RESEARCH ATOLL BULLETIN NO. 369
CHAPTER 5 SPECIES COMPOSITION, DISTRIBUTION AND ABUNDANCE OF ALGAE AND SEAGRASSES OF THE SEYCHELLES ISLANDS BY
A. A. KALUGINA-GUTNIK, L. P. PERESTENKO AND T. V. TITLYANOVA
ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. June 1992
CHAPTER 5 SPEC= COMPOSITION, DISTRIBUTION AND ABUNDANCE OF ALGAE AND SEAGRASSES OF THE SEYCHELLES ISLANDS
A.A. Kalugina-~utnik*, LP. ~erestenko** T.V. Titlyanova and
***
INTRODUCTION
Marine algae and seagrasses of the Republic of t h e Seychelles remain poorly studied. Macrophytic algae reported for the area (about 120 species) were collected between 1899-1990 by J. Stanley Gardiner during'the Sealark Expedition to the Indian Ocean (Gepp and Gepp 1909; 1911, Weber van Bosse 1913a, 1913b). Aleem (1984) reported 9 species of seagrasses and 22 algal species for the macrophyte communities of MahB, Latam, Aldabra, Comoro, Farquhar and Amirantes Islands. Subsequently, 5 seagrasses and 33 algal species were recorded (Titlyanova and Butorin 1987) for MahB and Wetivy Islands, half of these representing new records. There are no data on either algae or seagrasses for 12 of the Seychelles island groups. The present study of these island groups focused on: (1) macrophytic species composition; (2) distribution of algae and seagrasses at different depths for typical ecotopes; and (3) t h e structure of benthic plant communities (phytocoenoses), including determination of biomass for macroalgae and seagrasses.
METHODS AND MATERIALS
Studies were conducted at Wetivy Atoll, Desroches, African Banks, Providence, Farquhar Atoll, Aldabra Atoll, St. Joseph Atoll, Cosmoledo Atoll, Astove Atoll, MahB, Praslin and La Digue Islands. Quantitative and qualitative samples were collected using transects and square quadrats from the upper intertidal zone to a depth of 30-50 m, using a combination of snorkle and SCUBA diving techniques. In each intertidal horizon, four 25 x 25 cm quadrats were selected and sampled in representative habitats with vegetation coverage averaging 10-15%. In the subtidal zone, one 50 x 50 cm quadrat was harvested at each station. A total of 687 samples (479 quantitative harvest samples) were taken at 257 stations. Vegetation was described for 28 intertidal and 37 subtidal transects. About 4000 plant specimens were collected and duplicates are deposited in the herbaria of all three authors' institutions. Altogether, 327 algal species and 8 seagrass species were documented for the 12 island systems (Table 1). Intertidal zones were surveyed during periods of low water at which time type of substratum, bottom relief, vegetation coverage and the width of vegetation belts were determined. Laboratory Institute of South Seas Biology, USSR Academy of Sciences of the Ukraine, Sevastopol, USSR
** Botanical Institute, USSR Academy of Sciences, Leningrad, USSR *** Institute of Marine Biology, Far East Branch, USSR Academy of Sciences, Vladivostok,
690032, USSR
processing included determination of the population density (individuals-m-2),wet biomass (gmm2), plant height and species composition. Macroalgae were identified using the following works: Bt#~rgesen (1914, 1915, 1916, 1917, 1918, 1919, 1920, 1940, 1941, 1942, 1943, 1944, 1945, 1946, 1948, 1949,1950,1951,1952,1953,1954,1957), Cordero (1977), Colinvaux (1968), Dawson (1954), Durairatham (1%1), Egerod (1975), Hartog (1970), Hillis (1959, 1980), Kraft (1986), Misra (1%6), Olsen-Stojkovich (1985), Pham-Hoang Ho (1967, 1969), Tanaka and Pham-Hoang H o (1962), Taylor (1%0) and Valet (1%9). The structure of phytocoenosis (community associations) was analyzed with the use of the Shannon species diversity index (Wilhm 1%8),
where: H - species diversity of a sample; Wi - biomass of the i-th species sampled (gm-2); W - total biomass of the sample (g.m-2) and s - number of species in the sample. A t H = 0 - 0.8 the phytocoenosis structure w s considered to be monodominant; at H = 0.9 - 1.5 - oligodominant and a at H > 1.5 - polydominant.
The Seychelles islands surveyed can be divided into three groups by their structure and the type of bottom vegetation. The first group includes: CC)etivy, Desroches, African Banks and Providence; the second: Farquhar, Aldabra, St. Joseph and D'Arros, Cosmoledo and Astove; and the third: Mahe, Praslin and La Digue. CtJetivy, Desroches, African Banks and Providence are calcareous and fringed with barrier reefs. Farquhar, Aldabra, St. Joseph and D'Arros, Cosmoledo and Astove are atolls with shallow sand lagoons and sand banks exposed at low waters. On their ocean sides, these atolls are fringed with a broad (250-1000 m) reef-flat, pre-slope platform and reef slope. The intertidal zone of these islands is mostly wide (100-1000 m), sloping and uniform in structure. The upper and the middle horizons of the intertidal zone consist of sand. The lower horizon is sandy with pits, or built of dead consolidated corals covered with sand. The subtidal zone has different relief and substrata: the upper reef edge is sandy. The reef edge is built of coral plates and blocks above a steep sandy slope covered with dead corals with live coral colonies. The upper intertidal horizon is usually devoid of vegetation; a wide belt of seagrasses spreads from the middle horizon of the intertidal zone down t o depths of 15-20 m. Greater depths are occupied by Halimeda communities. CC)etivy is the most thoroughly studied island in the first group. Mahe, Praslin and La Digue, the third group, are high granitic islands with numerous inlets and are fringed by narrow reefs. Vegetation of the carbonate islands (groups 1 and 2) is dominated by seagrasses, and red and green algae. While on these granitic islands, brown algae (Phaeophyta), mostly Sargassum, predominate in plant communities along with seagrasses. RESULTS AND DISCUSSION Chtivy Island The benthic vegetation was surveyed along nearly the entire coastline (Fig. 1) and showed uneven distributional patterns. At the northern point of the island, the intertidal zone is narrow (510 m in width), sandy and devoid of vegetation. Macrophytic growth begins at a depth of 2-3 m, a
distance of 100-150 m from the shoreline. The southernmost intertidal zone is broad (500-1000 m in width) with numerous intertidal pools at low waters. Along the eastern and western coasts, the intertidal zone does not exceed 500 m in width. Settlements of Enteromorpha kylinii sometimes s occur on individual coral blocks in the upper intertidal horizon. The middle horizon i occupied by the seagrasses Syringodium isoetifolium, Halodule uninervis and Thalassodendron ciliatum (Fig. 2, Table 2) characterizing the major associations (or phytocoenoses, characterized by dominant species). The species composition of these associations is limited, including among the dominant species, Jania adhamens, @pnea pannosa, Diciyosphaeria setchellii, BoaIlea composita and Halimeda gracilis. The lower intertidal horizon is dominated by Thalassodendron ciliatum. Halimeda gracilis, H. stuposa, H. micronesica, Dictyurus purpurascens and Laurencia sp. are subdominants in all associations dominated by T. ciliatum. Dense mats of Gelidiella acmosa, Laurencia parvipapillata, L. paniculata and Halimeda opuntia develop at the stem bases and on the rhizomes of T. ciliatum, while the stems are populated by the algal epiphytes Haloplegma dupereyi, Gelidiella myrioclada, Champia parvula, Ceramium fastigiatum, Lophosiphonia villum, Poiysiphonia sp. and Centrocmas apiculatum. Among t h e o t h e r organisms noted were Dictyosphaeria cavernosa, D. setchellii, Caulerpa sertularioides, Udotea argentea, U. orientalis, Laurencia obtusa, Dictyurus purpurascens, Jania ungulata and Valonia aegagropila. Scattered individuals of Lobophora variegata and lkrbinaria ornata occur on the reef edge. 'helve associations were distinguished in the intertidal zone (Table 2), ranging in structure from mono- to oligo-dominant (i.e., H = 0 - 0.8 and 0.9 - 1.6, respectively) with well developed plant coverages ranging from 70-100%. In the sublittoral zone, associations of Thalassodendron ciliatum Halimeda dominate the sandy reef slope among coral debris (Fig. 2). Biomass of these two dominant species contributes 80-100% to the association (Fig. I), which includes almost all the species recorded for the lower intertidal horizon. In addition, Rhipilia tomentosa, Heterosiphonia sp., Lophocladia trichoclados, Grimhsia subcylindrica, Chlorodesmis comosa, Kh1l)menia sp., Coelarthm boergesenii, Tricleocarpa oblongata, Liagora divaricata and Botlyocladia skottsbergii were also found. The lower border of the association extends to a depth of 15-20 m with isolated specimens of T. ciliatum occuring to depths of 30-33 m. One of the most prominent species of subtidal coenoses is Halimeda opuntia. Off the southwestern border of the island, phytocoenoses of Halimeda appear at 1-m deep on the sand plateau and extend down to depths of 30-50 m and over vast areas. The biomass of Halimeda increases from 429 to 5078 gmv2at depths between 1-5 m (Table 2; Fig. 1; Stations 15-20). Species composition of the phytocoenosis is limited, with the associated species being T. ciliatum, Lobophora variegata, Caulerpa cupressoides, Haloplegma duperreyi, Dictyurus pupurascens and Hetmosiphonia sp. At depths of 30-50 m, dominants wereAvrainvillea amadelpha f. submersa, Halimeda copiosa, Caulerpa sp., Dasya sp., Struvea elegans, Antithamnion sp., Chtysymenia pynYormis, Peyssonnelia sp. and Anadyomene wrightii. The associations are characterized by a monodominant structure with sparse and low-statured vegetation coverage (Table 2). The H index of species diversity ranges mainly from 0 to 0.7, but in some phytocoenoses it exceeds 1.0 (Table 2). The distributional pattern of common species of algae and seagrasses around Cetivy is shown in Figure 3. The lower intertidal horizon and the upper sublittoral horizon down to a de th of 7-10 m (Fig. 4) show the greatest biomass of macrophytes (mean of 30% k 393 S.D. gnm-e2). Ranking second in biomass (1597 k 272 g-m-2)are phytocoenoses at 10-20 m in depth along the eastern coast and in the lower intertidal zone of the southern end of the island. Closest to the shore, within the middle horizon of the intertidal zone, the mean biomass of macrophytes is 878 k 306 g-m-2. The lowest biomass (557 k 157 w s found at greater depths. The total area of algal and seagrass a growths around Coetivy Island i approximately 13,190 ha, with an estimated total standing stock of s
1%,316 t wet weight. On average, 1 ha supports 15 t phytomass. Altogether, 113 plant species were found at Coetivy, 4 of which are seagrasses. The 109 algal species include 58 Rhodophyta, 46 Chlorophyta, 4 Phaeophyta and 1Cyanophyta (Table 1).
Desmhes Island
The marine benthic vegetation of the island was studied in considerable detail (Figs. 5-7). The upper intertidal zone is similar to that of Coetivy Island. An association of Thalassia hemprichii with Potolithon gardineri forms in the middle horizon. The phytocoenosis is oligodominant in structure, vegetation coverage averages 80-100%. Population density and biomass do not exceed 1536 individuals-m-2and 2471 respectively (Table 2). Among the associated species, Udotea orientalis, Caulerpa cupressoides, C. semlata, Dictyosphaeria cavernosa and Jania capillacea are common. In the lower intertidal horizon, Thalassodendron ciliatum forms three associations (characterized by their dominant species): Thalassodendron ciliatum T. hemprichii - Halimeda micronesica, T. ciliatum - Halimeda opuntia and T. ciliatum alone. Biomass of T. ciliaturn accounts for 60-100% of the total biomass of the three phytocoenoses (Fig. 5). Population density and and biomass are high, up to 2600 indi~idua1s.m.~ 5095 g.m-2 (Table 2), respectively. The stalks of Thalassodendron ciliafum are densely overgrown with epiphytes. The most common associated species are Laurencia parvipapillata, L. papillosa, Dasya mollis, Dictyurus purpurascens, Smtvea anastomosans, Haloplegma duperreyi, Hypnea esperi, H. spinella, Gelidiella lubrica, G. myrioclada, Hetmsiphonia crispella, Jania capillacea, J. ungulata, Dictyosphaeria cavemosa, D. vmslyusii, Caulerpa semlata, C. cupressoides, Valonia aegagropila, V: fastigiata and Halimeda stuposa, with Laurencia, Jania and Halimeda forming mats.
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In terms of depth of habitat, community structure, species composition and the main quantitative characteristics, subtidal associations of Thalassodendron ciliatum-Halimeda at Desroches Island do not differ appreciably from corresponding associations at metivy Island (Table 2). In addition to species that were also common on CC)etivy Island, we found Neomeris bilimbata, Caulerpa mericana, Halymenia sp., Liagora pennata, Galaxaura maqinata, llkleocarpa oblongata, G. rudis, Haliptilon subulafum and Lomentaria mauritiana. In the depth range between 15-20 and 35-38 m, a steep cliff partly fringing the island is covered with sparse algal growths. Below the cliff, a Halimeda phytocoenosis extends to a depth of 50 m (Table 2), with high species diversity of associated algae, most of them being common in the upper subtidal and lower intertidal zones. Typical of these deep waters were Avrainvillea amadelpha f. submersa, Halimeda copiosa, H. tuna, Dudresnaya sp., Caloglossa sp., IIypoglossum sp. and Caulerpa mericana. Altogether, 132 plant species, including 3 seagrasses and 129 algal species (72 red algae, 50 green algae, 5 brown algae and 2 blue-green algae) were collected near Desroches Island. In general, the bottom vegetation of Desroches Island is characterized by relatively high quantitative indices (Fig. 8). The total area covered by benthic vegetation is about 711 ha and supports about 16,581 t wet plant biomass. African Banks
The South Island's lower intertidal and upper subtidal zones (Figs. 9-11) are characterized by associations of Thalassia hemprichii Thalassodendron ciliatum - Microdictyon montagnei. Thalassodendron ciliatum is responsible for 40-100% of the total biomass of all phytocoenoses (Fig. 9). In the subtidal zone, this association is replaced by the T. ciliafum association which extends along the slope of the reef down to a depth of 20-25 m. The population density and biomass in the T. ciliatum association is significantly lower than on Cetivy and Desroches Islands (Table 2). Common associated species were Halimeda micronesica, H. sfuposa, H. pacilis, H. macroloba, H. opuntia,
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Dictyosphaeria cavernosa, D. verluysii, Rhipilia tomentosa, Anadyomene plicata, Struvea anastomosans, Tricleocapa oblongata, Haloplegma duperreyi and Lobophora variegata. The deepwater association of Halimeda sp. (25-37 m) is characterized by a rich species diversity of associated algae similar to that near Desroches Island. Most characteristic were Halimeda stuposa, H. micronesica, Caulerpa cupressoides,Avrainvillea amadelpha f. submersa and Udotea orientalis. Several species were found only at the 37 m depth (Station 200): Halophila stipulacea, Caulerpa sp., Boergesenia forbesii, Boodleopsis pusilla, Anotrichium tenuis, Galaxaura rudis and Chrysymenia p r o m i s . Zjdemania gardinen', Halophila stipulacea and Anadyomene plicata were found only near South Island. Altogether, 84 plant species were collected near South Island: 3 seagrasses and 81 algae (47 red algae, 31 green algae, 3 brown algae and 3 blue-green algae).
Providence Atoll
The interdidal zone of the southern coast of Cerf Island (Fig. 12,13) is sandy, almost devoid of vegetation, with scattered coral debris covered by Enteromopha clathrata. The distribution of plant communities in the subtidal zone was typical of the other islands of this group. At a depth of 20-23 m, the Thalassodendron ciliafum association is replaced by communities dominated by Halimeda gracilis extending the down to 32 m (Table 2). Thalassodendron ciliafum associates include Microdictyon montagnei, Herposiphonia secunda f. tenella, Dictyurus purpurascens, Griffithsia subcylindrica and Cottoniella arcuata. The Halimeda communities included Halimeda micronesica, Micrdiciyon montagnei, Valonia fastigiata, Caulerpa brachypus, Tricleocapa oblongata, Galaxaura rudis and other minor species. Two seagrass species and 58 algal species (40 red algae, 16 green algae and 2 brown algae) were collected near Cerf Island. The intertidal vegetation of islands of the first and second groups is similar in structure and distribution. The upper, and sometimes mid-tidal horizons, are devoid of plant coverage. The middle a n d lower i n t e r t i d a l horizons s u p p o r t associations of Thalassia hemprichii + Thalassdendron ciliafum. In the lower intertidal and subtidal horizons, this association is replaced by communities of T. ciliafum and green algae, which become more frequent in all phytocoenoses compared with those of the islands of the first group.
Farquhar Atoll
At Farquhar Atoll, Thalassia hemprichii Thalassodendron ciliatum associations of North Island, South Island, Bird Island and the lagoon (Figs. 14-17) had moderate population densities and biomasses (Fig. 15, Table 2). Plant coverage near South Island ranges from 5 to 40% and 100% near North Island. Macrophyte associations include Boodlea struveoides, Valonia aegagropila, Cladophoropsis sundanensis, Diciyosphaeria cavemosa and D. verluysii. The green algae Cladophora socialis and B. sfruveoides form monodominant intertidal phytocoenoses. Boodlea sfruveoides, Caulerpa cupressoides and Valonia aegagropila are included as subdominant species in both the Thalassdendron ciliafum - B. struveoides and T. ciliatum - K aegagropila - C. cupressoides (Table 2) plant associations. The phytocoenoses also include Microdictyon okamurai, Dictyosphaeria cavernosa, Cladophoropsis sundanensis, Jania adhaerens, J. ungulata, Gelidiella pannosa, G. rnyrioclada, G. Iubrica, Lophosiphonia villum, Ceramium fastigiatum and Hypnea esperi. Haloplegma duperreyi, Liagora ceranoides, Laurencia papillosa, L. obfusa and Lobophora variegata are epiphytic on the stems of T.ciliafum at the reef edge. The subtidal zone vegetation is poorly developed on the reef slope, because abundant coral populations extend almost to the reef-front. The following phytocoenoses were distinguished at depths of 3-17 m: Caulerpa cupressoides + Boodlea struveoides, Halimeda opuntia + C. cupressoides
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and Thalassodendron ciliatum - Halimeda. With increasing depth these were replaced by phytocoenoses of M. okamurai B. struveoides, Microdicfyon and Udotea + Caulerpa (Table 2). Plant coverage ranges from 5 to 90%, but more often 5-30%. The height of vegetative cover is not great (1-2, seldom more than 1cm)and the biomass is small (261-520 g.m-2).
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Species richness of algae in the subtidal zone is low, and does not change with depth. Such species as M. okamurai, C, cupressoides, B. struveoides, H. opuntia and Struvea anastomosans, are widespread at 35 m. The epiphytes Haloplegma dupeweyi, Jania ungulata, Gelidiella myrioclada, G. lubrica, Gnmhsia subcylindiica, G. metcalfii, Heterosiphonia &pella and Lophocladia nichoclados inhabit the stems of T.ciliatum. Turbinaria omata and Lobophora variegata also occur but are rare. Sargassum ilicifolium was found between 1-4 m deep on a sunken vessel at the southwestern end of the atoll representing the only low island location where the genus occurred. Ulva rigzda, Boodlea struveoides, Dictyosphaena cavemosa, Struvea anastomosans, Cladophora laetevirens, Dictyopteris delicatula, Padina sp., Sphacelaria rigidula, Gelidium pusillum, Jania ungulata, Iiypnea espen and H pannosa occurred with the Sargassum. The vegetation of the lagoon is considerably richer than that of the outer reef slope. It is characterized by the Thalassodendron - Halimeda phytocoenosis (Figs. 14-17; Table 2). The central part of the lagoon is covered with sand and corals and is devoid of vegetation. In the southern region of the lagoon, sites with Thalassodendron ciliatum phytocoenoses are rare. The belt of Thalassia hemprichu T. ciliatum, including species of Laurencia and Halimeda, runs along the northwestern reef edge. Northern and western parts of the lagoon are occupied by dense populations of T. ciliatum, which is responsible for 60-95% of the plant biomass. Halimeda opuntia, Caulerpa cupressoides and Microdicfyon okamurai are community subdominants. Halimeda gracilis, Caulerpa serrulata, C. peltata, C. racemosa, C. msricana, Valonia aegagropila, M l e a struveoides, Laurencia obtusa, L. papillosa, Iiypnea esperi, Jania ungulata, J. capillacea, Centroceras apiculatum, Ceramium fastigiatum and Gelidiella lubrica are of frequent occurrence. At Farquhar Atoll, an estimated 195,139 t of benthic phytomass occupies an area of 10,085 ha. Altogether near Farquhar Atoll, 2 seagrasses and 115 algal species (including 66 red, 39 green, 7 brown and 3 blue-green species) were found.
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Aldabra Atoll
The structure of the Aldabra Atoll differs considerably from some of the other island groups, since its coasts are built primarily of dead consolidated corals and are steeply undercut and overhang the water. Surveys were conducted mostly on the western coast of the atoll near Bua Passage in the lagoon and on the outer part of the reef, as well as the southeastern parts of the atoll (Figs. 18-20). A polydominant association of Thalassodendron ciliatum - Thalassia hemprichu - Halimeda forms in the lower intertidal horizon (Table 2). Thalassodendron ciliatum is responsible for 40-90% of the community biomass, while T. hempnchii, Halimeda opuntia, Caulerpa cupressoides and Laurencia combine t o contribute 60-10%. In addition to the above species, the phytocoenosis includes Laurencia, Hypnea and Jania species forming dense mats. The stems of T. ciliatum are overgrown with epiphytes, including Ulva rigida, Hypnea esperi, Dasya mollis, Dictyuruspurpurascens, Heterosiphonia sp., and occasionally Lobophora variegata and Synngodium isoetifolium are present. At the lower border of the intertidal zone, the association of Thalassodendron - Thalassia - Halimeda is replaced by a T. ciliatum association which forms a belt 400-500 m in width along the entire western coast. On the reef slope, T. ciliatum populations disappear at 10 m in depth on a sandy bottom with abundant dead coral debris. The most common associated species are Ulva rigida, Halimeda opuntia, H. tuna, H. gracilis, H. micronesica, Lobophora vanegata, Turbinaria ornata, Spyridia filamentosa, Acanthophora spicifera, Dasya mollis, Caloglossa adnata and species of
Cmamium, M n e a and Gelidiella. Wurdemannia miniata, Gelidiella myrioclada and Ulva rigida grow on the stems of T. ciliatum. Depths t o 12-35 m are occupied by simple monodominant phytocoenoses of Halimeda. These phytocoenoses include a considerable number of species similar to that of the T. ciliatum association. Additionally, Chlorodesmis comosa, Avrainvillea amadefpha f. submmsa, Acetabularia parvula, A. clavata, Boergeseniaforbesii and Chondriapolyrhiza occur here.
The lagoon is connected with the ocean by the Bua Passage (depth of 5-6 m). The passage bottom is swept by strong currents and is hard and sandy with settlements of black gorgonians. Often the dead coral debris is populated by extensive growths of Thalassodendron ciliatum (Station 93). Thalassodendron leaves and stems are devoid of epiphytes, probably because of the strong current in the passage. Seagrass growths spread into the lagoon and occupy a zone 600-800 m from the shore where the depth decreases sharply. On shallow intertidal areas of the lagoon, vegetative coverage is composed of numerous assemblages, the most common of which is a phytocoenosis dominated by S ' i a jilamentosa. A phytocoenosis of Thalassia - Halimeda occupies the area near the coast along the northern part of the lagoon (Fig. 18; St. 90; Table 2). Opposite the passage, phytocoenoses of Turbinaria - Halimeda (St. 91) and Thalassodendron - Laurencia - Halimeda (St. 89) are present. Plant communities of the lagoon have oligo- to polydominant structures, mosaic patterns of the phytocoenoses and rich macrophyte compositions. Near Aldabra Atoll, 4 seagrass species and 119 algal species (i.e., 73 Rhodophyta, 40 Chlorophyta, 4 Phaeophyta, and 2 Cyanophyta) were collected.
Cosmoledo Atoll
The bottom vegetation of the reef slope and lagoons was surveyed at depths from 4 to 42 m near Wizard and Pagoda islands (Figs. 21,22). The vegetative coverage at 4-20 m depths is poorly developed and consists of separate groups with dominants consisting of species of Laurencia, Turbinaria, Galaraura, Caulerpa and Boodlea. Below 20 m, Halimeda, Microdic&w and Avrainvillea species predominate (Table 2). Thalassodendron ciliatum, Microdictyon and Avrainvillea species predominate (Table 2). Thalassodendronciliatum phytocoenoses, common in the majority of the atolls surveyed, were not found in the subtidal zone. Most of the species found at this coastal site occur at all depths examined and do not reveal any definite regularity in vertical distribution. The following species were found rarely or for the first time: Neomeris vanbosseae, Caulepa webbiana, C. sp., Acetabularia pusilla, Mesophyllum mesomorphum, Hydrolithon breviclavium, Wrangelia argus, Cmamium huysmansii, Sphacelaria tribuloides and Phaeophila dendroides. The lagoon contains a rich vegetative cover. The vast sandy shallow water areas between the islands Wizard and Polit and the coastal zone of Wizard are occupied by an association dominated by Halodule uninervis. An association of Thalassia Qnodocea is situated somewhat farther from the shore. An association dominated by Thalassodendron ciliatum - Laurencia was found between 0.45.5 m in depth. The central and western parts of the lagoon are occupied by an association of T. ciliaturn - Halimeda opuntia (Table 2). The coenobionts Laurencia papillosa, Caulerpa racemosa, C. cupressoides, C. sp., Microdictpn okamurai and Chaetomorpha capillare are typical members of the association.
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The species composition of the lagoon algae is uniform and includes mostly green algae. In addition t o the above mentioned species, Boergesenia forbesii, Dictyosphaeria cavernosa, NeomeTis annulata, Lobophora variegata and Haloplegma duperreyi were found here. Cosmoledo Atoll contained 4 seagrass species and 106 algal species, including 60 red, 42 green and 6 brown algae.
St. Joseph Atoll
The southern coast of D'Arros and Resourse islands and the eastern, northeastern and southern coasts of St. Joseph Island were surveyed. The distribution of seagrass communities over the intertidal zone of D'Arros and St. Joseph islands is typical of the other systems (Figs. 23-25; Table 2). A phytocoenosis dominated by Thalassodendron - Thalassia is replaced by a phytocoenosis of Cladophoropsis sundunensis+ Valonia fastgiata in the lower intertidal horizon near D'Arros island. The dominant species are all mat formers. Boodlea smtveoides, Dictyosphaeria verluysii, Microdictyon montagnei Jania capillacea and Gelidiella pannosa also occur here. The reef edge is exposed to a heavy wave action and is devoid of vegetation. Separate populations of Halimeda gracilis and Botryocladia skonsbergii occur at depths of 3-5 m. Deeper substrata are covered by crustose algae, such as Fosliella, Peyssonnelia and Neogonwlithon, and by corals.
Resourse Island
Two communities dominated by Thalassodendron - Microdictyon or Thalassodendron form in the lower intertidal and upper subtidal zones of Resourse island. Halimeda communities occupy depths of 7-37 m (Table 2). Thalassia hemprichii dominates the areas exposed to air at low waters. In the lagoon of St. Joseph Island, T. ciliatum tends to predominate with increasing depth. Microdictyon okamurai, M. montagnei, Boodlea struveoides, Dicfyosphaeria cavernosa, D. verluysii, Valonia fastigiata, Udotea orientalis, Halimeda stuposa, H. micronesica and H, opuntia dwell among the Thalassodendron. Caulerpa serrulata and C. cupressoides inhabit bottoms of sandy depressions. Phytocoenoses of T. ciliatum with typical species composition were found on parts of the northeastern coast of the subtidal zone of St. Joseph Island (Fig. 25). Halimeda communities with rich species composition of associated algae dominate on the reef slope.
Cladophoropsis sundanensis, Boodlea struveoides and Valonia aegagropila form mats along the reef edge. Populations of Turbinaria omata, Lobophora variegata, Dictyosphaeria cavemosa, D. verluysii, Dictyuruspurpurascens, Rhipilia tomentosa and Udotea argentea occur at depths of 3-5 m. Deeper to 35-38 m, rubble and blocks are covered with dense low growing populations of numerous red algae such as Gelidiella myrioclada, Gelidium pusillum, Gelidiopsis scoparia, Poiysiphonia sp., Lophosiphonia villum, Herposiphonia secunda f. tenella, Heterosiphonia crispella, Ceramium fastgiatum, Centroceras apiculatum, M n e a spinella and Laurencia sp. The green algae Microdictyon okamurai, Struvea anastomosans, Caulerpa sp., Halimeda micronesica, H. stuposa, Valoniopsis pachynema and Valonia aegagropila are the most frequent, while Codium tomentosum, C. geppii, Cladophora laetevirens, Lophosiphonia reptabunda, Lomentaria mauritiana, L. hawaiiana, Chondria d q h y l l a , Jania decussatodichotoma and Halophila stipulacea occur less frequently. Anadyomene plicata, Scinaia and Gracilaria cylindrica are occasional and sparcely distributed.
St. Joseph and D'Arros Islands contain 4 seagrasses and 120 algal species, including 69 Rhodophyta, 46 Chlorophyta, 4 Phaeophyta and 2 Cyanophyta.
Astove Atoll
As a rule, t h e intertidal zone of Astove Atoll is characterized by a mosaic structure of phytocoenoses and massive development of Laurencia. The intertidal and upper subtidal vegetative cover differs considerably from that of the other islands by its structure and species composition. There is a polydominant phytocoenosis of Thalassodendron ciliatum - Caulerpa sertularioides Acanthopora spicifera formed on the vast silt-sand shallow area along the coast (Fig. 27; Table 2). In the lower horizon, this is replaced by a phytocoenosis formed by T. ciliatum and several Laurencia
+
species, including L. decumbens (Fig. 26, Sts. 247-249; Table 2). Maximal quantitative development of macrophytes occurs a t the reef edge (Fig. 27), with associates consisting of Laurencia parvipapillata, L. cotymbosa, L. sp., Liagora sp., Hypnea spinella, Chaetomopha crassa, Halimeda opuntia, Dictyosphaeria cavernosa, D. verluysii, Valonia aegagropila and Boodlea struveoides.
Thalassodendron ciliatum - Halimeda hederacea - Laurencia sp. occurs from 0-3 m in depth. Deeper, coral populations appear, and between them oligodominant phytocoenoses of Halimeda copwsa + Caulerpa sp. are located on sandy areas containing dead coral debris (Table 2). Valonia aegagropila, Dictyosphaeria cavernosa, D, verluysii, Halimeda opuntia, H. m a , Awainvillea amadelpha f. submersa, Laurenciapatentisamea, L. cotymbosa and Gelidiella myrioclada are commonly associated species
(Fig. 28). The distinctive feature of the subtidal vegetation of Astove Island is the development of massive populations of Caulerpa sp. at 20-50 m depths. The benthic vegetation of Astove Island was not examined fully due to time constraints, so only 3 seagrasses and 61 algal species (32 red, 27 green and 2 brown algae) were found.
MahC and adjacent islands
Two phytocoenoses can be distinguished on the subtidal reef slope. A phytocoenosis of
The northern, eastern and northeastern coasts of the island and three small islands just outside Victoria Harbor (Figs. 29-31) were studied. The intertidal zone of the northeastern coast is open to the sea and narrow (60-80 m), with a slight slope. A phytocoenosis of Cladophoropsis sundanensis Centroceras clavulatum, along with the associated species Ulva rigzda, Cladophora socialis, Valonia aegagropila, Champiaparvula, Cladophorapatentiramea, Hypnea sp. and Jania sp., populate stone blocks in the middle intertidal horizon. An oligodominant association dominated by SargassumGracilaria multifirrcata occupies the lower intertidal horizon spreading almost to the reef-flat edge (Fig. 30: Table 2). Gracilaria crassa develops in shallow water habitats, deeper it is replaced by populations of G. multifurcata, G. sp., Amphiroa foliacea, Gelidiella acerosa, Jania capillacea and Cheilosporum spectabile, forming mats. The species composition of the associated algae is rather rich with Gelidiopsis scoparia, Hypnea valentiae, Laurencia papillosa, Sargassum cristaefolium, S. microcystum, Colpomenia sinuosa, Dictyopteris delicatula, along with species of Padina, Dictyota, Turbinaria and Sphacelaria being found here.
+
Sargassum cristaefolium and Turbinaria decuwens occur on the fringing reef-flat of the northeast and east coasts and dominate on stony reef slope sites lacking corals in the subtidal zone (Fig. 29; Table 2; Sts. 164-167) where Tricleocarpa oblongata is a subdominant. Associates include Hypnea valentiae, Ulva rigida, Caulerpa taxifolia, Cladophora patentiramea, Chlorodesmis comosa and Syringodium isoetifolium.
Seagrass communities develop southwards into the harbor with m o d o c e a rotundata inhabiting a narrow band along the shore, which is then replaced by Thalassodendron ciliatum occupying the entire area exposed at low waters. Associations of Sargassum cristaefolium occupy the southeastern coast's lower intertidal and upper subtidal zones (Figs. 29-31; Table 2). In the intertidal zone, the lower level of the Sargassum phytocoenosis includes Gracilaria crassa, Cheilosporum spectabile, Jania
longiartha, J. ungulata, Hypnea pannosa, H. nidulans, Laurencia parvipapillata, Amphiroa foliacea, Cladophoropsis sundanensis and Gelidiella acerosa, which form mats. Isolated populations of Dictyota indica, D. dichotoma, D. divaricata, Turbinaria omata, Padina sp. and Caulelpa racemosa
occur as well.
Sargassum phytocoenoses have mono- and oligodominant structures (H = 0-1, I), provide 100% cover and high biomass (Table 2). Minimal biomass was found in the upper intertidal zone while maximal biomass occurred near the reef edge (Fig, 30). The biomass of the various phytocoenoses
decreases with depth. At 8-10 m deep, Sargassum phytocoenoses are replaced by those dominated by Halimeda, which are distributed around the island in a form of belt, as well as Sargassum spp. Seagrass phytocoenoses, including Thalassia hemprichii, Thalassodendron ciliatum, Halodule uninervis and Halophila ovalis, develop on shallow sites in the middle intertidal horizon where they attain maximal biomass. Sargassum phytocoenoses of the subtidal zone include species such as Turbinaria decumens, Lobophora variegata, Dictyota indica, Padina sp. and W l e o c a p a oblongata.
St. Anne Island
The association of Sargassum polycysntm + Turbinaria omata occupies the middle intertidal horizon of the south part of the island (Fig. 29), whereas in the lower horizon it is replaced by an association of Sargassum cristaefolium. The lower level of this association is made up of dense mats of Amphiroa foliacea and Gelidiella acerosa. Thalassia hemprichii and Halodule uninervis grow along this coast in small shallow bays, while Halophila ovalis occupies more silty sites. The lower border of the Sargassum phytocoenosis extends to a depth of 0.8-2 m. Coral populations occupy the bottom deeper.
Cerf Island
Sargassum microcystum and S. cristaefolium dominate the lower intertidal and upper subtidal zones. The main coenobiont of this community is firbinaria ornata. The lower level is formed by Amphiroa foliacea, Cheilospomm spectabile, Jania capillacea, Haliptilon subulantm, Gelidiella acerosa and Halimeda opuntia. Populations of Padina sp., Dictyota sp. and Qmodocea semlata occur rarely and near the shore. Dense mats of Gelidiopsis scoparia cover dead coral blocks among Sargassum spp. in the upper subtidal zone and Cladophoropsis membranacea grows on the sandy bottom among the blocks. Below 3-4 m in depth, Sargassum spp. communities are replaced by populations of coral.
Anonyme Island
The marine vegetation of the north side of Anonyme Island, situated at the south end of Victoria Harbor, was investigated. Monodominant continuous phytocoenoses of Sargassum micraysntm + S. cristaefolium occupy sandy-stony bottoms at 0.5-1.2 m in depth (Fig. 29; Sts. 169-173; Table 2). The species composition of these phytocoenoses is limited and includes such species as Hypnea
nidulans, Jania capillacea, J. ungulata, Centroceras clavulatum, Leveillea jungermannioides, Sphacelaria rigidula, Gelidium pusillum, Chlorodesmis comosa, Acetabularia parvula, &yopsis pennata and Gelidiella acerosa. The north reef slope is exposed to wave action and consists of blocks with poorly developed vegetation (Fig. 12; Table 2; Sts. 173-175). Turbinaria ornata, Lobophora variegata, Dasya baillouviana and crustose algae occur rarely.
Near MahC and its adjacent islands, 7 seagrasses and 173 algal species were collected, including 86 reds, 53 greens, 27 browns and 7 blue-greens.
Praslln Island
The marine vegetation of the eastern and central areas of Grande Anse Bay and the southern and southwestern coasts of Praslin were investigated to a depth of 40 m (Figs. 32-34). In the middle intertidal horizon of Grande Anse Bay, an association of Gelidiella acerosa is formed on separate rocky blocks, lower it is replaced by Gracilaria crassa associations. Sandy bottom sites between rocky
blocks are overgrown by Sargassum ilicifolium. Phytocoenoses of the seagrasses Qmodocea serrulata + Syringodium isoetifolium Thalassodendron ciliatum, S. isoetifolium and Halodule uninervis are situated in the nearshore sandy intertidal zone. C a u l q a serrulata predominates in the association of C. serrulata + S. isoetifolium and is responsible for 60-70% of the biomass. From the lower intertidal border, to 1-1.2 deep, the T. ciliatum association inhabits sandy to stony substrata (Table m 2 Fig. 3 ) The width of the T. ciliatum belt reaches 800-1000m. Halimeda stuposa is typically found ; 4. in this association.
+
m), At shallow sites (0.3-0.5 the above seagrass belt is replaced by a phytocoenosis of Sargassum cristaefolium - Padina sp. The species composition of this association is reduced and includes Lobophora variegata, Dictyota indica and Chlorodesmis comosa. With increasing depth, populations of S.~taefolium- Padina sp. are replaced by multiple Sargassum assemblages that extend down to 8-10m in depth. Maximal cover and biomass are found between depths of 1 to 3 m (Table 2. The ) species composition of Sargassum-dominated phytocoenoses is diverse and includes Caulerpa cupressoides, C. serrulata, C. racemosa, C. sertularioides, C. mexicana, C, taxifolia, C. ambigua, Dictyosphaeria cavernosa, Enteromorpha clathrata, Chlorodesmis cosmosa, Cladophoropsis sundanensis, Struvea anastomosans, Dictyopteris polypodiodes, Dictyota indica, D. friabilis, D. divaricata, Lobophora variegata, Turbinaria conoides, Sphacelaria rigidula, S. tribuloides, Centroceras clavulatum, C. apiculatum, Gelidiopsis scopariu, Gelidiella acerosa, Champia parvula, Tolypiocladia glomerulata and Wurdemannia miniata. Below 9 m, the bottom is sandy with large calcium carbonate blocks and coral colonies. Hard substrata are overgrown with crustose and small algae such as Gelidiella lubrica, Ceramium fastigiatum, Lophosiphonia villum, Gelidium pusillum and Champia parvula. Among the larger macroalgae, Lobophora variegata and Laurencia species are the most frequent. At the 40 m depth, Chlorodesmis comosa, Cladophora laetevirens, Halimeda sp., Cladophoropsis sundanensis, Dictyota divaricata, Champia salicomioides, Dasya sp., Antithamnion herminien and Chondria sp. are common.
La Digue Island
The eastern intertidal zone is exposed to strong wave action and is narrow and devoid of vegetation, whereas the western coast is more protected from wave effects. The upper and middle intertidal horizons consist of a sandy beach. Algae overgrow sandy-stony reef-flat habitats. The oligodominant association of Thalassdendron ciliatum - Hypnea pannosa occurs near the shoreline (Table 2; St. 180;Fig. 34)and includes Zhrbinaria omata, which deeper forms a 15-20m wide belt. Sargassum turbinarioides dominates the middle of the reef-flat (Table 2. Distinctive phyto&oses ) develop at lower levels, consisting mainly of Hypnea nidulans and H. pannosa mats which provide up to 8-20% of the total biomass. Populations of Heterosiphonia sp. often occur along the edge of the reef-flat among the Sargassum phytocoenoses. In the subtidal zone, Sargassum phytocoenoses extend down to 8 m in depth. Deeper, down to 30-31m, there is a belt of crustose algae with mats of small red algae and separate patches of Lobophora variegata and Laurencia sp. The species composition of algae in this region is similar to that of deep waters off Praslin Island. Phytocoenoses of granitic islands tend to have oligodominant structures, considerable biomass and limited species composition of associated species. Sargassum communities have monodominant structure, great biomass and rich species composition, which all become reduced with depth. Near Praslin and La Digue Islands, 6 seagrass and 136 algal species, including 72 red, 35 green, 22 brown and 7 blue-green species were recorded.
CONCLUSIONS
The benthic vegetation of the Seychelles Islands shows several common characteristics, presumably related to similar geomorphological features of the coastal zones and the tropical oceanic location of the islands:
1. Rich species composition of algae at every island. 2. Wide range of vertical distribution of most the macrophyte species. 3. Domination of red (4544.2%) and green (25.5-42.4%) algal species and poor development of brown algae (2.5-16.1%) in the floras of the low carbonate islands. 4. A great number of genera are common to all islands surveyed: green algae - Caulerpa, Halimeda, Microdictyon, Boodlea, Anadyomene, Cladophoropsis, Avrainvillea, Udotea, Acetabularia, Valonia, Diclyosphaeria and Struvea; brown algae - Lobophora, Thrbinaria and Dictyopteri~; algae - Galaxaura, Gelidiella, Wurdemannia, Peyssonnelia, Jania, Melobesia, red Hjpnea, Champia, Haloplegma, Anotrichium, Chondria, Centroceras, Dasya, Heterosiphonia, Lophocladia, Herposiphonia a n d Laurencia; and t h e seagrasses - Thalassia and Thalassodendron. 5. Thalassodendron ciliaturn and Thalassia hemprichii dominate the overall vegetative cover of soft bottoms. In the middle and lower intertidal horizons, seagrasses occur in the following sequence: Halophila ovalis - Halodule uninervis - Thalassia hemprichii - Cymodocea s m l a t a - C. rotundata - Syringodium isoetifolium - T. ciliatum. Thalassodendronciliatum consistently forms a belt that dominates in the subtidal zone. 6. Thalassodendron ciliatum plays an important role in the stabilization of unconsolidated substrata in the coastal ecosystems of the atoll islands. 7. The benthic vegetation is characterized by relatively high standing stocks. In habitats with highly developed vegetative cover approaching loo%, the biomass reaches 2-4 kg.m-'. Maximal biomass was found w e r the depth range of 0.5-0.3 m, and averaged 4-8 kg-m-'. 8. Commercially exploitable stocks of macrophytes were found: mainly Thalassodendron ciliatum (biomass = 1-4 kg-m-') on atoll island sand and on the granitic islands, primarily species of Sargassum (biomass = 2-10 kgm-') and Gracilaria (biomass = 2-6 kg-m-'1.
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Table 1. Preliminary checklist of benthic marine algae and seagrasses collected near Seychelles Islands. M = Mahk, P = Praslin, Co = COetivy, F = Farquhar, D = Desroches, A = Aldabra, AB = African Banks, S = St. Joseph, Pd = Providence, Cs = Cosmoledo and As = Astove. CoF A D Island P MABS PdCsAs
Oscillatoriales Oscillatoriaceae 1.Oscillatoria margaritifera (Kutz.) Gom. 2. Oscillatoria miniata (Zanard.) Hauck 3. Lyngbya confervoides Ag. 4. Lyngbya majuscula (Dillw.) Ham. 5. Lyngbya martensiana Menegh. 6. Lyngbya sp. 7. Symploca hydnoides (Ham.) Kutz. Nostocales Rivulariaceae 8. Calothrir confervicola (Dillwyn) Ag. 9. Calothrir aeruginea (Kutz.) Thuret Nostocaceae 10. Calothrirparietina (Nlg. ex Kutz.) Thuret Nostocales 11.Homothamnium solutum Born. et Grunow
PMAB PM PMAB FADPMABS F A ' M F M S P
Co
Goniotrichales Goniotrichaceae 12. Chroodactylon omatum (C. Ag.) Basson (=Aster@ omata;Aster@ ramosa) 13. Stylonema akidii (Zanard.) Drew Bangiales Erythropeltidaceae 14. Etythrotrichia camea (Dillw.) J. Ag. 15. Eryhrocladia sp. Floridepphyceae Acrochaetiales Acrochaetiaceae 16.Acrochaetium occidentale B&g. 17.Acrochaetium robustum B@g. 18.Acrochaetium seriatum B@g. 19.Acrochaetium sp. 20. Chromastnrm crassipes (B&-g.)Papenf. (=Acrochaetium crassipes)
F
A
D
M
ABS
FADPMABS CoF A M ABS
\
Cs
Pd Cs
Taxon Nemaliales Liagoraceae 2 1 Liagora ceranoides Lam. . 22. Liagora divaricata Tseng 23. Liagora pinnata Haw. 24. Liagora sp. 1 25. Liagora sp.2 26. Liagora sp.3 Galaxauraceae 27. Scinaia complanata (Collins) Cotton 28. Galuxaura marginata (Ell. et Sol.) Lam. 29. Tricleocarpa oblongata (Ell. et Sol.) Huisman et Borowitzka (=Galaxaura oblongata) 30. Galaxaura mgosa (Ellis et Soland.) Lam. ( =Galaxaura rudis) 31. Galaxaura sp. Gelidiales Gelidiaceae 32. Gelidium crinale f. colymbosa (Kutz.) J. Feldm. et Hamel 33. Gelidium pusillum (Stackh.) Le Jolis 34. Pterocladia caloglossoides (Howe) Dam. Gelidiellaceae 35. Gelidiella acerosa (ForsskAl) J. Feldm. et Hamel 36. Gelidiella adnata Dam. 37. Gelidiella lubrica (Kutz.) J. Feldm. et Hamel 38. Gelidiella myrioclada J. Feldm. et Hamel 39. Gelidiella sanctamm J. Feldm. et Hamel 40. Gelidiellapannosa J. Feldm. et Hamel (=Gelidiella tenuissima) Bonnemaisoniales Bonnemaisoniaceae 41. "Falkenbergiahillebrandii" (N.B. - stage ofAsparagopsis) Corallinales Corallinaceae 42. Fosliella farinosa (Lam.) Howe 43. Neogoniolithon sp. 44. Pheophyllum confervicola (Kiitz.) Chamb. (=Mefobesia minutufa) 45. Hydrolithon breviclavium (Fosl.) Fosl. 46. Porolithon gardinen' (Fosl.) Fosl. 47. Spongitesreinboldii (W.V. Bosse et Fosl.) Penrose et Woelkerling
Island C o F A D P MABS
PdCsAs
F Co F Co
A D
S
S D CoF A P M Pd Pd Cs Pd M CoFADPM P P Co Co Co Co Co F F F F A A A D D P P P P M M M M AB AB S ABS
S
CoFADPM D
Cs Cs As Cs As Cs As
A
.Co F
D
P
M
Table 1. Continued. Taxon
48. Melobesia sp. 49. Mesophyllum mesomorphum (Fosl.) Adey 50. Sporolithon ptychoides Heydrich ( =Spongites erythraeum) 5 1. Spongites sporolithon 52. Lithophyllum kotschianum (Fosl.) 53. Cheilosporum spectabile Haw. 54. Haliptilon subulatum (Ell. et Sol.) Johan. 55. Jania adhaerens Lam. 56. Jania capillacea Haw. 57. Jania decussato-dichotoma (Yendo) Yendo 58. Jania longiartha Daws. 59. Jania ungulata (Yendo) Yendo f. brevior (Yendo) Yendo 60. Jania sp. 61.Amphiroa anastomosans W.V.Bosse 62.Amphiroa foliacea Lam. 63.Amphiroa ji-agilissima (L.) Lam. 64. Metagoniolithon stelligerum (Lam.) W.V.Bosse Cryptonemiales Peyssonneliaceae 65. Pqssonnelia dubyi Crouan 66. Pqssonnelia sp. 1 67. Pqssonnelia sp.2 Dumontiaceae 68. Dudresnaya sp. 69. Gibsmithia hawaiiensis Doty Halymeniaceae 70. Halymenia sp. Kallymeniaceae 71. Kallymenia sp. Gigartinales Gymnophlocaceae 72. Titanophora sp. 73. Portieria homemannii (Lyngbye) Silva Hypneaceae 74. Hypnea cenomyce J. Ag. 75. Hypnea cervicomis J. Ag. 76. Hypnea comuta (Kutz.) J. Ag. 77. Hypnea esperi Bory 78. Hypnea nidulans Setchell 79. Hypnea pannosa J. Ag. 80. Hypnea spinella (C. Ag.) Kutz. 81. Hypnea valentiae (Turn.) Mont. 82. Hypnea sp.
CoF
A
D
Island P MABS M AB M M M M M M M M M M S
PdCsAs
ABS S AB S S
C
o
F
A
D
P
M
S
Cs PdCsAs Pd Cs
D
P
M
P C C P oFADP P oFADP FADP DP FA M M M S MAB M M S S Pd Cs As PdCsAs Cs As As As
Table 1. Continued. Taxon Plocamiaceae 83.Plocamium sp. Caulacanthaceae 84. Caulacanthus sp. Wurdemanniaceae 85.Wurdemannia miniata (Sprengel) J. Feldm. et Hamel Phyllophoraceae 86. Gymnogongrus chnoosporoides Tan. et Pham-Hoang 87.GymnogongrusWgmaeus J. Ag. Gigartinaceae 88. Gigartina sp. Gracilariaceae 89.Gracilaria crassa Ham. 90.Gracilaria cylindrica B+rg. 91.Gracilaria multifircata B@g. 92.Gelidwpsisgracilis (Kiitz.) Vickers 93.Gelidiopsis scoparia (Mont. et Mill.) Schmitz 94.Gelidwpsis sp. 95.Ceratodictyonspongiosum Zanard. Rhodymeniales Rhodymeniaceae 96.Botyocladia s k o t t ~ b e (B4rg.) Levr. ~i 97.Chrysymenia enteromorpha Ham. 98.Chrysymeniapyri~omis B@g. 99.Chrysymenia sp. 100.Coelarthrum boergesenii W.V. Bosse 101.Rhodymenia anastomosans W.V.Bosse 102.Rhodymenia sp. Lomentariaceae 103.Lomentaria corallicola Bq5rg. 104.Lomentaria mauritiana Bcprg. Champiaceae 105.Champia globulifea Bcprg. 106. Champia indica B&g. 107.Champiaparvula (C. Ag.) Ham. 108.Champia salicomoides Ham. 109.Champia viellardii Kiitz. 1 1 .Champia sp. 0 Ceramiales Ceramiaceae 111 Crouania attenuata (C. Ag.) J. Ag. . 112. Antithamnion fherminieri Nasr. 113.Antithamnion sp. CoF A D Island P MABS PdCsAs
F
D
P
M
S
Pd
As
F
A
P
M
ABS
Cs As
CoF Co CoF CoF
A A A
D
P
M
ABS Pd AB S
Cs As
D
AB S M
Pd
D
D D D
CoF F A A D A Co
A
P P
MABS M
Pd As PdCsAs Pd
D D
P P
MABS M S M S AB S M ABS
D D D
Pd Cs
Table 1. Continued. CoF 114. Platythamnion sp. 115.Antithamnionella sp. 116. Ceramium brevizonatum var. caraibicum H. Petersen et B&. 117. Ceramium fastigiatum (Wulf. ex Roth) Haw. f. flaccidum H. Petersen 118. Ceramium jimbriatum Setch. et Gardn. 119. Ceramium gracillimum (Kiitz.) Zanard. 120. Ceramium huysmansii W.V.Bosse 121. Ceramium mazatlanense Daws. 122. Ceramium taylorii Daws. 123. Ceramium sp.1 124. Ceramium sp.2 125. Ceramium sp.3 126. Centroceras clavulatum (C. Ag.) Mont. 127. Centroceras apiculatum Yam. 128. Spyndiajilamentosa (Wulf.) Haw. 129. Wrangelia argus (Mont.) Mont. 130. Callithamnion byssoides Am. 131.Aglaothamnion sp. 132. Pleonosporium borreri (Smith) N2g. 133. Haloplegma duperreyi Mont. 134. Grifithsia globulifera (Haw.) Kiitz. 135. Grifithsia metcalfii Tseng 136. Grifithsia subcylindrica Okam. 137.Anotrichium tenuis (C. Ag.) Nag. 138. Grifithsia weber-van-bosseaeB@g. 139. Grifithsia sp. 140. Gymnothamnion elegans (Schousboe ex C. Ag.) J. Ag. Delesseriaceae 141. Caloglossa adnata (Zanard.) De Toni 142. Caloglossa leprieurii (Mont.) J. Ag. 143. Caloglossa stipitata Post 144.Hypoglossum attenuatum Gardn. 145. Taeniomaperpusillum (J. Ag.) J. Ag. 146. Cottoniella arcuata B#q. 147. Platysiphonia sp. 148. Martensia sp. Dasyaceae 149. Dasya mollis Haw. 150.Dasya baillouviana (Gmel.) Mont. 151.Dasya sp.1 152. Dasya sp.2 153. Dasya sp.3 A D Island P MABS AB F Co Co F Co F CoF CoF A A A A A F CoF F D D D D P D D P P P P M M M M M M MA M M CoF CoF Co F A A A A P A A A A A D Pd D D P CoF Co A A A D D M Cs P M S P AB S AB S Cs S Pd Cs D M D D D D P M AB S ABS ABS S Pd Cs Cs S ABS Pd Cs AB S ABS S ABS S B AB S S S Pd Cs Cs Cs Cs Cs Cs As Cs Cs A D P M Pd Cs PdCsAs
ADP ADP A A AD
Table 1. Continued. CoF 154. Heterosiphonia crispella (C. Ag.) Wynne (=Heterosiphonia wurdemannii) 155. Heterosiphonia sp.1 156. Heterosiphonia sp.2 157. Dictyuruspurpurascens B o ~ y Rhodomelaceae 158. Polysiphonia coacta Tseng 159. Polysiphonia mollis J. Hook. et Haw. 160. Polysiphonia ferulacea Suhr ex J. Ag. 161. Polysiphonia scopulorum Haw. 162. Polysiphonia subtilissima Mont. 163. Polysiphonia sp. 164. Truialia sp. 165. Tolypiocladia glomerulata ( Ag.) Schmitz 166. Lophocladia trichoclados (C. Ag.) Schmitz 167. Boshychia binderi Haw. 168. Herposiphonia secunda (C. Ag.) Ambronn f. tenella (Ag.) Wynne 169. Leveillea jungermannioides (Hering et Martens) Haw. 170. Lophosiphonia reptabunda (Suhr.) Kyl. villum (J. Ag.) Setch. et Gardn. 171. ~o~hosiphonia 172. Lophosiphonia sp. 173. Chondria dayphylla (Woodw.) C. Ag. 174. Chondria poiyrhiza Coll. et Hew. 175. Chondria repens B+rg. 176. Chondria sp. 177.Acanthophora spicifera (Vahl) B+rg. 178. Laurencia articulata Tseng 179. Laurencia corymbosa J. Ag. 180. Laurencia decumbens Kiitz. 181. Laurencia distichophylla J. Ag. 182. Laurencia obtusa (Huds.) Lam. 183. Laurenciapatentiramea (Montagne) Kiitz. 184. Laurencia par/ipapillata Tseng. 185. Laurencia papillosa (Ag.) Grev. 186. Laurencia~gmaeaW.V.Bosse 187. Laurencia sp.1 188. Laurencia sp.2 189. Laurencia sp.3 190. Laurencia sp.4 191. Laurencia sp.5 CoF Co CoF CoF Co F P P Co FAD CoFADP Co F D P Co F A D P Co CoF Co F Co F A A D P P P P MAB S M M M Cs AB S Pd As Cs M ABS A A A A A D D D D Island P MABS P P P MABS S AB ABS PdCsAs PdCs Cs As PdCsAs
MABS M M S AB S S
PdCsAs Pd Cs Cs Cs Cs PdCs Cs As Pd Cs As Cs As
A A A
D
Co F F
P A A P A A A A A A A A D P P D P
MABS M S S M M M M M M M ABS S S AB S
F Co F Co F CoF F CoF
D D
P
Cs As PdCsAs PdCsAs As
AB S
Table 1. Continued. Taxon CoF A D Island P MABS PdCsAs
Ectocarpales Ectocarpaceae 192. Ectocarpus sp. 193. Hincksia mitchelliae (Haw.) Silva 194. Hincksia breviarticulata (J. Ag.) Silva 195. Streblonema sp. Scytosiphonales Scytosiphonaceae 196. Colpomenia sinuosa (Mert. ex Roth) Derb. et Sol. 197. Hydroclathrus clathratus (C. Ag.) Howe 198. Rosenvingea intricata (J. Ag.) B4rg. Chnoosporaceae 199. Chnoospora minima (Hering) Papenf. Sphacelariales Sphacelariaceae 200. Sphacelaria rigidula Kutz. 201. Sphacelaria nibuloides Menegh. Dictyotales Dictyotaceae 202. Dictyota bartayresii Lam. 203. Dictyota ceylanica Kutz. 204. Dictyota divaricata Lam. 205. Dictyota friabilis Setch. 206. Dictyota indica Sond. ex Kiitz. 207. Dictyota patens J. Ag. 208. Dictyota sp. 209. Dictyopteris delicatula Lam. 210. Dictyopterispolyposioides (DeCandalle) Lam. 211. Padina minor Yam. 212. Padinapavonica (L.) Thivy 213. Padina sp. 214. Lobophora variegata (Lam.) Womers. Fucales Cystoseiraceae 215. Hormophysa cuneiformis (Gmelin) Silva Sargassaceae 216. Sargassum cristaefolium C. Ag. 217. Sargassum ilicifolium (Turn.) C. Ag. 218. Sargassum mcclurei Setch. 219. Sargassum microcystum J. Ag. 220. Sargassum piluliferum (Turner) C. Ag.
D P P
M M M
M M M M
F Co
A
P P
MAB M
Cs Cs
A A
D
P P P P P P P P P P
Co F F
D
M M M M M M
S S
Pd
CoF
A
D
MABS M M M M M
PdCsAs
P P P
Table 1. Continued. Taxon
221. Sargassum polycystum C. Ag. 222. Sargassum turbinarioides Grun. 223. Sargassum sp. 224. Turbinaria conoides (J. Ag.) Kutz. 225. Turbinaria decurrens Bory 226. Turbinaria omata ( u n ) Ag. T r .J. 227. Turbinaria sp.
CHLOROPHYTA
CoF
A
D
Island P MABS M M M M M AB S
PdCsAs
A C o F A D
P P P P P
Cs As Cs
Phaeophilales Phaeophilaceae 228. Phaeophila dendroides (P. et H. Crouan) Batt. Dasycladales Dasycladaceae 229. Neomeris annulata Dickie 230. Neomeris bilimbata Koster 231. Neomeris vanbosseae Howe 232. Neomeris sp. Polyshysacaea 233. Polyphysa sp. 234. Acetabularia clavata Yam. 235. Acetabularia &gua Solms-hub. 236. Acetabulariaparvula Solms-hub. 237. Acetabulariapusilla (Howe) Collins 238. Acetabularia sp. Bryopsidales Caulerpaceae 239. Caulerpa brachypus Haw. 240. Caulerpa cupressoides (Vahl) Ag. 241. Caulerpa cupressoides var. mamillosa (Mont.) W.V.Bosse 242. Caulerpafastigiata Mont. 243. Caulerpa mexicana Sond. ex Kutz. 244. Caulerpa microphysa (W.V.Bosse) J. Feldm. 245. Caulerpapeltata Lam. 246. Caulerpa racemosa (ForsskAl) J. Ag. var. macrophysa (Sond. ex Kutz.) W. Taylor var. occidentalis (J. Ag.) B@g. var. peltata (Lam.) Eubank 247. Caulerpa serrulata (ForsskAl) J. Ag. var. serrulata f. spiralis (W.V.Bosse) Gilbert 248. Caulerpa setularioides (Gmel.) Howe f. farlowii (W.V.Bosse) B@g.
A A A A
D D D D P P
M M M M
Cs Cs
F F F F F F F F F
A
D D D
P P P P P P P P P
ABS
Cs Pd Cs Cs
A A
S M M M M M M M S S ABS
Cs Cs Cs Cs Cs Cs
A
D
Table 1. Continued. Taxon 249. Caulerpa tarifolia (Vahl) C. Ag. 250. Caulerpa ambigua Okam. 251. Caulerpa webbiana Mont. 252. Caulerpa sp. 1 253. Caulerpa sp.2 254. Caulerpa sp.3 Udoteaceae 255. Chlorodesmis comosa Haw. et Bail. 256. Boodleopsispusilla (Collins) W. Taylor, Joly et Bernatowicz 257. Avrainvillea amadelpha f. submersa Gepp 258. Tydemania gardinen A et B. Gepp 259. Rhipiliopsk reticulata (van den Hoek) Farghaly 260. Rhipilia tomentosa Kiitz. 261. Udotea argentea Zanard. 262. Udoteaflabellum (Ell. et Sol.) Howe 263. Udoteajavensis (Mont.) A Gepp et E. Gepp 264. Udotea orientalis A Gepp et E. Gepp Halimedaceae 265. Halimeda gracilis Haw. ex J. Ag. 266. Halimeda copiosa Goreau et Graham 267. Halimeda macroloba Decne. 268. Halimeda micronesica Yam. 269. Halimeda opuntia (L.) Lam. 270. Halimeda stuposa W. Taylor 271. Halimeda tuna (Ell. et Sol.) Lam. 272. Halimeda sp. Codiaceae 273. Codiumformosanum Yam. 274. Codium geppii O.C. Schmidt 275. Codium tomentosum (Huds.) Stackh. 276. Codium sp. Bryopsidaceae 277. Derbesia marina (Lyngb.) Solier 278. Derbesia sp. 279. Halicystis ovalis (Lyngb.) Aresch. (N.B. - sporophyte stage of Derbesia marina) 280. Bryopsis pennata Lam. 281. Tnchosolen sp. Siphonocladales Valoniaceae 282. Valonia aegagropila C. Ag. 283. Valoniafastigiata Ham. es J. Ag. 284. Valonia utriculari. (Roth) Ag. f. crustacea Kuck. CoF CoF Co F A Co C o F A CoF Co Co F Co F Co Co F CoF Co F CoF CoF CoF CoF A A A A A A A A A A D P M AB M ABS AB S AB S ABS ABS Pd PdCsAs Cs As Pd As Cs PdCsAs PdCs As AB S S Pd Cs As DP DP D D D D D D D D D D D D D D P P P A A D Island P MABS P P M M
S S
PdCsAs
Cs Cs
Cs Cs As
M M M
MABS AB S MABS AB S M ABS AB S ABS AB S M M M S S
F
A P
M M
A
D
P
M S
Co F Co
D D D
MABS M ABS S
PdCsAs Pd Cs
Cs
Table 1. Continued. Taxon
285. Ventricaria ventricosa (J. Ag.) Olsen et West 286. Diclyosphaeria cavemosa (ForsskAl) B+rg. 287. Diclyosphaenh versluysii W.V.Bosse 288. Valoniopsispachynema (Mart.) B+rg.
CoF F Co F CoF Co
A A A A A
D
Island P MABS M M M M M P M M M M
PdCsAs Cs Pd Cs As Cs As Cs As Cs Pd Cs As
D D D D D
P P
AB S ABS AB S AB S ABS ABS S
Siphonocladaceae
289. Boergeseniaforbessii (Ham.) J. Feldm. 290. Siphonocladus rigidus Howe Co 291. Siphonocladus tropicus (P. et H. Crouan) J. Ag. 292. Cladophoropsisherpestica (Mont.) Howe 293. Cladophoropsismembranacea (Ag.) B4rg. 294. Cladophoropsis modonensis (Kutz.) B+rg. 295. Cladophoropsissundunensis Reinb. CoF
Cs Cs
A
D D D D D D
P P
Boodleaceae 296. Boodlea composita (Ham.) Brand Co 297. B d l e a siamensis Reinb. 298. BoodIea struveoides Howe .CoF 299. Boodlea sp. 300. Struvea anastomosans (Ham.)Picc. et Grun. Co F 301. Struvea elegans B4rg. F 302. Struvea sp. 1 Co Co 303. Struvea sp.2 Anadyomnaceae 304. Microdiclyon okamurae Setch. CoF 305. Microdidyon montagnei Ham. Co 306. Microdicyon sp. Co 307. Anadyomeneplicata C. Ag. 308. Anadyomene wrightii Ham. ex J.E. Gray CoF Cladophoraceae 309. Cladophora vagabunda (L.) van den Hoek 3 10. Cladophora laetevirens (Dillw.) Kutz. Co 31 1. Cladophorapatentiramea (Mont.) Kutz. 312. Cladophora socialis Kutz. F 3 13. Cladophora sp. 314. Rhizoclonium implexum (Dillw.) Kutz. F (=Rhizoclonium kenten) 315. Chaetomorpha ligustica (Kutz.) B4rg. Co 316. Chaetomorpha crassa (C. Ag.) Kutz. Co F Chaetophorales Ulvellaceae 317. Entocladia sp. F 318. Phaeophila dendroides (P. and H. Crouan) Batt. Ulvales Ulvaceae 3 19. Enteromorpha clathrata (Roth) Grev. Co 320. Enteromorphajlexuosa (Wulf.) J. Ag.
A A
P P P P
MABS S M AB S
PdCsAs Pd
A A A
D
M
D
ABS S S AB ABS S S S
P d C s As Pd Cs PdCsAs Cs Cs
P A
M M M M M M M M
Cs As As
P ADPMAB D
Cs Cs
Table 1. Continued. CoF
321. Enteromorpha intestinalis (L.) Link ex Nees 322. Enteromorpha hylinii Bliding 323. Enteromorpha linza (L.) J . Ag. 324. Enteromorpha ralfsii Harv. 325. Enteromorpha sp. 326. Ulva rigida C. Ag.
SPERMATOPHYTA
A
D D D D
Island P MABS P P
PdCsAs
CoF
A A
CoF
A
Potamogetonaceae 1. Halodule uninervis (Forsskal) Aschers. 2. Cymodocea rotundata Ehrenb. and Hemprich ex Aschers. 3. Cymodocea semlata (R. Br.) Aschers. et Magnus 4. Syringodium isoetifolium (Aschers.) Dandy Co 5. Thalassodendron ciliatum (Forsskal) den Hartog Co F Hydrocharitaceae 6. Thalassia hemprichii (Ehrenb.) Aschers. CoF 7. Halophila ovalis (R. Br.) Hooker f. hawaiiana (Doty et Stone) den Hartog 8. Halophila stipulacea (Forsskal) Aschers.
P P P A A A D D D P P M M M M AB S ABS ABS S P d C s As P d C s As
Table 2. Quantitative characteristic of bottom vegetation of the Seychelles Islands (12 January - 12 March, 1989). In zone heading, "rnih" = middle intertidal horizon, "lih" = lower intertidal horizon. "s" = sand, "st" = stones, "c" = corals, "cd" = coral debris and "Hs" Halimeda sand. = Depth in # of Biomass, % Growth Coast Station m, zone Substrate species.m-' in g.m-2 cover height Association
H
COetivy Island
N-W N-W N-W N-W N-W N-W 1 2 3
mih mih mih mih lih lih mih
s, &,st s, cd s, cd s,cd S, cd, S, cd, s, cd, s, cd, S, cd, s,cd s, cd, s, cd, S, cd, S, cd, s, cd s, cd s, cd s, cd s, cd s, cd s, cd s, cd S, cd s, cd s, cd S, cd s,
5 100 100 70
80
4 5
6
100
90 100 100 90 90
S S S
S
S
N N-E W W W W W W W W W W E E E E E E
7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
mih
lih mih mih 1-2 1 8 1.5 2.5 3.5 3.5 4.0 4.5 3.0 4.5 5-6 mih mih lih 2
90 100 50
80
100 100 100 80 30 100 95 50 10
N-E N-E N-E N-E N-E
- Enteromorpha kylinii 15 Thalassia hemprichii 10 Halodule uninervis +Thalassia hemprichii 6 Halodule uninervis Thalassia hemprichii 15 Thalassodendron -Halimeda+Hypnea 20 Thalassodendron-Halimeda -Dictyurus 20 Thalassodendron 15 Thalassia-Caulerpa 10 Halimeda+Dictyuius +Lawencia 15 Thalassia Thalassodendron 20 Halimeda Thalassodendron ciliarum-Laurencia 25 Thalassodendron-Halimeda 25 Thalassodendron-Halimeda 30 Thalassodendron-Caulerpa 10 Halemeda 10 Halemeda 20 (Tha1assodendron)-Halemeda 10 Halemeda 10 Halemeda 10 Halemeda 21 Thalassodendron ciliatum 22 Thalassodendron-Halimeda 25 Thalassodendron 12 Thalassiu 25 Thalassodendron-Thalassia 25 Thalassodendron-Dictyuius 20 Thalassodendron-Halimeda -Dictyurus 20 Thalassodendron-Halimeda 12 (Thalassodendron)-Caulelpa -Halimeda 18 (Tha1assodendron)-Caulerpa-Halimeda 10 Halimeda +Boodles +Microdictycon 23 Thalassodendron-Halimeda Thalassodendron-Halimeda 27 Thalassodendron-Halimeda
+
+ +
Table 2. Continued. Depth in # of Biomass, % Growth Coast Station m, zone Substrate ~ ~ e c i e s . m -g.m-2 cover height Association in ~
H
N-E N-E N-E N-E
S S S
W W W W W W
35 mih 36 mih 37 lih 38 32 39 5 40 10 41 12 42 43 44 45 46 47 48 49 50 51
w.
W W W
mih mih mih lih 3 3 6 7 17 1.5
Thalassia Thalassodendron+ Thalassia Thalassodendron-Dictyurus Avrainvillea-Halimeda Thalassodendron Thalassodendron (Thalassodendron)-Halimeda -Microdictyon Thalassia +Halodule Syringodium + Thalassodendron Halodule + Thalassia Thalassodendron Thalassodendron Thalassodendron Thalassodendron Thalassodendron Thalassodendron-Dictyurus Thalassodendron
Farquhar Island
S S S S
52 mih 53 lih 54 lih 55 lih 6 15 25 mih lih lih 6-7 12h 22 3 7 9
15 Thalassodendron+Thalassia 13 Thalassodendron 1-2 Boodlea 15 (Thalassodendron)-Caulerpa
-Bodlea
56 57 58 N 59 N 60 N 61 N-E 62 N-E 63 N-E 64 Lagoon 65 Lagoon 66 Lagoon 67
S S S
3 Caulerpa 1-2 Microdictyon +Boodlea 1-2 Microdictyon 8 Thalassodendron-Boodlea 1-2 Boodlea struveoides 1-2 Microdictyon +Boodlea 2 Caulerpa+Boodlea 2 Caulerpa 2-5 Udotea+Caulerpa 2 Caulerpa cupressoides 2 Caulerpa cupressoides 2 Halimeda opuntia
Lagoon Lagoon Lagoon Lagoon Lagoon Lagoon
68 2.5 69 8 70 12 71 3 72 1.2 73 1.0
+Caulerpa cupressoides 28 Thalassodendron
---
35 Thalassodendron-Halimeda 24 Thalassodendron-Halimeda 28 Thalassodendron-Caulerpa
Table 2 Continued. .
# of Biomass, % Growth Depth in Coast Station m, zone Substrate species.m-2 in g.m-2 cover height Association
Lagoon Lagoon Lagoon Lagoon Lagoon Lagoon Lagoon N N
74 8 75 10 76 12 77 6 78 3-4 79 2 80 1 81 mih 82 lih 83 lih 84 4 85 1
0 0 0 1 100 100 100 100 100 100 100 70
--
--
-- Heterosiphonia
30 Thalassadendron-Halimeda 126 Thalassadendron-Thalassia 20 Thalassadendron-Caulerpa 14 Thalassadendron-Caulerpa 12 Thalassadendron- Valonia Microdictyon 1 Thalassadendron-Valonia 2 50 Sargassum ilicifolium 25 Thalassodendron
+
Aldabra Island
E 86 12 E 87 25 E 88 12 Lagoon 89 lih
s, cd s, cd Halimeda s 518 S, cd s, cd s, cd s, cd s,cd s, black c s, black c 376 284 584 960 1004 960
60 20 70 70 80 80 100 100 100 100 100
10 1 0 30 18 12 25 1 7 24 24 1 5 1 7 20
Lagoon Lagoon Lagoon Lagoon W W
90 91 92 93 94 95
lih lih mih 5 1 lih
96 lih 97 lih
s, black c 768 s, black c
-
Halimeda Halimeda lOHalimeda (Thalassadendron)-Lawencia +Halimeda Thalassia +Lawencia Thalassia +Halimeda Thalassia +Halimeda Thalassia Thalassadendron Thalassadendron Thalassia -Halimeda Thalassadendron +Thalassia -Halimeda Thalassadendron +Halimeda
+
Desroches Island
N-W N-W N-W N-W N-W N-W N-W
98 mih 99 mih 100 101 102 103 lih 2 6 12
662 2400 3864 2164 1572 1700
80
100 100 70 100 50 25
20 Thalassodendron +Thalassia -Halimeda 30 Thalassadendron-Halimeda -Haloplegma 42 Thalassadendron 40 Thalassadendron 42 Thalassadendron 25 Thalassadendron-Halimeda Udotea 10 Halimeda
+
Table 2. Continued.
# of Biomass, % Growth Depth in Coast Station m, zone Substrate species-m-2in g-m-2 cover height Association
H
N-W N-W N-W N-W N-W S-E S-E W W W W S S S S S S S S-E S-E S-E S-E S-E S-E S-E S-E S-E S-E S S S S S S S S-W S-W S-W S-W S-W S-E
105 30 106 mih 107 mih 108 10...