia dendritica !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Appendix 2 (Con’td)
Genus
species
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
27
28

56
Leveillea jungermannioides !! !! !! !! !! !! !! !! !! !! "! !! "! !! "! !! !! !! !! !! !! !! !! !! !! !! !!
Lithophyllum bamleri "!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Lithophyllum kotschyanum "! !! "! !! "! "! !! !! !! !! !! !! !! !! !! "! !! !! !! !! "! "! !! "! "! !! "!
Lithothamnion proliferum "! !! !! "! "! !! "! !! !! "! "! !! !! "! !! !! !! "! !! !! "! "! !! "! "! "! "!
Lithothamnion #$!
!
!
!
"! "!
!
!
!
!
"!
!
!
!
"!
!
!
!
"!
!
!
"! "!
!
"! "! "!
!
Lomentaria corallicola
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Martensia fragilis !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! "! !!
Martensia #$!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
"!
!
!
Mesophyllum erubescens !! !! !! !! !! !! !! !! !! "! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Mesophyllum #$!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
Myriogramme heterostroma !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! "! !! !! !!
Myriogramme melanesiensis
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
Myriogramme #$! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! "! !! !! !!
Neogoniolithon brassica-florida
!
!
!
!
!
!
!
!
"!
!
!
!
"!
!
!
!
!
!
!
!
"! "!
!
!
!
!
!
Neogoniolithon frutescens "! "! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! "! "! !! !! !! !! !!
Neogoniolithon laccadivicum
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
Neosiphonia apiculata !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !!
Neosiphonia ferulacea
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
Nitophyllum adhaerens !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Palisada
parvipapillata
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Peyssonnelia cf. boergesenii !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !!
Peyssonnelia inamoena
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
"!
!
!
!
!
!
Polysiphonia delicatula !! !! !! "! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Polysiphonia sertularioides
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
Portieria hornemannii !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Pterocladiella caespitosa
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
Pterocladiella caloglossoides !! !! !! !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Rhodymenia #$%"!
!
!
!
!
!
!
!
!
!
"! "!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Rhodymenia #$%&! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !!
Rhodymenia #$%'!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
Rhodymenia #$%(! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! "!
Seirospora orientalis
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
!
!
!
!
!
Spirocladia barodensis !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !!
Sporolithon ptychoides
!
!
!
"! "! "! "!
!
!
!
"!
!
!
"!
!
!
!
"!
!
!
!
!
!
"!
!
!
"!
Thuretia #$%! !! !! "! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Tolypiocladia glomerulata
!
!
!
!
!
!
"!
!
"!
!
!
!
"!
!
!
!
!
!
!
!
!
"!
!
!
!
!
!
Appendix 2 (Con’td)
Genus
species
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
27
28

57
Wrangelia sp. inedit !! !! !! !! !! !! !! !! !! !! !! !! !! !! "! !! !! !! !! !! !! !! !! !! "! !! !!
Syringodium isoetifolium !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
Thalassia hemprichii !! "! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !! !!
!
Appendix 2 (Con’td)
Genus
species
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
27
28

58
Appendix 3
Rhodophyta 1/4
Gibsmithia hawaiensis!Gibsmithia dotyii!!
Botryocladia skottsbergii Cryptonemia umbraticola
Corynocystis prostrata
Asteromenia anastomosans
!

9]
Rhodophyta 2/4
!
Actinotrichia!"#! Actinotrichia fragilis
!
!
Acanthophora pacifica Amphiroa tribulus
Dictyurus purpurascens!
Kallymenia thompsonii
!

60
Rhodophyta 3/4
Halymenia durvillei
!
Coelothrix irregularis
Gelidium isabelae!Heterosiphonia crispella
Martensia fragilis
!
Myriogramme heterostroma
Myriogramme!"#$!
!
Peyssonnelia "#$!

61
Rhodophyta 3/4
Hydrolithon gardneri
!
Lithothamnion proliferum
!
Hydrolithon "#$!
!
Sporolithon ptychoides.!

62
Chlorophyta 1/4
Caulerpa serrulata!!
Caulerpa racemosa var. peltata
Caulerpa diligulata Tydemania expeditionis
Avrainvillea lacerata!
Rhipidosiphon javensis
!

63
Chlorophyta 2/4
Halimeda cuneata Halimeda distorta
Halimeda gracilis Halimeda micronesica
Halimeda minima
Halimeda opuntia!!
!
!

64
Chlorophyta 3/4
Bryopsis pennata!
Cladophoropsis vaucheriaeformis
Chlorodesmis fastigiata!
Phyllodictyon anastomosans
!!
Cladophora feredayoides!
Microdictyon umbilicatum
!

65
Chlorophyta 4/4
Valonia fastigiata!
Valonia ventricosa!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
Boergesenia forbersii !Boodleopsis pusilla!!
!
!
!
!
!
!
!

66
Phaeophyceae 1/1
Lobophora variegata Padina okinawensis!
Dictyota friabilis
!
Dictyota humifusa
Magnolophyta 1/1
!
Cymodocea serrulata!Syringodium isoetifolium