ATOLL RESEARCH BULLETIN NO. 590
BIODIVERSITY, RESOURCES, AND CONSERVATION
OF
BAA ATOLL (REPUBLIC OF MALDIVES):
A UNESCO MAN AND BIOSPHERE RESERVE
Edited by
Serge Andréfouët
Issued by
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. U.S.A.
JANUARY 2012
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%"+)>'!"'$,%#'.;.)-5'
we give a general description of the representative macrophyte communities of Baa Atoll
in relation to the geomorphology of reefs.
________________________________________________________
1
!"#$%$&$'()'*)+,)-+,)'./&-'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;-%")'*)#);-+,'!"#$%$&$)5'=;.)'
Town, South Africa
32
INTRODUCTION
Z)-K'0%4%$)('%"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;&$,/-#'FN]]OH> The species list is given in Appendix 1. Records belong to 17 orders,
Y9'B;4%0%)#';"(']`'V)")-;'FI;E0)'WH>
34
I;E0)'N>'D;4.0%"V'#%$)#'(%#$-%E&$%/"'%"'$,)'#)3)"',;E%$;$'+0;##)#'()A")('E;#)('/"'
geomorphology.
Class
Habitat
number
of sites
Oceanic
reef flat
(seagrass)
Lagoon
reef flat
and slope
Lagoon
patch reef
Lagoon
reef flat
Deep
lagoon
Oceanic
reef flat
Oceanic
reef slope
(1)
(10)
(5)
(3)
(1)
(2)
(5)
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
Site
label
Figure 1. Location of the sampling sites in Baa Atoll.
35
I;E0)'W>':&4E)-'/B'L-()-#5'Q;4%0%)#5'\)")-;';"('D.)+%)#'/B'4;+-/.,K$)#'%()"$%A)('B-/4'
Baa Atoll
Macroalgae
Seagrasses
Rhodophyta
Chlorophyta
Phaeophyceae
TOTAL
Magnoliophyta
Orders
10
5
2
17
1
Families
22
11
2
35
2
Genera
64
25
5
]`
2
Species
108
58
10
176
2
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)-";$/<%+l'<,%+,'/++&--)('
on faros’ sandy bottoms. (Appendix 2).
Species Richness Distribution in Baa Atoll.
Species richness per site. Species richness per site (Fig. 2) ranged from two at
Baa 23 (deep lagoon) to 38 species at Baa 11 (lagoon patch reef) and Baa 7 (oceanic reef
slope) (Fig. 1). These two sites, coral built and exposed to strong water movements, were
the most diverse and displayed the highest species richness with 38 spp.
Figure 2. Distribution of species richness per sites.
36
Because only two species have been collected from the sandy bottom of the
()).)#$'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).
Figure 3.Spatial distribution of the species richness in Baa Atoll.
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'.;$+,)#5'
pinnacles and deep lagoon sections.
No contrasting spatial variation could be observed between the different sections
of the atoll rim (Table 3). The sampling effort in each geographic area was not strictly
similar and could partially affect the resulting values. The average species richness did
not contrast strongly from North (21 spp.) to South (28 spp.) and from East (26.2 spp.)
to West (27 spp.). Nevertheless, the lowest species richness was observed in the northern
rim section. No contrasting difference could be found between the average species
richness of the global atoll rim (25.2 spp.) and the lagoonal faros (24.7 spp.).
37
Table 3. Species richness per geographic area (sites 2 and 20 were excluded because not
properly prospected)
Geographic Area
Northern
Rim
Southern
Rim
Eastern
Rim
Western
Rim
Atoll Rim
Atoll
Lagoon
Number of sites
(4)
14, 15,
16, 17
(3)
1, 3, 4
(5)
^5'P5']5
NP5'N]
(2)
27, 28
21
28
26.2
27
(14)
1, 3, 4, 7,8,
]5'N`5'N95
16, 17, 18,
N]5 27, 28
25.2
(8)
5, 6,10,
11,12,13,
21, 22, 23,
24, 25
24.7
Site label (BaaX)
Average species
Species richness per geomorphological habitat type. Average species richness
varied within each geomorphological habitat type (Fig. 4). Figure 4 shows that the deep
0;V//"';"('$,)'#);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;$';"('#0/.)a';"('n= 5 sites for the ‘lagoon patch reef’). The oceanic slope
showed an average of 30 spp. for the 5 sites.
The spatial distribution of the species varied with bathymetry and some
species were observed at a wide range of depths. This is true for instance for the green
algae Caulerpa diligulata, Halimeda discoidea, Rhipidosiphon javensis, Tydemania
expeditionis and the red Botryocladia which were found from 5 m down to 40 m,
independently of geographical location. Other species were found to be restricted to deep
l/")#5'#&+,';#@'Padina okinawensis, Cladophora feredayoides, Microdictyon okamurae,
Caulerpa sedoides, or cryptic and only in shaded areas such as Cryptonemia umbraticola
or Corynocystis prostrata. Many other species were restricted to shallow waters such as
Valonia aegagropila, Dictyurus purpurascens, Halymenia actinophysa, Hypnea spp. and
Turbinaria ornata.
Figure 4. Variation of species richness per habitat type.
38
Species rarity. Patterns of the algal vegetation in the Baa Atoll were characterized
by the dominance of species with very low occurrences at all the prospected sites (Fig. 5).
Half of the species (n=87) were found in less than 8 % (2/27) of the sites thus revealing
;',%V,'E)$;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^'#%$)#>'
Ninety percent of the species occurred in only 13 sites which represents less than half of
the prospected sites. Less than 10% of the species were represented by a single specimen.
Figure 5. Histogram of species occurrences at the 27 sites.
Main Algal Assemblages.
The most common species are illustrated in Appendix 3.
!"#$"%&'#'($()*+%*+%"$#**+%,''-%.$(/%$+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.
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);#$'$,-))'/&$'/B'A3)'
sites visited in this same geomorphologic habitat type. The assemblage was dominated by
large green Tydemania expeditionis, as well as Halimeda minima and H. opuntia with a
lesser abundance of by H. cuneata and H. gracilis, the bright green pompom-like Chlorodesmis fastigiata, the dark green Avrainvillea lacerata and Asteromenia anastomosans.
Caulerpa diligulata /++&--)(';$';00'#%$)#'+0;##%A)('%"'$,%#'V)/4/-.,/0/V%+;0',;E%$;$'$K.)>'
Most of the investigated sites showed turf assemblages associated with dead coral including mostly Gelidiopsis intricata, Dictyota humifusa, and Champia compressa. Several
species such as !8$+(9*19*,$%1$8)68$, Caulerpa diligulata, Cladophoropsis vaucheriaeformis, C. herpestica were also common component of the oceanic reef slopes assemblages 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 proliferum was easily recognizable thanks to its pink crust and numerous short knobs. Along
the slope several Rhodymeniales including Leptofauchea spp. and Rhodymenia spp. occurred 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 javensis 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 anastomosans 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 okinawensis. 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 classiA)('%"'$,)'/-()-'80%#4;$;0)#'F"/4)"+0;$&-)'E;#)('/"'.,K0/V)")$%+'#$&(%)#'87\!!!5'WXX]H>'
In tropical regions, they are almost permanently immersed in sheltered marine and estuarine 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)")-;$)',/4/V)")/&#'
environmental forcing thus limiting habitat diversity and in turn leading to a more or
0)##',/4/V)")/�K'(%#$-%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&)#$%/"'$,)'+/"+).$'/B'b-).-)#)"$;$%3)'
protected area” in larger marine ecosystem like the Maldives regions atoll complex.
Table 4. Species richness, Sorensen’s similary Index (SI =2x/2x+y+z; where x is
the number of shared species, y'$,)'"&4E)-'/B'$,)'$/$;0'#.)+%)#'/B'$,)'A-#$'%#0;"('
and z is the total species of the second island or group) and % of common species
calculated between species diversity in Baa atoll and other archipelagoes of the
West Indian Ocean.
Rhodophyta
Chlorophyta
Phaeophyceae
Total species
Shared species
% Baa
Sorensen index
(SI)
Baa
108
58
10
176
Maldives
200
]^
24
321
63
35.7
0.24
Laccadives
71
41
20
132
45
25.5
0.22
Chagos
26
34
7
68
31
17.6
0.2
Seychelles
212
102
54
374
76
43.1
0.21
Reunion
122
58
36
215
52
W].5
0.21
!
!
ACKNOWLEDGMENTS
Thanks are expressed to Serge Andréfouët and Shiham Adam who provided
&#'$,)'/../-$&"%$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#/'$,;"T'$,)':/;,'$);4'B/-',;3%"V'
made our life onboard easy and enjoyable. Finally, our thanks are addressed to Dr Roy
Tsuda for his helpful comments on the manuscript.
44
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algae, Rhodophyceae. Transactions of the Linnean Society of London, Second Series, Zoology NO@'WO]GYXO>
Wynne M.J.
N]]Y>' '6)"$,%+'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`>
Appendix 1. Taxonomic list of the macrophytes recorded for Baa Atoll during the present study
Rhodophyta
Bonnemaisoniales Bonnemaisoniaceae Asparagopsis
Crouania
Ceramiales
Callithamniaceae
Seirospora
Centroceras
Ceramiaceae
Centroceras
Ceramium
Ceramium
Corallophila
Cryptonemia
Gayliella
Griffithsia
Dasya
Dasyaceae
Dasya
Dasya
Dictyurus
Heterosiphonia
Thuretia
Hypoglossum
Delesseriaceae
Martensia
Martensia
Myriogramme
Myriogramme
Myriogramme
Nitophyllum
Acanthophora
Rhodomelaceae
Chondria
Chondria
Chondria
Chondria
Chondrophycus
Coelothrix
Dipterosiphonia
taxiformis
minutissima
orientalis
clavulatum
minutum
maryae
mazatlanense
apiculata
umbraticola
transversalis
heteromorpha
anastomosans
baillouviana
palmatifida
purpurascens
crispella
sp
simulans
fragilis
sp. 'petit'
heterostroma
melanesiensis
sp.
adhaerens
pacifica
arcuata
bullata
ryukyuensis
simpliciuscula
succisus
irregularis
dendritica
(Delile) Trevisan
Yamada
G. T. Kraft
(C. Agardh) Montagne
Yamada
Weber-van Bosse
E.Y. Dawson
(Yamada) R.E. Norris
E.Y. Dawson
FQ>D>'=/00%"#')$'d)-3)KH'I>L>'=,/')$'Q-)()-%+n
Kützing
(Weber-van Bosse) M.J. Wynne
(S. G. Gmelin) Montagne
(Weber-van Bosse) A.J.K. Millar et E. Coppejans
Bory de Saint-Vincent
(C. Agardh) M.J. Wynne
M.J. Wynne, Price et Ballantine
Harvey
N'Yeurt, M.J. Wynne et Payri
N'Yeurt, M.J. Wynne et Payri
M. J. Wynne
(Setchell) Kraft
Hollenberg
N'Yeurt et Payri
Yamada
Weber-van Bosse
(A.B. Cribb) K.W. Nam
(Harvey) Børgesen
(C. Agardh) F. Schmitz
47
Rhodophyta
Ceramiales
Corallinales
48
Appendix 1 (Con’td)
Rhodomelaceae
Wrangeliaceae
Corallinaceae
Herposiphonia
Laurencia
Laurencia
Laurencia
Laurencia
Laurencia
Laurencia
Laurencia
Leveillea
Neosiphonia
Neosiphonia
Palisada
Polysiphonia
Polysiphonia
Tolypiocladia
Wrangelia
Amphiroa
Amphiroa
Amphiroa
Amphiroa
Amphiroa
Hydrolithon
Hydrolithon
Hydrolithon
Hydrolithon
Jania
Lithophyllum
Lithophyllum
Lithothamnion
Lithothamnion
Mesophyllum
Mesophyllum
Neogoniolithon
Neogoniolithon
secunda
cf minuta
distichophylla
sp. 1
sp. 2
sp. 3
sp. 4
sp. 5
jungermannioides
apiculata
ferulacea
parvipapillata
delicatula
sertularioides
glomerulata
sp. inedit
foliacea
fragilissima
rigida
tribulus
sp
gardneri
onkodes
reinboldii
sp
adhaerens
bamleri
kotschyanum
proliferum
sp
erubescens
sp
brassica-florida
frutescens
(C. Agardh) Ambronn f. tenella (C. Agardh) M.J. Wynne
Vandermeulen, Garbary et Guiry
J. Agardh
(K. Hering et G. Martens) Harvey
(Hollenberg) Masuda et Kogame
(Suhr ex J. Agardh) S.M. Guimarães et M.T. Fujii
(C. K. Tseng) K. W. Nam
Hollenberg
(Grateloup) J. Agardh
(C. Agardh) F. Schmitz
Lamouroux in Quoy et Gaimard
(Linnaeus) Lamouroux
J.V. Lamouroux
(Ellis et Solander) Lamouroux
!"#$%&'()*'+,'&-).)/+012,#33')456)7'&6'
(Heydrich) D. Penrose et Woelkerling
(Weber-van Bosse et Foslie) Foslie
Lamouroux
(Heydrich) Heydrich
Unger
Foslie
(Foslie) M. Lemoine
(Harvey) Setchell et L.R. Mason
!"#$%&'()8'9:,'%%).);<7<=5$#6
Appendix 1 (Con’td)
Rhodophyta
Gelidiales
Gelidiaceae
Gelidiellaceae
Gigartinales
Corynocystaceae
Dumontiaceae
Hypneaceae
Kallymeniaceae
Peyssonneliaceaea
Halymeniales
Rhizophyllidaceae
Halymeniaceae
Nemaliales
Galaxauraceae
Rhodymeniales
Champiaceae
Faucheaceae
Leptofaucheaceae
Rhodymeniaceae
Neogoniolithon
Caulacanthus
Gelidium
Gelidium
Pterocladiella
Pterocladiella
Gelidiella
Gelidiella
Corynocystis
laccadivicum
ustulatus
isabelae
sp
caespitosa
caloglossoides
acerosa
myrioclada
prostrata
!"#$%&'()*'+,-'%%).)/0$#1
(Turner) Kützing
W.R. Taylor
Gibsmithia
Gibsmithia
Hypnea
Hypnea
Hypnea
Kallymenia
Peyssonnelia
Peyssonnelia
Portieria
Halymenia
Halymenia
Halymenia
Actinotrichia
Actinotrichia
Galaxaura
Champia
Champia
Coelothrix
Gloiocladia
Leptofauchea
dotyi
hawaiiensis
nidulans
pannosa
spinella
thompsonii
cf. boergesenii
inamoena
hornemannii
actinophysa
durvillei
maculata
fragilis
sp
filamentosa
compressa
parvula
irregularis
iyoensis
sp
Kraft et Ricker
Doty
Setchell
J. Agardh
(C. Agardh) Kützing
Abbott et McDermid
Weber-van Bosse
Pilger
(Lyngbye) P.C. Silva
M. A. Howe
Bory de Saint-Vincent
J. Agardh
(Forsskål) Børgesen
Asteromenia
Botryocladia
Botryocladia
anastomosans
skottsbergii
tenuissima
(Kylin) Santelices
(M.A. Howe) Santelices
(Forsskål) Feldmann et G. Hamel
(Børgesen) Feldmann et G. Hamel
G.T. Kraft
R. Chou
Harvey
(C. Agardh) Harvey
(Harvey) Børgesen
(Okamura) R. Norris
`]
(Weber-van Bosse) G. W. Saunders, C. E. Lane, C. W.
Schneider et Kraft
(Børgesen) Levring
W.R. Taylor
50
Appendix 1 (Con’td)
Rhodophyta
Rhodymeniales
Rhodymeniaceae
Chlorophyta
Sporolithales
Bryopsidales
Sporolithaceae
Bryopsidaceae
Caulerpaceae
Codiaceae
Halimedaceae
Chlorophyta
Bryopsidales
Halimedaceae
Chamaebotrys
Gelidiopsis
Halichrysis
Lomentaria
Rhodymenia
Rhodymenia
Rhodymenia
Rhodymenia
Spirocladia
Sporolithon
Bryopsis
Bryopsis
Bryopsis
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Codium
Codium
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
boergesenii
intricata
irregularis
corallicola
sp.1
sp.2
sp.4
sp.5
barodensis
ptychoides
pennata
plumosa
sp
cupressoides
diligulata
filicoides
nummularia
racemosa
sedoides
serrulata
sertularioides
taxifolia
tongaensis
arabicum
geppiorum
cuneata
discoidea
distorta
fragilis
gracilis
micronesica
minima
opuntia
taenicola
(Weber-van Bosse) Huisman
(C. Agardh) Vickers
Kützing
Børgesen
Børgesen
Heydrich
J.V. Lamouroux
(Hudson) C. Agardh
(Vahl) C. Agardh
G.T. Kraft et A.J.K. Millar
Yamada
Harvey ex J. Agardh
(Forsskål) J. Agardh var. peltata (Lamouroux) Eubank
C. Agardh
(Forsskål) J. Agardh
(S. Gmelin) M. Howe
(Vahl) C. Agardh
Papenfuss
Kützing
O.C. Schmidt
Hering
Decaisne
(Yamada) Hillis-Colinvaux
W.R. Taylor
Harvey ex J. Agardh
Yamada
(W.R. Taylor) Colinvaux
(Linnaeus) Lamouroux
W.R. Taylor
Appendix 1 (Con’td)
Chlorophyta
Chlorophyta
Cladophorales
Dasycladales
Halimeda
Halimeda
Halimeda
Avrainvillea
Udoteaceae
Boodleopsis
Boodleopsis
Chlorodesmis
Chlorodesmis
Rhipidosiphon
Rhipilia
Rhipiliella
Rhipiliopsis
Tydemania
Anadyomene
Anadyomenaceae
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Microdictyon
Siphonocladacaeae Boergesenia
Boodlea
Cladophoropsis
Cladophoropsis
Dictyosphaeria
Dictyosphaeria
Phyllodictyon
Valonia
Valoniaceae
Valonia
Valonia
Valoniopsis
Neomeris
Dasycladaceae
Halimedaceae
velasquezii
xishaensis
sp
lacerata
pusilla
sp.
fastigiata
hildebrandtii
javensis
crassa
verticillata
gracilis
expeditionis
wrightii
dotyana
feredayoides
goweri
prehendens
rupestris
vagabunda
sp.
okamurae
forbesii
composita
herpestica
vaucheriaeformis
cavernosa
versluysii
anastomosans
aegagropila
fastigiata
ventricosa
pachynema
annulata
W.R. Taylor
S>e>1/"V'C'=>r>I#)"V
Harvey ex J. Agardh
(F.S. Collins) W.R. Taylor, A.B. Joly et Bernatowicz
(C. Agardh) Ducker
A. Gepp et E.S. Gepp
Montagne
A.J.K. Millar et Kraft
G.T. Kraft
Kraft
Weber-van Bosse
Harvey ex J. Gray
Gilbert
Kraft et Millar
A.H.S. Lucas
Kraft et Millar
(Linnaeus) Kützing
(Linnaeus) Hoek
Setchell
(Harvey) J. Feldmann
(Harvey) F. Brand
(Montagne) M.A. Howe
(J.E Areschoug) Papenfuss
(Forsskål) Børgesen
Weber-van Bosse
(Harvey) Kraft et M.J. Wynne
C. Agardh
Harvey ex J. Agardh
J. Agardh
(G. Martens) Børgesen
Dickie
51
Chlorophyta
Bryopsidales
parvulus
flexuosa
repens
bartayresiana
ceylanica
friabilis
grossedentata
humifusa
sp.1
variegata
okinawaensis
ornata
(Solms-Laubach) S. Berger et al.
(Wulfen) J. Agardh
(Okamura) Børgesen
Lamouroux
Kützing
Setchell
De Clerck et Coppejans
Hörnig, Schnetter et Coppejans
Sargassaceae
Parvocaulis
Ulva
Dictyopteris
Dictyota
Dictyota
Dictyota
Dictyota
Dictyota
Dictyota
Lobophora
Padina
Turbinaria
Cymodoceacea
Hydrocharitaceae
Syringodium
Thalassia
isoetifolium
hemprichii
(Ascherson) Dandy
(Ehrenberg) Ascherson
Ulvales
Phaeophyceae Dictyotales
Polyphysaceae
Ulvaceae
Dictyotaceae
Phaeophyceae Dictyotales
Dictyotaceae
Fucales
Magnolophyta Alimastales
!
52
Appendix 1 (Con’td)
(Lamouroux) Womersley ex Oliveira
Ni-NI-_%"5'D>'8-;%'C'd>'r;<;%
(Turner) J. Agardh
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
Avrainvillea
Boergesenia
Boodlea
Boodleopsis
Boodleopsis
Bryopsis
Bryopsis
Bryopsis
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Caulerpa
Chlorodesmis
Chlorodesmis
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Cladophora
Cladophoropsis
Cladophoropsis
Codium
wrightii
lacerata
forbesii
composita
pusilla
#$%!
pennata
plumosa
#$%!
cupressoides
diligulata
filicoides
nummularia
racemosa
sedoides
serrulata
sertularioides
taxifolia
tongaensis
fastigiata
hildebrandtii
dotyana
feredayoides
goweri
prehendens
rupestris
vagabunda
#$%!
herpestica
vaucheriaeformis
arabicum
!! !! "! !! !!
!! !! "! !! !!
!!
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"!
!! !! !! !! !! !! !! !! !! "! !!
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!! !! !! !! !! !!
!!
!!
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!! !! !! !! !! !! !!
!! !! !! !! "! !! !!
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!!
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"! "!
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"! "!
!! !!
"! !!
"!
"! !! !! !! !!
!!
"!
!!
"! !!
!! !! !! !! !!
"!
"!
"!
"! !!
!!
"!
"!
"! !!
!!
!!
"! !!
"!
!! !!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
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!!
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!!
!!
!!
!!
!!
!!
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!!
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!!
!!
"! !!
!!
!!
!!
!!
!!
!!
"!
!!
"! !!
"! !!
!!
!!
!!
!!
!!
!!
!!
!!
"! !!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
!!
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"!
"!
!!
"!
"!
53
!! !! !! "! !!
!! !! !! !! !!
"!
"!
!! !! !! !! !!
Appendix 2 (Con’td)
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
Codium
Dictyosphaeria
Dictyosphaeria
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Halimeda
Microdictyon
Neomeris
Parvocaulis
Phyllodictyon
Rhipidosiphon
Rhipilia
Rhipiliella
Rhipiliopsis
Tydemania
Ulva
Valonia
Valonia
Valonia
Valoniopsis
Dictyopteris
Dictyota
Dictyota
Dictyota
Dictyota
Dictyota
Dictyota
Lobophora
Padina
geppiorum
cavernosa
versluysii
cuneata
distorta
gracilis
micronesica
minima
opuntia
taenicola
velasquezii
xishaensis
#$%"
okamurae
annulata
parvulus
anastomosans
javensis
crassa
verticillata
gracilis
expeditionis
flexuosa
aegagropila
fastigiata
ventricosa
pachynema
repens
bartayresiana
ceylanica
friabilis
grossedentata
humifusa
#$%!"
variegata
okinawaensis
!"
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54
Genus
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Appendix 2 (Con’td)
species
Turbinaria
Acanthophora
Actinotrichia
Actinotrichia
Amphiroa
Amphiroa
Amphiroa
Amphiroa
Amphiroa
Asparagopsis
Asteromenia
Botryocladia
Botryocladia
Caulacanthus
Centroceras
Centroceras
Ceramium
Ceramium
Chamaebotrys
Champia
Champia
Chondria
Chondria
Chondria
Chondria
Chondrophycus
Coelothrix
Corallophila
Corynocystis
Crouania
Cryptonemia
Dasya
Dasya
Dasya
Dictyurus
Dipterosiphonia
ornata
pacifica
fragilis
#$%!
foliacea
fragilissima
rigida
tribulus
sp.
taxiformis
anastomosans
skottsbergii
tenuissima
ustulatus
clavulatum
minutum
maryae
mazatlanense
boergesenii
compressa
parvula
arcuata
bullata
ryukyuensis
simpliciuscula
succisus
irregularis
apiculata
prostrata
minutissima
umbraticola
anastomosans
baillouviana
palmatifida
purpurascens
dendritica
1 2 3 4
!! !! !!
"!
"! !! "!
"!
!! !! !!
"!
"!
!! !! !!
!! !! !!
5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 27 28
"! "!
!! !! "! "! !! !! "!
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!! !! !! !! !!
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"! "!
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!!
!! !! !! !! "! !! !!
!! !! "! !! !! !! !!
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!!
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55
Genus
Appendix 2 (Con’td)
species
1 2 3 4
5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 27 28
jungermannioides
bamleri
kotschyanum
proliferum
#$!
corallicola
fragilis
#$!
erubescens
#$!
heterostroma
melanesiensis
#$!
brassica-florida
frutescens
laccadivicum
apiculata
ferulacea
adhaerens
parvipapillata
cf. boergesenii
inamoena
delicatula
sertularioides
hornemannii
caespitosa
caloglossoides
#$%"!
#$%&!
#$%'!
#$%(!
orientalis
barodensis
ptychoides
#$%!
glomerulata
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56
Genus
Leveillea
Lithophyllum
Lithophyllum
Lithothamnion
Lithothamnion
Lomentaria
Martensia
Martensia
Mesophyllum
Mesophyllum
Myriogramme
Myriogramme
Myriogramme
Neogoniolithon
Neogoniolithon
Neogoniolithon
Neosiphonia
Neosiphonia
Nitophyllum
Palisada
Peyssonnelia
Peyssonnelia
Polysiphonia
Polysiphonia
Portieria
Pterocladiella
Pterocladiella
Rhodymenia
Rhodymenia
Rhodymenia
Rhodymenia
Seirospora
Spirocladia
Sporolithon
Thuretia
Tolypiocladia
Appendix 2 (Con’td)
Genus
species
1 2 3 4
Wrangelia
Syringodium
Thalassia
sp. inedit
isoetifolium
hemprichii
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5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 27 28
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57
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
Coelothrix irregularis
Halymenia durvillei!
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 diligulata
Caulerpa racemosa var. peltata
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
Cladophoropsis vaucheriaeformis
Bryopsis pennata!
Chlorodesmis fastigiata!
Phyllodictyon anastomosans!!
Microdictyon umbilicatum
Cladophora feredayoides!
!
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
lydiane mattio