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The morphotaxonomy and phytogeographical distribution of sinuosa (Mertens ex Roth) Derbes & Solier (Scytosiphonales, Phaeophyta) fro....

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The morphotaxonomy and phytogeographical distribution of (Mertens ex Roth) Derbes & Solier (Scytosiphonales, Phaeophyta) from Myanmar

Khin Khin Gyi1 and Soe Htun2

Abstract The plants of the brown algal genus Colpomenia Derbes & Solier collected from the Rakhine Coastal Region and the Ayeyarwady Delta and Gulf of Martaban Coastal Region from 1971-2010 and deposited in the Department of Marine Science, Mawlamyine University in the forms of herbarium and wet-stack specimens, and fresh specimens collected from Setse coastal areas, were identified as Colpomenia sinuosa (Mertens ex Roth) Derbes & Solier, mainly based on the morphological studies. The distribution ranges of C. sinuosa (Mertens ex Roth) Derbes & Solier along the two Coastal Regions of Myanmar and the geographical regions of the world oceans were presented here. Some ecological notes and potential uses of this species were briefly described. Keywords: Colpomenia sinuous (Mertens ex Roth) Derbes & Solier, distribution, morphology, Myanmar, Phaeophyta, Scytosiphonales, .

Introduction There are 11 species in the genus Colpomenia (Endlicher) Derbes & Solier commonly occur in all oceans from tropical to temperate regions of the world: C. bullosa (Saunders) Yamada, C. durvillei (Bory de Saint-Vincent) Ramı´rez, C. ecuticulata Parsons, C. expansa (Saunders) Lee, C. mollis Taylor, C. nainativensis Durairatnam, C. peregrina Sauvageau, C. phaeodactyla Wynne et Norris, C. ramosa Taylor, C. sinuosa (Mertens ex Roth) Derbes & Solier, and C. tuberculata Saunders (Saunders 1898, Taylor 1945, Parsons 1982). Very recently, it was recorded 8 species of the genus Colpomenia, viz., C. bullosa, C. ecuticulata, C. expansa, C. phaeodactyla, C. peregrina, C. nainativensis, C. sinuosa, and C. tuberculata in the Asiatic waters (Guiry 2011, Boo et al. 2011). Among these species, 5 species of Colpomenia, namely C. ecuticulata, C. nainativensis, C. peregrina, C. sinuosa, and C.

1 Demonstrator, Department of Marine Science, Mawlamyine University 2 Professor and Head, Department of Marine Science, Mawlamyine University 2 Universities Research Journal 2012, Vol. 5, No. 1 tuberculata were reported from the Indian Ocean Region (IOR) (Silva et al. 1996). Interestingly, the only species of the genus Colpomenia, C. sinuosa (Mertens ex Roth) Derbes & Solier, occur in India (Misra 1966, Krishnamurthy and Joshi 1970, Jha et al. 2009), Bangladesh (Islam 1998), Thailand (Lewmanomont and Ogawa 1995 and Aungtonya and Liao 2002) and Malaysia (Phang 1998, 2006), although two species of the genus Colpomenia, C. sinousa (Mertens ex Roth) Derbes & Solier and C. nainativensis Durairatnam, commonly predominates along the coastal areas of Sri Lanka (Durairanam 1961, Guiry 2011), in the Bay of Bengal (Silva et al. 1996). In Myanmar, Kyi Win (1972), Kyaw Soe and Kyi Win (1977), Soe- Htun et al. (1997, 1999, 2009a) and Soe-Htun (1998, 2005) reported the only species in the genus Colpomenia, C. sinuosa (Mertens ex Roth) Derbes & Solier, from the Rakhine, the Ayeyarwady Delta and Gulf of Mottama (Martaban), and the Tanintharyi Coastal Regions. Recently, Hlaing Hlaing Htoon (2009) also recorded this species from Setse coastal areas. C. sinuosa is used as human food and then prepared as a salad due to the presence of antioxidant properties, a good source of phenols, vitamins, folic and folinic acids, and amino acids and can be used as animal feed, fertilizer and an indicator of metal pollution (Trono 2001). Likewise, forming a considerable amount of C. sinuosa in nature, it can be considered as a raw material for the extraction of alginic acid and alginates on commercial basis beside Sargassum tenerrimum (Levring, Hoppe and Schmid 1969, Rizvi 2003). The objectives of this study are: 1) to revise and update the systematics of the species of Colpomenia based on the morphology of vegetative and reproductive structures, and 2) to know the distribution ranges of this species along both the three Coastal Regions of Myanmar and the world oceans. Materials and Methods Voucher materials of the genus Colpomenia collected from the Rakhine and Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Regions were preserved in 4% formalin in seawater and lodged in the Herbarium of Department of Marine Science, Mawlamyine University, Mawlamyine, Myanmar (MMB) from 1971 to 2010, in the forms of herbarium and wet- stack specimens as well as the fresh specimens recently collected from the Universities Research Journal 2012, Vol. 5, No. 1 3 upper intertidal zones of Setse coastal areas (Lat. 15˚ 56′ N, Long. 97˚ 35′ E) in 2010, were used for identification. Specimens examined.– Tanintharyi Coastal Region: No Data; Ayeyarwady and Gulf of Mottama (Martaban) Coastal Region: Setse (Witt Yi Win, 4.iii.2006; MMB 05220; Win Win Aye, 4.iii.2006; MMB 05253; Myat Thway Myint Aung, 4.iii.2006; MMB 05224; Aung Min Htaik, 4.iii.2006; MMB 05260; Nyunt Sandar Aung, 4.iii.2006; MMB 05264; Myo Min Win, 4.iii.2006; MMB 05233; Thet Naung Soe, 4.iii.2006; MMB 05240; Htay Htay Lwin, 4.iii.2006; MMB 05249; Su Su Mon, 4.iii.2006; MMB 05246; Aung Myo Tun, 4.iii.2006; MMB 05274; Thein Tin Oo, 4.iii.2006; MMB 05283; Nay Zaw Aung, 4.iii.2006; MMB 05290; A Zan Ko; 4.iii.2006; MMB 05292; Mya Mya Thet, 4.iii.2006; MMB 05295; Ingyin, 4.iii.2006; MMB 05301; Myat Thu, 4.iii.2006; MMB 05306; Thuyein Tun, 4.iii.2006; MMB 05310; Aung Hlaing Oo, 4.iii.2006; MMB 05303; Group 3, 19.xii.1994; MMB 06533; 4th Year, 12.xii.1980; MMB 06534; Group 6, 12.xii.1980, MMB 06535; Yi Yi Cho, 12.xi.1979; MMB 01004, 06539; Nyunt Wai, 6.ii.1994; MMB 06543; Theikpan Soe, 6.ii.1994; MMB 06544; Win Myint, 7.ii.1994; MMB 06545; Khin Thandar Linn, 4.iii.2006; MMB 06587; 4th Year (Group 2), 20.xii.1994; MMB 07033; Group 4, 4.iv.2000; MMB 07034; Hla Kathy & Tin Zar Yu Yu Kyaw, 4.iv.2000; MMB 07035; Soe Thiha & Thu Thu Min, 4.iv.2000; MMB 07036; Group 3, 4.iv.2000; MMB 07037; Myo Thu, 4.iv.2000; MMB 0703; Group 2, 20.xii.1994; MMB 07039; Hlaing Hlaing Htoon, 15.x.2008; MMB 10180-10185; Khin Khin Gyi, 17.i.2010; MMB 10277-10281; 4.ii.2010; MMB 10282-10286: Vegetative; M.Sc. Students, 28.iv.2010; MMB 10511: Vegetative); Kyaikkhami (Mu Mu Aye & Myint Myint Than, 12.xii.1980; MMB 00855; Mu Mu Aye, 20.xi.1981; MMB 06540; Hla Tint, 26.ii.1981; MMB 0654: Vegetative); Rakhine Coastal Region: Grang Gyaing (Ma Yi Shwe, 4.ii1977; MMB 06538: Vegetative); Leik I. (Aung Myint, 25.ii.1967; MMB 00890: Vegetative); Cape Negrais (San San Oo, 2.iii.1984; MMB 06531: Vegetative); Chaungtha (Mu Mu Aye, 4.iii.2007; MMB 10276: Vegetative); Wetthey Gyaing (Maung Maung Nyunt, 4.iv.1996; MMB 03131: Vegetative); Phoe Htaung Gyaing (Soe-Htun, 3.iv.1996; MMB 03284: Vegetative); Lonetha (San Tha Tun, 18.iii.1987; MMB 03044: Vegetative); Ngapali (Aung Myint, 28.i.1977; MMB 06536: Vegetative); Makyee I. (Ramree) (Phyu Tun Aung, 8.i.1975; MMB 06537: Vegetative); Ta Kyet I. (Dept. of Botany, Mawlamyine College, 8.xii.1971; MMB 01009: Vegetative); Zin Chaung (Phyu Tun Aung, 2.iii.1975; MMB 4 Universities Research Journal 2012, Vol. 5, No. 1

01010, 06542: Vegetative); Kyauk Phyu (San Tha Tun, 25.ii.1987; MMB 03009: Vegetative); Taung Pyo (Dept. of Botany, Mawlamyine College, 30.xii.1971; MMB 06532: Vegetative). Sections of vegetative and reproductive structures of the materials were made by hand using double-edged razor blades. Microscopic measurements of the plants were recorded in micron meter (µm) by using the calibrated ocular meter attached to a BX–41 microscope (Olympus, Tokyo, Japan). External and internal morphologies of vegetative and reproductive structures were photographed with a Panasonic (Lumix) DMC- TZ 15 digital camera and processed using Adobe Photoshop 7.0. This study followed the classification system used by Guiry (2011). Local distribution of C. sinuosa was recorded from the collection sites of the specimens deposited in the Herbarium of Department of Marine Science, Mawlamyine University. Phytogeographical distribution and potential uses of C. sinuosa were consulted with worldwide published literature records available. Moreover, habitats and associated algae of C. sinuosa were also recorded in the field.

Results and Discussion A classification system of the genus Colpomenia Phylum: Phaeophyta (= Heterokontophyta) Class: Phaeophyceae Order: Scytosiphonales Family: Genus: Colpomenia (Endlicher) Derbes & Solier Species: C. sinuosa (Mertens ex Roth) Derbes & Solier

Colpomenia sinuosa (Mertens ex Roth) Derbes & Solier Saunders 1898: 164, pl. XXXII, figs. 7-8; Matsumdra 1904: 64; Boregesen 1914: 20-21, fig. 12; Britton and Millspaugh 1920: 590; Setchell and Gardner 1930: 143-144; Dawson 1944: 232-233; Dawson 1945: 31; Taylor 1945: 83-85; Segawa 1956: 37, pl. 21, fig. 169; 1968: 37, pl. 21, fig. 169; Arasaki 1964: 34, fig. 103; Taylor 1960: 260, pl. 36, fig. 1; Durairatnam 1961: 32-33, fig. 1; Dawson 1966: 97, 158, 160, fig. 9-10 A- H; Misra 1966: 115-116, fig. 59, Pl. II; Papenfuss 1968: 38; Pham-Hoang Universities Research Journal 2012, Vol. 5, No. 1 5

Ho 1969: 339-340, fig. 3.39; Smith 1969: 127-128, pl. 20, fig. 1; Chihara 1970: 37, pl. 19, fig. 2; 1975: 67; Tsuda 1972: 99-100; Abbott and Hollenberg 1976: 202-204, fig. 168; Wynne and Norris 1976: 2-5, fig. 1 a- b, fig. 2 a-b; Kyaw Soe and Kyi Win 1977: 82-83, figs. 137-138; Saraya and Trono 1979: 31, pl.9, fig. 4; Menez and Mathieson 1981: 8; Parsons 1982: 291-293, figs. 2-3 and figs. 11-12; Tseng 1983: 184, fig. 4, Pl. 93; Womersley 1987: 297, figs. 107 A, 108 E,F; Lewis and Norris 1987: 13; Lewmanomont and Ogawa 1995: 61; Silva et al. 1996: 627-628; Kogame 1997: 227-231, figs. 1-6; Trono 2001: 138-139, figs. 1-5; Abbott and Huisman 2004: 178-180, fig. 67A-B; Nunes and Paula 2004: 14-16, figs. 4- 6; Abbas and Shameel 2009: 1921-1926, figs. 1-4, figs. 5-8, figs. 9-10; Hlaing Hlaing Htoon 2009: 37-38; Jha et al. 2009: 80, figs. a-b; Guiry 2011. Synonyms.- Ulva sinuosa Mertens ex Roth 1806, Saunders 1898: 164. Tremella cerina Clemente 1807; Tremella rugosula Clemente 1807 (as cited in Guiry 2011), Enchoelium sinuosum (Mertens ex Roth) C. Agardh 1820; C. Agardh 1824: 262; Asperococcus sinuosus (Mertens ex Roth) Bory de Saint-Vincent 1832, Saunders 1898: 164; Stilophora vesicata Harvey 1834; Asperococcus sinuosus (C. Agardh) Zanardini 1841; sinuosus (Mertens) ex Roth) Zanardini 1843: 39; Hauck 1888 (1886-1889): 92; Enchoelium vesicatum (Harvey) Kutzing 1849; Soranthera leathesiformis P.L. Crouan et H.M. Crouan 1865 (as cited in Guiry 2011). Type locality.- Near Cadiz, (Silva et al. 1996: 627). Type. - BM (Guiry 2011). Morphology: Vegetative features.- Plants of Colpomenia sinuosa, are hollow, greenish brown to light-golden brown, solitary or clustered, hollow, sessile, spherical or variable in shape, becoming flattened, expanded and irregularly convoluted, individually reaching a diameter of 1.2-8.0 cm (Fig.1), growing on rocks or bottom surface in the upper intertidal zones, by mean of rhizoids or by a broad base. Small plants are approximately spherical, smooth and contorted in some portions filled with water when young but collapsed later. Vegetative thalli consist of rounded to hexagonal shaped cells, 60- 230 µm long and 60-180 µm broad with chromatophores in surface view (Fig. 2) and with 2 portions comprised 4-6 layers of cells, 280-360 µm in thickness, in cross section: (1) the cortex composed of 1-2 layered 6 Universities Research Journal 2012, Vol. 5, No. 1 epidermis, small, thin wall and consisting of short rectangular cells, 4-20 µm in length, 4-24 µm in width, with rounded corners and many chromatophores, and (2) the medulla, composed of 3-5 layers of large, colorless, rectangular and thin- walled cells of various shape and size with lacking pits, 28-220 µm in length, 16-80 µm in width, intercellular space absent, without chromatophores (Figs. 4-6). Several oil globules are found in the apical portion of the thallus (Fig. 7). Rhizoidal filaments are 20-180 µm long, forming in large groups on lower surfaces (Fig. 3). Reproductive features.- When matured, thalli formed reproductive structures as plurilocular zoidangia which are scattered singly or in groups on epidermal layer of the thallus (Fig.8), yellowish brown in color, with cylindrical, uniseriate or biseriate filaments, 25-30 μm in length, 7.5-9.0 μm in width and aggregated together (Figs. 9-10). Phytogeographic distribution: (i) Myanmar distribution.- Tanintharyi Coastal Region: No data; Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Region: Setse and Kyaikkhami; Rakhine Coastal Region: Grang Gyaing, Leik I., Cape Negrais, Chaungtha, Wetthey Gyaing, Phoe Htaung Gyaing, Lonetha, Ngapali, Makyee I., Ta Kyet I., Zin Chaung, Kyauk Phyu, Taung Pyo. (Fig. 11). (ii) Phytogeographical distribution.- Pacific Ocean: Hong Kong (Guiry 2010), China (Tseng 1983, Guiry 2011), Korea (Guiry 2011), Japan (Yamada 1950, Segawa 1956, 1968, Arasaki 1964, Chihara 1970, 1975, Kogame 1997,Guiry 2011), Taiwan (Lewis and Norris 1987, Guiry 2011), South Australia, New South (Guiry 2011), Queensland (Cribb 1961,Guiry 2011), Lord Howe I., New Zealand, Chatham I. (Guiry 2011), Hawaii I. (Abbott and Huisman 2004, Guiry 2011), French Polynesia, Mariana I. (Guiry 2011), Micronesia (Tsuda 2006, Guidry 2011), Fiji I. (Guiry 2011); California (Abbott and Hollenberg 1976, Wynne and Norris 1976, Guiry 2011), Mexico (Taylor 1960, Ortega et al. 2001, Guiry 2011), Galapagos I. (Guiry 2011), Colombia (Daiz-Pulido & Daiz-Ruiz 2003), Peru, Chile (Guiry 2011); Atlantic Ocean: Florida (Taylor 1960, Guiry 2011), Georgia (Guiry 2011), North Carolina (Taylor 1960, Guiry 2011); Costa Rica (Guiry 2011), Panama (Taylor 1960, Guiry 2011), Uruguay (Guiry 2011), Brazil (Taylor 1960, Nunes and Paula 2004, Guiry 2011), Venezuela (Taylor 1960, Guiry 2011), Trinidad & Tobago (Taylor 1960, Guiry 2011), Barbados (Guiry 2011), Virgin I., Puerto Rico (Taylor 1960, Universities Research Journal 2012, Vol. 5, No. 1 7

Guiry 2011), Lesser Antilles, Caribbean (Guiry 2011), Bahamas, Jamaica (Taylor 1960, Guiry 2011), Trinidad (Guiry 2011), Cuba (Taylor 1960, Guiry 2011), , , France, Spain, , Malta, Italy, Adriatic Sea, Greece, Turkey, Madeira, Canary I., Cape Verde I., Morocco, Algeria, Tunisia, Libya, Mauritania, Senegal, Guinea, Sierra Leone, Ghana, Togo, Equatorial Guinea, São Tome & Príncipe, Gabon, Angola (Guiry 2011); Indian Ocean: Egypt, Sudan, Yemen, Saudi Arabia, Oman, Bahrain, Kuwait, Iran, Ethiopia, Somalia, Kenya, Tanzania, Mozambique, South Africa, Madagascar, Seychelles, Laccadive I., (Guiry 2011), Pakistan (Abbas and Shameel 2009, Guiry 2011), India (Misra 1966, Silva et al.1996, Krishnamurthy and Joshi 1970, Jha et al. 2009,Guiry 2011), Andaman I. (Guiry 2011), Sri Lanka (Durairatnam 1961, Guiry 2011), Bangladesh (Islam 1998, Guiry 2011), Myanmar (Kyaw Soe and Kyi Win 1977, Soe-Htun 1998, Soe-Htun et al. 2009, Present study), Thailand (Lewmanomont and Ogawa 1995, Aungtonya and Liao 2002); Indo-Pacific Region: Malaysia (Phang 1998, 2006, Guiry 2011), Singapore, Vietnam (Guiry 2011), Philippines (Saraya and Trono 1979, Silva et al. 1987, Guiry 2011), Indonesia (Guiry 2011), Western Australia (Huisman and Borowitzka 2003, Guiry 2011) (Fig. 12). Ecological notes.- This species attaches to rocky substrates and the pneumatophores of mangrove trees such as Sonneratia graffithii and Avicennia marina, in intertidal zones subjected to slow to moderate water movement and is commonly found from November to April. The thalli are easily removed from the substrate by strong wave action and may be transported as drift materials. Associated algal species are Acanthophora spicifrea and Gracilaria canaliculata. Potential uses.- Colpomenia sinuosa can be considered as a raw material for the production of alginic acid and alginates (Levring et al. 1969). It is used as human food, as salad, and animal feed, due to the presence of antioxidant properties, a good source of phenols, vitamins, folic and folinic acids, and amino acids such as alanine, aspartic acid, glutamic acid, glutamine, glycine, serine, threonine, tyrosine and valine, and fertilizer as well as indicator of metal pollution (Trono 2001). In addition, C. sinuosa contains 9.7-15.8% protein, 0.57-1.33% lipids, 4.4-5.98% crude fibre, 0.0021-0.0815% of iodine, 20.19-25.8 % of ash, 11.9-15.4 % of moisture, 5.59-7.7% of sulphate, 34.93-41.26% of carbohydrate (Nyo Nyo Tun 1999). The yields of alginic acid and sodium alginate extracted from C. sinuosa 8 Universities Research Journal 2012, Vol. 5, No. 1 from Setse varied from 7.0 to 11.5 and from 6.60 to 10.95 %. The viscosity value of 1% sodium alginate solution varied from 166 to 250 cps (Myint Myint Cho 2011). C. sinuosa contains 28623 ppm of Ca, 12991 ppm of Fe, 19263 ppm of K, 10028 ppm of Mg, 330 ppm of Mn, 5700 ppm of Na, 147 ppm of Zn, 1.7 ppm of Pb, 0.8 ppm of Cd and 1.9 ppm of Hg, respectively (War War Nwe 2011). Taxonomic assessment on Colpomenia sinuosa (Mertens ex Roth) Derbes & Solier Silva et al. (1996) had reported 5 species of Colpomenia, namely C. ecuticulata, C. nainativensis, C. peregrina, C. sinuosa, and C. tuberculata from the Indian Ocean Region (IOR). Among these, stalk is uniquely formed in C. bullosa which possess 1-3 hollow sacs arising from a crustose lobed base by with thickened medullary cells of 5-9 layers with pits, but not in C. sinuosa which is characterized by sessile globular or folded and convoluted thallus with thin walled medullary cells of 3-5 layers, lacking pits and sorus with cuticle, punctuate, usually with a central hair pit, C. peregrina that is formed globose or vesicle liked thalli with 2-5 medullary layers of large colorless, thin-walled cells lacking pits, C. ecuticulata was distinguished by its slightly folded, globose or vesicle liked thalli with 3-4 medullary layers of large colorless, thin-walled cells lacking pits and C. nainativensis which shows irregularly lobed thallus hollow with spine-like projections and 3-layered medullary cells (Parsons 1982, Boo et al. 2011). In the present study, the gross morphology of C. sinuosa agrees well with the descriptions of specimens described by Smith (1969: 127-128, pl. 20, fig. 1), Saraya and Trono (1979: 31, pl. 9, fig. 4), Tseng (1983: 184, fig. 4, pl. 93), Lewmanomont and Ogawa (1995: 67), Abbas and Shameel (2009: 1921-1926, figs. 1-4, figs. 5-8, figs. 9-10) and Jha et al. (2009: 80, figs. a-b). The size of thalli in Myanmar plants is variable, reaching about 1.2-8.0 cm in diameter, as compared with the plants found in other countries showing around 8-12 cm in diameter (Taylor 1960, Smith 1969, Saraya and Trono 1979, Abbott and Huisman 2004). The habitat of the plants of C. sinuosa is closely similar to that of the plants in India (Misra 1966; Jha et al. 2009), the Philippines (Saraya and Trono 1979), Indonesia (Trono 2001), and Hawaii (Abbott and Huisman (2004). The present observations on numbers of layers in the cortex, size and shape of cells in C. sinuosa closely corresponded to the observations on those of Parson (1987), Womersley (1987) and Boo (2011) but not in the Universities Research Journal 2012, Vol. 5, No. 1 9 plants which composed of 3-7 layers of cortical cells from Pakistan (Abbas and Shameel 2009). In this study, the medulla composes of 3-5 layers of large, rectangular and thin-walled cells of various shape and size, 28-220 μm in length, 16-80 μm in width, intercellular space absent without chromatophores (Fig. 4-6) but mostly over 5 layers with relatively thick- walled cells in Hawaiian specimens (Abbott and Hollenberg 1976, Abbott and Huisman 2004), five to six layered cells in Indian specimens (Misra 1966, Jha et al. 2009) and 5-8 rows of cells in the Philippines’s specimens (Saraya and Trono 1979). The morphology of inner cells which are normally large and colorless, progressively smaller and more pigmented outwardly is closely similar to that of inner cells described by Abbott and Hollenberg (1976), Kyaw Soe and Kyi Win (1977) and Abbott and Huisman (2004). The morphology of rhizoidal filaments, 20-180 μm long, forming in a large group on the lower surface (Fig. 3), is nearly the same as the description of that mentioned by Misra (1966) and Abbott and Huisman (2004). The species of Colpomenia belonging to the Family Scytosiphonaceae, showed a heteromorphic life-history pattern with an alternation of generation between gametophytic, erect parenchymatous macrothalli (bearing plurilocular zoidangia and sporophytic, prostrate pseudoparenchymatous microthalli bearing ectocarpoid plurilocular and unilocular zoidangia. Plurizoids released from erect macrothalli developed into prostrate microthalli. Swarmers released from unilocualr zoidangia of prostrate microthalli directly developed into erect macrothalli (Kogame 1997, Cho et al. 2001, Trono 2001). In the present study, the reproductive structures in the forms of plurilocular zoidangia (= plurilocular gametangia) (25-30 μm in length, 7.5-9 μm in breath) are normally found in matured macrothalli of C. sinuosa as described by (Taylor 1960), Durairatnam (1961), Misra (1966), Dawson (1966), Tseng (1983), Kogame (1997), Reine and Trono (2002), Abbott and Huisman (2004), Nunes and Paula (2004), Abbas and Shameel (2009) and Jha et al. (2009). Sporophytic phase as in the form of microthalli in C. sinuosa which will be dominant in the monsoon season from June to October is still unknown for Myanmar specimens. 10 Universities Research Journal 2012, Vol. 5, No. 1

1 2 3

1 cm 50 µm 60 µm 4 5 c c

m m

100 µm 100 µm 6 7 c c

m

100 µm 100 µm

8 9 10

50 µm 5 µm 10 µm

Figs.1-10. Morphology and anatomy of Colpomenia sinuosa (Mertens ex Roth) Derbes et Solier: 1) Habit of a plant (MMB 10277); 2) Surface view of the thallus, showing rounded or hexagonal shaped cells (MMB 10278); 3) Rhizoidal filaments (arrow) of C. sinuosa (MMB 10279); 4) Cross section of thallus showing 3 layers of medulla (m) and 1-2 layers of cortex (c) cells (MMB 10282); 5) Cross section of thallus showing 4 layers of medulla (m) and 1-2 layers of cortex (c) cells (MMB 10283); 6) Cross section of thallus showing 5 layers of medulla (m) and 1-2 layers of cortex (c) cells (MMB 10284); 7) Oil Universities Research Journal 2012, Vol. 5, No. 1 11

globules (arrowheads) (MMB 10285). Figs. 8-10. Formation of reproductive structures in C. sinuosa: 8) Sori on surface of thallus, with hair pits (arrow) immediately surrounded by the darker paraphyses; 9-10) Ectocarpoid plurilocular zoidangia (arrowhead) in cross section of thallus.

Moreover, the molecular results of neighbor-joining tree (NJ), maximum- likelihood tree (ML) and maximum parsimonious tree (MP), based on sequences of plastid-encoded RuBisCo spacer region revealed that C. sinuosa which was allied with Hydroclacthrus clathratus, differed from C. claytonia which was sister to the above clade and the monophyly of C. bullosa which was allied with Petalonia binghamiae, P. fascia, Scytosiphon dotyii, S. gracilis and S. lomentaria, showing polyphyly of the genus Colpomenia (Cho et al. 2001). In addition, Cho et al. (2005) demonstrated that Colpomenia sinuosa mostly encountered in subtropical to tropical waters, was consistently the sister taxon to C. claytonia normally occurred in temperate to subtropical waters, in their rbcL tree derived from analysis of the ITS sequences of both species. In Myanmar, the local distribution of C. sinuosa (Mertens ex Roth) Derbes & Solier, Kyaw Soe and Kyi Win (1977) was firstly reported an account of this species from Ngapali and Mawtin Point (the Rakhine Coastal Region), Kyaikkhami (the Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Region) and Pine Tree Is. (the Tanintharyi Coastal Region) and Soe-Htun et al. (1997) also recorded that the presence of C. sinuosa which occurred along all the three Coastal Regions of Myanmar. Soe-Htun (1998) described that this species distributed along the Rakhine and Tanintharyi Coastal Regions except for the Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Region. However, Soe-Htun et al. (1999) also reported the horizontal distribution of C. sinuosa along the Kyaikkhami and Setse coastal areas of the Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Region. Moreover, Soe-Htun (2005) and Soe-Htun et al. (2009) documented occurrence of the species of C. sinuosa along the three Coastal Regions of Myanmar. Furthermore, Hlaing Hlaing Htoon (2009), Sein Moh Moh Khine (2012) recorded the presence of C. sinuosa (Mertens ex Roth) Derbes & Solier from Setse coastal areas. 12 Universities Research Journal 2012, Vol. 5, No. 1

16 94˚ 96˚ N M Y A N M A R 20˚ 20˚

15 T H A I L A N D 14 12 13 11 18˚ Bay 10 9 8 of 4 7 3 16˚ 6 16˚ 5 2 Bengal Andaman 14˚ Sea

12˚ 100 0 100 1 10˚N km 10˚ 92˚E 94˚ 96˚ 98˚

Fig. 11. Distribution of Colpomenia sinuosa (Mertens ex Roth) Derbes et Solier along Myanmar coastal areas: 1. Pine Tree I., 2. Setse, 3. Kyaikkhami, 4. Bilugyun I., 5. Leik I. (Diamond I.), 6. Mawtin point, 7. Cape Negrais, 8. Chaungtha, 9. Wetthey Gyaing, 10. Phoehtaung Gyaing, 11. Lonetha, 12. Ngapali, 13. Makyee I., 14. Zin Chaung, 15. Kyauk Phyu, 16. Taung Pyo. Symbols: Potential distribution range; Southern- most and Northernmost recorded limits; Present study, Previous records (Kyaw Soe and Kyi Win 1977). Universities Research Journal 2012, Vol. 5, No. 1 13

In the present study, it is noted that C. sinuosa distributes along the two Coastal Regions of Myanmar: Rakhine Coastal Region, from Leik I. (Diamond I.) (Lat. 15˚ 51′ N, Long. 94˚ 17′ E) to Taung Pyo (Lat. 21˚ 13′ N, Long. 92˚15′ E) and the Ayeyarwady Delta and Gulf of Mottama (Martaban) Coastal Region, from rocky shores of Setse Yele Pagoda (Lat. 15˚ 56′ N, Long. 97˚ 35′ E) to rocky shores of Kyaikkhami Yele Pagoda (Lat. 16˚ 04′ N, Long. 97˚33′ E) (Fig. 11), according to the specimens deposited at the Herbarium of the Department of Marine Science, Mawlamyine University (MMB). The local distribution range of C. sinuosa is very wide from the southern limit of Pine Tree Island of the Tanintharyi Coastal Region to the northern limit of Taung Pyo of the Rakhine Coastal Region (Table. 1). However, the distribution of C. sinuosa along the Tanintharyi Coastal Region is not known, due to lack of voucher specimens previously recorded by Kyaw Soe and Kyi Win (1977) and Soe-Htun et al. (1997), in the Herbarium of the Department of Marine Science, Mawlamyine University until now. C. sinuosa can be encountered along the Tanintharyi Coastal Region, facing the Andaman Islands where the plants of C. sinuosa are encountered (Silva et al. 1996). However, there were no plants observed in Kampani coastal areas, the northern part of the Tanintharyi Coastal Region (Myo Min Tun 2012). Further collections of C. sinuosa growing along the three Coastal Regions of Myanmar are still required to confirm the range of distribution of this species in Myanmar.

Table 1. The distributional range of Colpomenia sinuosa (Mertens ex Roth) Derbes & Solier in the three Coastal Regions of Myanmar. Coastal Regions Sr. TCR ACR RCR No. From To From To From To 1. Pine Tree I. No data Setse Bilugyun I. Leik I. Taung pyo (Lat. 10º (Lat. 15º (Lat. 16º (Lat. 15º (Lat. 21º 20′ N, 56′ N, 13′ N, 52′ N, 13′ N, Long. 98º Long. 97º Long. 97º Long. 94º Long. 92º 17′ E) 35′ E) 32′ E) 15′ E) 15′ E) 14 Universities Research Journal 2012, Vol. 5, No. 1

The very wide distributional ranges of C. sinuosa occurs lattitudinally from Norway of the Atlantic Ocean as northernmost recorded limits to Chatham I. (New Zealand) of the Pacific Ocean as southernmost recorded limits (Fig. 12). In the Pacific Ocean, there are the only species of Colpomenia, C. sinuosa, in Malaysia (Phang 1998, 2006, Guiry 2011), Vietnam (Pham-Hoang-Ho 1969, Guiry 2011) and the Philippines (Saraya and Trono 1979, Silva et al. 1987, 1996, Guiry 2011), two species, C. bullosa, and C. sinuosa in China (Tseng 1983), four species such as C. bullosa, C. phaeodactyla, C. peregrina and C. sinuosa, in Japan (Yamada 1950, Segawa 1956, 1968, Arasaki 1964, Chihara 1970, 1975, Yoshida et al. 1985, 1990, 1995 and Kogame and Yamagishi 1997), six speices, viz., C. bullosa, C. peregrina, C. ecuticulata, C. phaeodactyla, C. expansa and C. sinuosa in Korea and one species, C. peregrina also in Russia (Guiry 2011). In geneal, C. peregrina is normally distributed in temperate to subtropical regions, whereas C. sinuosa widely predominates from tropical to temperate coastal regions in all oceans (Womesley 1987, Kogame 1997, Cho et al. 2001).

130°W 80° 30°W 20°E 70°E 120°E 170°E

60° 60°N N Atlantic Ocean 10°N Pacific 10°N 0 Ocean 0 Indian 40°S Ocean 40°S

90 ° 130° 80°W 30°W 20° 70° 120°E 170° W E E E Fig. 12. Phytogeographical distribution of Colpomenia sinuosa (Mertens ex Roth) Derbes et Solier. Symbols: Potential distribution range; Northernmost and Southernmost recorded limits of C. sinuosa. Universities Research Journal 2012, Vol. 5, No. 1 15

With regard to phylogenetic relationship among these two species, Cho et al. (2005) suggested that C. peregrina originated at nothern margin of the taxon that gave rise to C. sinuosa. Hence, this study confirmed the occurrence of the species of Colpomenia in Myanmar waters as C. sinuosa in its taxonomic distinctness based on vegetative and reproductive morphology, and its distribution patterns in the Indian Ocean Region (IOR). In general, this species widely distributed along the tropical, subtropical, temperate and subtemperate regions of the world oceans. Further studies on the reproductive biology of C. sinuosa from Myanmar are still needed to be done to know the complete life history of the plants growing in nature.

Conclusions

This study provided morphology-based taxonomy and distribution of Colpomenia sinuosa (Mertens ex Roth) Derbes & Solier The occurrence of the only species of the genus C. sinuosa in Myanmar was confirmed. The main characteristics of C. sinuosa collected from Myanmar are observed as follows: (1) Plants are spherical or globular, solitary or clustered, sessile, commonly epiphytic, and attached by means of rhizoids formed in large groups on lower surfaces, and greenish brown to light golden brown, growing on the rocky and muddy surface along the upper intertidal region, both in exposed and sheltered situations. (2) Small plants are solid at first, later becoming hollow, becoming lobed and water filled-sac or irregularly expanded and large plants variable in shape, individually reaching a diameter of 1.2-8.0 cm. (3) Thallus consists of 4-6 layers, 280-360 µm thick, the outermost layer closely packed pigmented cortical cells, 4-20 µm long and 4-24 µm broad and inner-layered cells becoming larger inwardly, 28-220 µm long and 16-80 µm broad, thick wall, without intercellular space. (4) Several prominent oil globules are remarkably found in the apical portions of the thallus. (5) Plurilocular zoidangia present in groups and scattered on surface of the thallus, yellowish brown in color, arising from epidermal layer and 16 Universities Research Journal 2012, Vol. 5, No. 1

forming as sori aggregated by cylindrical uniseriate or biseriate filaments measured 25-30 μm in length and 7.5-9 μm in width.

The present study reveals that Colpomenia sinuosa distributes along both Rakhine and Ayeyarwady Coastal Regions except for the former record which showed the occurrence of this species from the Tanintharyi Coastal Region. In general, C. sinuosa widely predominates from tropical to temperate coastal regions of the world oceans.

Acknowledgements We are indebted to Dr. Htay Aung, Rector of Mawlamyine University, for his encouragement and supports in preparing this work. We are very grateful to Dr. Min-Thein, Director (Retd), Ministry of Industry (1), Myanmar Pharmaceutical Factory (MPF), Sagaing, Myanmar for his valuable suggestions and constructive criticisms on this study and literature provided. We would like to express our sincere thanks to the local people who kindly help us in many ways during our field trips. Many thanks go to Daw Soe Pa Pa Kyaw and Mu Mu Aye, Assistant Lecturer and Lecturer of the Department of Marine Science, Mawlamyine University, for her assistance in preparations of the manuscript. The first author, Khin Khin Gyi, would like to thank her beloved parents, Professor Dr. Khin Maung Cho, Head of Department of Marine Science, Pathein University and Dr. Myint Myint Cho, Professor, Department of Marine Science, Mawlamyine University for their physical, moral and financial supports throughout this study and also to her best friends, Wah Wah Shwe and Cho Cho Latt, for their help and encouragement during this research work. Thanks are also due to all my respected teachers and colleagues for their encouragement. In addition, funding for this work from the Department of Higher Education (Lower Myanmar), the Ministry of Education and the Department of Marine Science, Mawlamyine University is also mostly appreciated.

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