Species Diversity and Distribution of Mangrove-Associated Red Alga Bostrychia (Rhodomelaceae, Rhodophyta) from Southern Thailand

International Journal of Applied Environmental Sciences ISSN 0973-6077 Volume 11, Number 1 (2016), pp. 55-71 © Research India Publications http://www.ripublication.com

Species diversity and distribution of mangrove-associated red alga Bostrychia (Rhodomelaceae, Rhodophyta) from southern Thailand

J. Seangkaew1, S. Bovonsombut1 and Y. Peerapornpisal1*

1Department of Biology, Faculty of Sciences, Chiangmai University, Chiangmai 50200, Thailand

Abstract

The diversity and distribution of Bostrychia in Thailand were conducted to fill up document in comparison with that of research in others areas and updated information. Four sites along the Andaman Sea: Ranong, Phangnga, Phuket, and Krabi provinces and three sites on the Gulf of Thailand: Chumphon, Surat Thani, and Nakhon Si Thammarat provinces were studied. Algal samples were collected every 2 months during October 2012 to August 2013. The algal distribution with respect to the zonation, water, and sediment characteristics were compared between sites, among transects and among quadrats (50 x 50 cm2). The results indicated that intertidal position within sites have the greatest influence on algal frequency. Individual species dominated in different intertidal zones. Eight species of Bostrychia was detected: Bostrychia binderi Harvey, B. calliptera (Montagne) Montagne, B. kelanensis Grunow, B. moritziana (Sonder ex Kützing) J. Agardh, B. radicans (Montagne) Montagne, B. radicosa (Itono) J. A. West, G. C. Zuccarello and M. H. Hommersand, B. simpliciuscula Harvey ex J. Agardh and B. tenella (Lamouroux) J. Agardh. Bostrychia moritziana, B. radicans and B. tenella were abundant and common on both coasts of Thailand, whereas B. calliptera was scarce and restricted to the Andaman Sea. These results suggest that the abundance and distribution patterns of algal communities were likely to be influenced by seasonal variation, wave action, sediment, nutrients and other chemical-physical parameters and habitat conditions.

Keywords: Diversity, Bostrychia, Thailand, Sediment, Water quality

56 J. Seangkaew et al

Introduction Mangroves are among the most productive and biologically complex ecosystems. The rich assemblage of species found associated with subtidal mangrove habitats (aerial roots, peat walls) offer the potential to serve as indicators of the biological and ecological status of this ecosystem (Tomlinson,1986[1]). They are sometimes called rainforests by the sea occupy tropical regions at river mouths, bays, coastal lagoons and islands. They grow in areas of desiccating heat, deep mud, and salt levels that would kill an ordinary plant within few hours and grow luxuriantly in the places where freshwater mixes with seawater and where sediment is composed of accumulated deposits of mud. Mangroves protecting a coastline by reducing erosions from wave currents and storm energy in the form of waves action, creating a natural breakwater that helps stop erosion, reducing from coastal storms are generally affected and property. expected increase of the cyclones, monsoon, tsunami, storm and hurricane winds intensity and rising sea levels, mangroves will become increasingly. Mangrove protecting coastal land and provides a buffer between the terrestrial and nearby marine environments and helping to maintain water quality. (Felício et al, 2008[2]) and provides a filter through the mesh of aerial roots that retains the particles that might pollute the lagoon environment (Chapin 2003[3]; Feller et al, 2010[4]). Mangrove habitats have demonstrated the high biological productivity and rich biodiversity of these ecosystems in tropical and sub-tropical regions. Their productivity values vary with the type of primary producers (mangroves, microalgae and macroalgae) found in this ecosystems, The root system is an environment they are provide habitats support a wide diversity of marine life by providing food sources for other species range of wildlife species including commercial for the smallest animals such as birds, mammals, worms, snails, shrimp, mollusks, nudibranchs, mussels, barnacles, clams, oysters, herbs, trees, fish and crustaceans (Aburto-Oropeza et al. 2008[5]). Mangroves also shelter and provide nursery grounds for many commercially and culturally important species of fish and invertebrates lay their eggs so their descendants might grow up and they are protected from most predators, and thus contribute to supporting of local bundance and regional distribution of fish and invertebrate populations (McIvor et al. 2012 [6]; Ley et al. 2002[7]). Mangroves maintain coastal water quality by abiotic and biotic retention, removal, and cycling of nutrients, pollutants, and particulate matter from land-based sources, and supply nutrients to adjacent coral reef, seaweed and seagrass communities. (Walters et al. 2008[8], Koch et al. 2009[9]). Mangrove root systems slow water flow, facilitating the deposition of sediment. For instance, terrigenous sediments and nutrients carried by freshwater runoff are first filtered by coastal forests. Toxins and nutrients can be bound to sediment particles or within the molecular lattice of clay particles and are removed during sediment deposition. Mangrove forests are significantly important as hot spots for tropical coastal biodiversity. Mangroves perform a vital ecological role providing the habitats for many wildlife species, including macroalgae. Mangrove macroalgae have an important role as primary producers in the estuarine Species diversity and distribution of mangrove-associated red alga Bostrychia 57 ecosystem and contribute to the total productivity of mangrove forests through the production of organic materials to nutrient cycling, as food sources for grazing marine animals and as habitat for small estuarine invertebrates (Kamer and Fong 2000[10]). Marine algae are an ecologically and economically important component of marine ecosystems worldwide. The macroalgal are rich in mangrove habitats where it contributes to primary producers and provide shelter, nursery grounds, and food sources for marine organisms. Mangrove macroalgal assemblages normally grow epiphytically on pneumatophores, stems and other hard substrates (Kamiya and Chihara 1987[11]; Zuccarello et al. 2001[12]), and have an important role as primary producers, as food sources for grazing marine animals and as habitat for small estuarine invertebrates (Kamer and Fong. 2000[10]). The most common red algae associated with the mangrove areas are Bostrychia Montage, Caloglossa (Harvey) Martens and Catenella Greville (West et al. 2006[13]). Bostrychia is filamentous alga that can be found in marine, brackish and even in freshwater habitats throughout the tropical and temperate regions (West et al. 2006[13]). This genus is currently composed of about 19 species (Zuccarello et al. 2015[14]). Traditionally, species-level classification of the genus is mainly base on a few morphological characters (e.g. number of tier cells per axial cell, type of attachment structure, absence or presence of cortication and branching pattern) (Zuccarello and West 2006[15]). In Thailand, a few studies have been done to observe the diversity and distribution of mangrove-associated marine algae (Lewmanomont. 1976[16]; Coppejans et al. 2011[17]). Previous taxonomic studies of the genus Bostrychia recognized 5 species from both Andaman Sea and The Gulf of Thailand, including B. moritziana (Sonder ex Kützing) J.Agardh, B. radicans (Silva et al. 1996[18]; Ruangchuay et al. 2006[19]). However, Thailand is located around an equator with hot and rainy climate and surrounded by marine ecosystems include the areas of the Gulf of Thailand in the Pacific Ocean and the Andaman Sea in the indian Ocean separated by the Southern Peninsular of Thailand, The Gulf is a part of Sunda Shelf and is a semi-enclosed sea located in Southeast Asia off of the South China Sea and is also very shallow (on average only 45 meters deep) and strong inflow from rivers draining the mainland makes it low in salinity and rich in nutrients and sediments. Circulation of currents in the Gulf are influenced by two seasonal monsoons and water is exchanged between the Gulf and the South China Sea very slow. The Andaman Sea is deeper and with higher salinity and lower turbidity levels than the Gulf (Kantachumpoo et al. 2014[20]). Andaman species composition are influenced by physical and biological environment of the Indian Ocean.

The aim of this study was therefore to look for seasonal distribution of the difference taxa of Bostrychia and sediment, some parameters of water quality (salinity, nitrate, phosphate) analysis. In the Upper Southern Ranong, Phangnga, Phuket, Krabi (Andaman sea) and Chumphon, Suratthani, Nakhon Si Thammarat (Gulf of Thailand) mangrove in relation to environmental parametry.

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Materials and Methods Algal samples were collected from 7 stations along the Gulf of Thailand and the Andaman Sea (Fig.1) from October 2012 to September 2013. Other samples were collected in the field and dried onto herbarium sheets. Voucher specimens were deposited at Applied Algal Research Laboratory, Department of Biology, Faculty of science, Chiang Mai University, Mueang 50200, Thailand. (Table.1). And some specimens were either preserved in silica gel or fixed in 4% formalin /seawater. Some samples were prepared for microscopy by soaking dried samples in sea water for 10 min and then simultaneously staining and preserving the specimens in 1 % aniline blue intensified with1 % HCl and mounted using a 50 % glucose syrup (Karo Syrup, Corn Products) on a microscope slides. Digital images were photographed by microscope digital camera Olympus DP20 (Olympus, Tokyo, Japan) and eventually edited using Photoshop Elements 6 (Adobe, San Jose, CA, USA).

Figure 1: Map of Thailand showing the collection sites.

Species diversity and distribution of mangrove-associated red alga Bostrychia 59

Results Morphological observation: Our Bostrychia samples collected from the southern Thailand were morphologically identified for the seven species (Table 1-2).

Table 1: Temporal changes in species diversity of Bostrychia species in the andaman sea from 2012-2013.

Species Ranong Phangnga Phuket Krabi Division Rhodophyta Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun Aug Class Florideophyceae Order Ceramiales Family Rhodomelaceae Bostrychia binderi / / / / / / / / / / / / / / / / / / / / / / / / B. calliptera / / / / / / B. kalensis / / / / / / B. moritziana / / / / / / / / / / / / / / / / / / B. radicans / / / / / / / / / / / / / B. radicosa / / / / / B.simpliciuscula / / / / / B. tenella / / / / / / / / / / / /

Table 2: Temporal changes in species diversity of Bostrychia species in the gulf of Thailand from 2012-2013.

Species Chumphon Surat Thani Nakhon Si Thammarat Division Rhodophyta Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun Aug Class Florideophyceae Order Ceramiales Family Rhodomelaceae Bostrychia kalensis / / / / / / / / / / / / / / / / / / B. moritziana / / / / / / / / / / / / / / / / / / B. radicans / / / / / / / / / / / / / / / / / / B. radicosa / / / / / / / / / / / / B. simpliciuscula / / / B. tenella / / / / / /

Eight species of Bostrychia were identified from mangrove forests in the upper southern Thailand. The descriptions of these taxa are as follows:

Bostrychia calliptera (Montagne) Montagne This species grow abundantly on the mangrove tree in upper zone. Thalli about 1-2 cm high, reddish purple to brown. Thalli erect with subpostrate branching pinnate alternate, corticated, polysiphonous thoughout each axial cell with two tier cells of 6-8 periaxial cells, peripherohaptera developed form ventral and lateral periaxial cells.

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Tetrasporangial sticihdia developed at apical of lateral branching, 0.8-1.2 mm long and 120-200 µm in diameter. No male and female observed. (Fig. 2. A-C).

Bostrychia kelanensis Grunow This species grow abundantly on the roots of the Avicennia marina and Rhizophora Xylocarpus. It is normally found as a mixed patchy assemblages with other Bostrychia species (e.g. B. moritziana and B. radicans). Thalli are about 0.1-1 cm high, which consist of both prostrate and erect system.. Thalli are polysiponous throughout, alternate branching, multiple new shoots alternate branches bonre from cladohaptera attachment disc. composing of three tier cells and cladohaptera. Thalli occasionally have a cortication on the main axis and determinate lateral branches. Spermatangial stichidia are borne below the tips of laterals branches, 0.3–1 mm long and 50–75 µm wide. Cystocarps are globular 300-400 µm in diameter, 350-500 µm long, producing at the terminal of lateral branches. Tetrasporangial stichidia are developed below tips of lateral branches, approximately 0.4–1 mm long and 30-70 µm wide (Fig.3. D-L).

Bostrychia moritziana (Sonder ex Kützing) J. Agardh Thalli are about 1-3 cm high, reddish purple to brown. Thalli postrate, alternate branching, The main axes polysiphonous throughout, ultimate branches monosiphonous. Thalli are composed with two tier cells of 4-8 periaxial cells to each axial cells and attached by cladohaptera on substrate. Usually ecorticated cells but sometimes have a cortication development throughout are associated with season. Tetrasporangial stichidia developed below tips of lateral branching, Cystocarps developed near the apical of lateral branching, ovoid to spherical 400-550 µm in diameter, 500-650 µm long. Spermatangial stichidia developed below apical of lateral branching 25-60 µm in diameter, 120-220 µm long. (Fig.3 A-E).

Bostrychia radicans (Montagne) Montagne Thalli are about 1-3 cm high, reddish purple to brown. Thalli postrate and suberect, alternate branching, The main axes polysiphonous throughout. Thalli are composed with two tier cells of 5-9 periaxial cells to each axial cells and attached by cladohaptera on substrate. Usually ecorticated cells but sometimes have a cortication development throughout are associated with season. Tetrasporangial stichidia developed below tips of lateral branching, Cystocarps developed near the apical of lateral branching, ovoid to spherical 400-520 µm in diameter, 500-650 µm long. Spermatangial stichidia developed below apical of lateral branching 25-60 µm in diameter, 120-220 µm long. (Fig.3 F-K).

Bostrychia radicosa (Itono) J. A. West, G. C. Zuccarello and M. H. Hommersand Thalli are about 0.5-1.5 cm high, reddish purple to brown. Thalli of erect, ventral on Species diversity and distribution of mangrove-associated red alga Bostrychia 61 creeping stolon alternate branching, polysiphonous throughout. Thalli are composed with two tier cells of 4 periaxial cells to each axial cells. Rhizoids borne from tier cells and can develop new erect shoots. Usually ecorticated cells but sometimes have a cortication development throughout are associated with season. Tetrasporangial stichidia developed below tips of lateral branching 0.1-2 mm long, 75-100 µm wide, Cystocarps branches borne within the polysiphonous of ultimate lateral branches, ovoid to spherical 300-450 µm in diameter, 350-500 µm long. Spermatangial stichidia no found. (Fig.4 A-F).

Bostrychia simpliciuscula Harvey ex J. Agardh Thalli are about 1-3 cm high, dark red, reddish purple to brown. Thalli postrate, alternate branching, The main axes polysiphonous throughout, ultimate branches monosiphonous at apical but polysiphonous at the base. Thalli are composed with two tier cells of 4-5 periaxial cells to each axial cells and attached by peripherohaptera on substrate. Usually ecorticated cells but sometimes have a cortication development throughout are associated with season. Tetrasporangial stichidia branches borne within the polysiphonous determinate branches 150-400 µm long, 80-160 µm in diameter, Cystocarps and Spermatangial stichidia on found. (Fig.4. G-L).

Bostrychia tenella (Lamouroux) J. Agardh. Thalli are about 1-5 cm high, dark red, reddish purple to brown. Thalli postrate, with 2-3 orders of alternate branching, The main axes with primary polysiphonous laterals and many monosiphonous laterals. Thalli are composed with two tier cells of 5-8 periaxial cells to each axial cells and attached by peripherohaptera on substrate, corticated with 1-4 layers. Tetrasporangial stichidia developed below tips 180-420 µm long, 20-160 µm in diameter of ultimate lateral branches, Cystocarps branches borne within the polysiphonous of ultimate lateral branches, ovoid to spherical 350-780 µm long, 300-600 µm in diameter and Spermatangial stichidia on found. (Fig.5. A-E).

Bostrychia binderi Harvey Thalli are about 1-5 cm high, dark red, reddish purple to brown. Thalli postrate, with 2-3 orders of alternate branching, The main axes heavily corticated primary laterals, secondary laterals, spine-like, with polysiphonous base and occasionally short monosiphonous tips. Thalli are composed with two tier cells of 5-8 periaxial cells to each axial cells and attached by peripherohaptera on substrate, corticated with 1-4 layers. Tetrasporangial stichidia developed below tips 160-400µm long, 20-180 µm in diameter of ultimate lateral branches, Cystocarps branches borne within the polysiphonous of ultimate lateral branches, ovoid to spherical 300-700 µm. long, 250- 500 µm in diameter and Spermatangial stichidia no found. (Fig.5. F-I).

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Fig. 2 Bostrychia calliptera, Bostrychia kelanensis. A-C. Bostrychia calliptera. A. Whole thallus. Scale bar 2 mm. B. Tetrasporophyte with tetrasporangial stichidia (arrows). Scale bar 0.5 mm. C. Peripherohaptera at node (arrowhead). Scale bar 50 µm. D-L. Bostrychia kelanensis. D. Whole thallus. Scale bar 3 mm. E. Tetrasporophyte with tetrasporangial stichidia. Scale bar 3 mm. F. Cystocarp with many carposporangia (arrowheads). Scale bar 100 µm. G. Lateral branch of male gametophyte with spermatangia stichidia (arrows). Scale bar 100 µm. H. Branch showing 3-tier cells per pericentral cell (arrowheads). Scale bar 20 µm. I. Apex of axis. Scale bar 100 µm. J. Many shoots arising from cladohaptera attachment disc. Scale bar 60 µm. K. Cladohaptera at node. Scale bar 20 µm. L. Branch with corticated (arrowhead). Scale bar 50 µm.

Species diversity and distribution of mangrove-associated red alga Bostrychia 63

Fig. 3 Bostrychia moritziana, Bostrychia radicans. A-E. Bostrychia moritziana. A. Aternate branching and ultimate branches monosiphonous (arrow). Scale bar 100 µm. B. Cladohaptera (arrow). Scale bar 50 µm. C. Lateral branch of female gametophyte with developing carposporophyte (arrow). Scale bar 100 µm. D. Lateral branch of male gametophyte with spermatangia stichidia (arrow). Scale bar 200 µm. E. Tetrasporophyte with tetrasporangial stichidia. Scale bar 150 µm (arrows). F-J. Bostrychia radicans. F. Female gametophyte with mature cystocarp (arrowheads). Scale bar 1 mm. G. Cladohaptera (arrow). Scale bar 50 µm. H. Branch showing 2 tier cells per pericentral cell (arrowheads). Scale bar 20 µm. I. Mature terminal cystocarp with many carposporangial. Scale bar 100 µm. J. Spermatangia stichidia (arrow). Scale bar 200 µm. K. Tetrasporophyte with tetrasporangial stichidia (arrow). Scale bar 100 µm.

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Fig. 4 Bostrychia radicosa, Bostrychia simpliciuscula. A-F. Bostrychia radicosa. A. Whole thallus. Scale bar 1 mm. B. Tetrasporophyte with tetrasporangial stichidia (arrow). Scale bar 100 µm. C. Branch showing 2 tier cells per pericentral cell. (arrowheads). Scale bar 20 µm. D. Rhizoids at node (arrow). Scale bar 40 µm. E. Apex of axis with corticated. Scale bar 50 µm. F. New shoots arising at node (arrow). Scale bar 70 µm. G-L. Bostrychia simpliciuscula. G, H. Branch with corticated. Scale bar 1 mm. I. Tetrasporophyte with tetrasporangial stichidia borne on lateral branches (arrow). Scale bar 1 mm. J. Alternate branching and ultimate branches monosiphonous (arrow). Scale bar 50 µm. K. Branch showing 2 tier cells per pericentral cell (arrowheads). Scale bar 10 µm. L. Peripherohaptera at node (arrow). Scale bar 20 µm.

Species diversity and distribution of mangrove-associated red alga Bostrychia 65

Fig. 5 Bostrychia tenella, Bostrychia binderi. A-E. Bostrychia tenella. A. Whole thallus. Scale bar 300 µm. B. Branch with polysiphonous primary branch and monosiphonous secondary lateral branches and heavily corticated primary laterals. Scale bar 25 µm. C. Peripherohaptera at node with many corticated (arrow). Scale bar 100 µm. D. Tetrasporophyte with tetrasporangial stichidia (arrow). Scale bar 60 µm. E. Female gametoptyte with mature cystocarp (arrow). Scale bar 75 µm. F-I. Bostrychia binderi. F,G. Branch short polysiphonous secondary lateral and heavily corticated primary laterals. F. Scale bar = 500 µm, G. Scale bar 50 µm. H. Female gametoptyte with mature cystocarp (arrow). Scale bar 0.5 mm. I. Tetrasporophyte with tetrasporangial stichidia (arrow). Scale bar 100 µm.

Assessment of Water Quality in the Studied Sites The chemical characteristics of an aquatic ecosystem is an important aspect that affects the distribution of Bostrychia, Caloglossa and Catenella species (Table 3). The trophic status of water in the studied sites was classified by some physical and chemical parameters including nitrate nitrogen, and soluble reactive phosphorus. The water salinity differed significantly between the Ranong, Chumphon, Surat Thani and Nakhon Si Thammarat sites (p < 0.05). In contrast, no significant differences in salinity were recorded between the Phangnga, Phuket and Krabi sites (p > 0.05).

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Water salinity during the month of April differed significantly from October and August with regard to sample time. Nitrogen concentrations in the water resulted in significant differences between the groups and within groups (p < 0.05). However, phosphorus concentrations were highest at the Krabi sites (p < 0.05).

Table 3: Water characteristics in the studied sites.

- 3- Characteristics Water Salinity (psu) NO3 (mg/L) PO4 (mg/L) Ranong 20.50 0.29 0.03 Phangnga 32.83 0.30 0.02 Phuket 32.83 0.32 0.02 Krabi 32.33 0.33 0.02 Chumphon 23.75 0.30 0.03 Surat Thani 25.00 0.29 0.04 Nakhon Si Tammarat 28.58 0.29 0.04

Mangrove Sediment Characteristics Sediment characteristics are important factors in the occurrence variations of Bostrychia species. The sediments of the mangrove sites were predominantly made up of clay-like silt. Differences were found among the three intertidal zones between the sites (p < 0.05) (Figure 6). Granules revealed significant differences in sediment type frequencies among the sites. Sand was found in Phuket resulting in significant differences between the study sites (p < 0.05). Silt was found in Phangnga resulting in significant differences between the study sites (p < 0.05). The sediment at Nakhon Si Thammarat was not significantly different than that at Ranong, Chumphon and Surat Thani (p > 0.05); and clay was not significantly different between the Phangnga, Phuket and Krabi sites. In contrast, significant differences were found between the Ranong, Chumphon, Surat Thani and Nakhon Si Thammarat sites (p < 0.05), while no significant differences were seen with regard to sampling time (p > 0.05).

Discussion Thailand is located in Southeast Asia, bordering the Andaman Sea and the Gulf of Thailand in the south. The upper southern area comprised of Ranong, Phangnga, Phuket and Krabi on the Andaman sea and Chumphon, Surat Thani and Nakhon Si Thammarat on the Gulf of Thailand differed in water salinity, sediments and food levels in this study. Thailand’s coastline has two different ecological regions that are influenced by their geographic setting, the Andaman coast (Indian Ocean) and the Gulf of Thailand coast (Pacific Ocean). Based on climate, light, temperature, rainfall, wind, most of the mangroves are only found in tropical zones where the tropical climate is suited for their growth (Duke et al. 1998[21]).

Species diversity and distribution of mangrove-associated red alga Bostrychia 67

Figure 6: Percentages of mangrove sediment components in intertidal zones of the sites (n =126).

Tides are the main factor influencing the habitat zones of macroalgae in the mangroves, because, tidal ranges influence the physical features of plant structures, especially, the aeration of roots above the soil which is important. Consequently, these factors affect pneumatophore height and density which in turn affects algal attachment, distribution and frequency of occurrence (Melville et al. 2005[22]). Waves and currents help spread the spores of the algae into other habitats, distribute nutrients to the sea that benefit other aquatic lives and coastal aquacultures. In addition, waves and currents are important factors in the coastal sedimentation process. However, nutrients, that are vital for the survival of organisms in the mangroves, are derived from both outside and inside the mangroves, including rain, rivers, sediments, the sea, and the decomposition of organic matter in the mangroves from algae as well as the salinity of water and soil, and these factors significantly affect the distribution of algae in the mangroves. Differences in salinity cause differences in the distributions of algae (Karsten et al. 1992[23], Fernandes et al. 2011[24]). In addition, mangrove soil results from the deposition process of sediments flowing with water from various sources as well as the decomposition of the organic matter in the mangrove forests. Sedimentation may have both direct and indirect impacts on macroalgal communities. Thus, soil is important for the growth and distribution of different species of mangrove trees and algal distribution and abundance in mangroves (Alongi et al. 1992[25]; Purcell and Bellwood, 2001[26]). The algal taxa primarily include Bostrychia, Caloglossa and Catenella (Dawes, 1996[27]). Red macroalgae associated with mangrove forests are commonly found in mangroves around the world. These algae usually growing abundantly are attached on roots of the Avicennia marina, Rhizophora and Xylocarpus in the mangrove areas by clusters of rhizoids. Algal diversity can also be quite high in mangrove environments.

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During this period the distribution of less tolerant algal communities was affected by adverse environmental conditions. Within each site sampling time, inter-site, and vertical factors did not appear to have a large influence on algal species distribution. For example, Bostrychia tenella, B. binderii, B. calliptera, B. kelanensis, B.radicans, B. moritziana, Caloglossa adherens and Catenella spp. displayed this common distribution feature throughout the study period (Steinke and Naidoo, 1990[28]), as did Bostrychia kelanensis, B.radicans, B. moritziana at all the sites studied. Similar findings have been reported in the Parramatta River area of Australia, Japanese mangroves (Tanaka and Chihara, 1987[11]) and Perequê and Sítio Grande mangroves (Yokoya et al. 1999[29]). Intertidal areas exposed to wave action show a distinct zonation pattern related to the gradient of exposure or submergence along the vertical slope (Melville et al. 2005[22]). In the present study the greatest frequencies of occurrence of Bostrychia tenella, B. binderii, B. calliptera, B. simpliciuscula and Caloglossa leprieurii were observed in the back zone. Previous research indicated that some taxa of Bostrychia and Caloglossa were found close to the water with an occasional sample found higher on the shore (Melville et al. 2005[22]). Besides, observations on northeastern Brazilian mangroves reported higher species diversity, due to the presence of species that are normally present on rocky shores (Miranda, 1986[30]). Moreover, Bostrychia kelanensis, B.radicans, B. moritziana, Caloglossa adherens and C.beccarii occurred very frequently throughout the intertidal zone (Pinheiro- Joventino and Lima-Verde 1988)[31]. Thus, salinity gradients create distinct ecotypes of Caloglossa in mangals along the Brisbane River, Australia (Mosisch, 1993)[32]. Tidal levels controlled the range of vertical distribution of macroalgae from Perequê mangroves while salinity variations were a determinant factor for the majority of the species from the Sítio Grande mangroves (Yokoyama et al. 2000)[33] and lower frequencies of occurrence for algal were observed in the front zone.

Acknowledgments We would like to thank Prof. Dr. Khanjanapaj Lewmanomont for valuable suggestions. Special thanks goes to the members of Laboratory of Applied Algae at Chiangmai University, Thailand. This research was partially supported by Phuket Rajabhat University, Thailand.

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