Marine algal flora of Hainan Island: a comprehensive synthesis

Tamara V. Titlyanova, Eduard A. Titlyanov and Tatyana L. Kalita

Coastal Ecosystems, 2014, vol 1, 28-53

© 2014 Society for Coastal Ecosystems Studies - Asia Pacific Titlyanov E et al. Algal flora of Hainan Island

Data Paper

Marine algal flora of Hainan Island: a comprehensive synthesis

Tamara V. Titlyanova, Eduard A. Titlyanov and Tatyana L. Kalita

A.V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, 690041, Russia Corresponding author : TV. Titlyanova, e-mail: [email protected]

Abstract

The present report contributes to the knowledge of the marine algal flora in northern part of the South China Sea. On the basis of literature data and own sampling, an annotated account of the of Hainan Island and nearby islets is presented. Three sets of algal collec- tions are involved: the first series of collection by Tseng and colleagues between the 1930s and 1970s (344 species), the second collection through two German-Chinese expeditions in October−December 1990 and March−April 1992 (214 species) and the third by the authors in 2008−2012 (251 species). Decadal changes in the benthic flora between the 1930s−1970s and 1990s resulted in a decline of epilithic with voluminous forms mainly of the families Liagoraceae (Rhodophyta, Rh), (, Oc), Caulerpaceae and Codiaceae (, Ch) and in an increase of epiphytic species with fine filamentous and membranous thalli forms: the families Ceramiaceae, Rhodomelaceae (Rh), Ulvaceae, Cladophoraceae (Ch). Changes in marine flora from the 1990s to 2008−2012 showed the same tendency. It is suspected that these historical changes in the marine flora occurred due to human impacts on the coastal ecosystems (in particular, coral reefs) of the island, includ- ing over-exploitation of resources and eutrophication of shallow waters, as well as natural catastrophes such as coral bleaching events of 1998.

Keywords: decadal changes, Rhodophyta, Ochrophyta, Chlorophyta, human impact

Introduction (SSS) fluctuates between 33.3 and 34.0 psu. However, SSS is strongly seasonal at Hainan Island, decreasing to a mean Hainan Island (Fig. 1) is located in the tropical northern value of 26.5 psu owing to freshwater run-off and rain during periphery of the Indo-Pacific Ocean in the South China Sea, the wet summer season. The mean tidal range is generally between 20°10’N, 110°41’E and 18°9’N, 109°34’E. Climate less than 1.5 m. The rainy season in the southern part of is subequatorial; the annual mean temperature is +24°C, Hainan occurs from May to October and accounts for 95% of with sunny weather during more than 300 days in a year. the yearly rainfall; the dry season occurs from November to The annual mean sea surface temperature (SST) is 26°C April. The island has an area of 33.920 km2 and a coastline with an average seasonal range of 12.1°C. The annual SST of more than 1600 km (after Titlyanov et al. 2011a, 2014а). maximum (30.8°C) and minimum (18.7°C) commonly occur Four main marine ecosystems such as coral reefs, seagrass in July and January, respectively. Mean sea surface salinity beds, estuaries and mangrove swamps border the coasts of

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Hainan Island. Coral reefs are the major coastal ecosystem in the intertidal and shallow subtidal zones of exposed and semiprotected areas along the shore. Seagrass beds mainly occupy the soft bottom of semi-protected bays, and mangrove swamps occupy brackish water lagoons connected with the sea. The highest biodiversity of the main coastal ecosystems was recorded between the 1950s and 1960s (Gurianova 1959) but decreased severely during the next 20 years (Hutchings & Wu 1987, Zhang et al. 2006). Almost 80% of the fringing reefs along the coastline of Hainan Island have been damaged due to human activities during the 1970s−1990s (fishing with dynamite, removal of corals for lime and construction). Recently, the eutrophication of Hainan coastal waters, particularly in the shallow bays, has increased due to greater tourist numbers, construction works along the coast, and mariculture in coastal ponds and pools with wastes draining into the sea (Hutchings & Wu 1987, Fiege, Neumann & Jinhe 1994, Zhang 1996, Zhang et al. 2004, Tadashi et al. 2008, Titlyanov et al. 2011b). Three major algal collections were conducted on Hainan Island between the 1930s and 2012. The first sampling campaign of marine seaweeds was performed by Tseng and colleagues in the 1930s at 19 localities on Hainan Island, followed by another series of sampling in Hainan including Xisha Islands between the late 1950s and the 1970s (Fig. 1A). This collection was processed by Chinese phycologists and deposited in the herbarium of the Institute of Oceanology of the Chinese Academy of Sciences (Tseng 2004). All publications relating to these collections were Figure 1. Collection sites at Hainan Island. A. Collection considered here: Tseng 1935, 1936, 1937, 1938, 1941a, b, sites of Tseng and coworkers in 1933–1935 with old spellings of names (after Tseng, 1936). B. Collection 1942a, b, 1983, 2004, 2009, Tseng & Gilbert 1942, Tseng sites during 1990/1992 (small dots, after Titlyanov et al. & Chang 1962, Tseng et al. 1962, 1980, 2001, 2005, 2011, 2011a) and 2008−2012 (large dots). Chang & Xia 1963, 1976, Tseng & Dong 1978, Zhang & Xia 1979, 1980, 1981, 1983, 1984, 1988a, b, c 1991, 1994, Zheng 1980, Zhu & Liu 1980, Lu & Tseng 1980, Xia & Zhang 1982, 1999, Tseng & Lu 1983, 1988, 1992, 1995a, b, c, identified by B. Xia and I. Bartsch in 1990−1994 and by T. 1997, 1999, 2000, 2002a, b, c, d; Xia & Abbott 1985, 1987, Titlyanova and E. Titlyanov in 2007−2009. On the basis Xia 1985, 2004, Dong & Tseng 1985, Xia & Yamamoto 1985, of these materials, the following papers were published: Doty 1988, Abbott, Zhang & Xia 1991, Chiang et al. 1992, Bischoff et al. 1994, Fiege, Neumann & Jinhe 1994, Titlyanov Masuda, Zhang & Xia 1994a, b, Luan & Luan 1995, Xia & et al. 2011a, Titlyanova et al. 2012 and one paper is in press Wang 1997, 1999a, b, 2000a, b, 2002, Millar & Xia 1999, (Titlyanov et al. 2014b). Tseng & Xia 1999, Zheng et al. 2001, Xia, Tseng & Wang The third series of collection was conducted in October 2004, Xia et al. 2004), Xia et al. 2002, Zeng et al. 2005, Ding 2008, April 2009, December 2010 and February-April 2012 et al. 2007, Liu 2008, Zheng & Li 2009. by T. Titlyanova, E. Titlyanov and Li Xiu Bao on the southern, In October−December 1990 and March−April 1992, eastern and northern shores of Hainan Island, including 8 the second major sampling campaign was conducted in localities: Luhuitou, Xiaodong Hai, Dadong Hai, Yalong Wan, the shallow waters of 16 localities (Fig. 1В) through two Wenchang, Haikou, Meixia and Xianhai (Fig. 1В). Algae German-Chinese expeditions. Algal samples were prepared were sampled from the splash, intertidal and shallow subtidal as herbarium specimens by B. Xia, I. Bartsch and B. Bischof (up to 5 m depth) zones. Results were partially published and one set of herbarium samples each (for all species/ (Titlyanov et al. 2011a, b, 2014 a, b, c, Titlyanov & Titlyanova forms) was deposited at the Alfred-Wegener-Institute for 2012, 2013). Polar and Marine Research (Germany) and at the Institute The purpose of the present study is to make an of Oceanology in Qingdao (China). Herbarium samples were annotated account of the seaweeds (Chlorophyta,

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Ochrophyta and Rhodophyta) of Hainan Island and generally rich in species. The low water mark, however, was nearby islets using all published information (1935−2014) characterized by dead coral blocks. Sandy beaches with plus our unpublished data of the last investigation in interspersed dead coral blocks were present at Xianhai. 2012. On the basis of earlier published works and this Poorly developed mangrove areas with extensive mudflats annotated account, some features of decadal changes and seagrass fields were present at Qukou and Qinglan in the marine flora of the island are noted, with a Gang. Near Haikou, algae were collected from fishponds discussion of possible reasons behind the changes. with variable salinity. At Meixia in the north of the island, extensive stony intertidal areas with offshore reef flats mostly comprising dead corals were dominant. An extensive reef flat Materials and Methods with many living corals above or between dead corals was located at Nanmai, a few hundred meters west of Meixia. Characteristic of sampling locations In 2008−2012, Luhuitou and Dadong Hai were characterized by damaged coral reefs (projective cover of In earlier publications on the marine flora of Hainan Island, live corals amounted from 30 to 50%, respectively), whereas descriptions of sampling locations were practically absent. Xiaodong Hai was characterized mainly by dead coral reefs For the 1990/1992 collection, sampling localities were and seagrass beds. The most studied site during this period partially described (Titlyanov et al. 2011a) according to the was the Luhuitou coral reef, situated opposite the Marine materials of the German-Chinese expeditions and personal Biological Station (South China Sea Institute of Oceanology, communications of the expedition participants (Inka Bartsch Chinese Academy of Sciences) (Fig. 2−4). and Xia Bangmei). During the 1990/1992 the intertidal/ The Xipai site (small rocky islet) in Yalong Wan Bay infralittoral reef flats (Linchang, Yalong Wan, Dadong Hai, was characterized by sandy substrates with unstable blocks Xiaodong Hai and Luhuitou) were characterized by unstable of dead corals, stones or boulders and rocky substrates blocks of dead corals; sometimes stones or boulders were near the islet. Algae were collected from intertidal rocks at interspersed with sandy areas. The offshore island of Ximao Wenchang and from materials cast ashore and fishery nets Zhou was the only place where the amount of living corals which were placed on (mostly dead) coral reefs at Xianhai. increased with depth in the shallow subtidal, and was The Meixia site was characterized by an extensive stony

Figure 2. The intertidal zone at Luhuitou Peninsula, April 2009. Inset: algal turf community in the middle intertidal zone (see Titlyanov et al. 2014d for details).

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Figure 3. The intertidal zone of coral reef at Luhuitou Peninsula, April 2009. Inset: algal turf community at low intertidal (for details, see Titlyanov et al. 2014d).

Figure 4. The upper subtidal zone at coral reef of Luhuitou Peninsula, March 2012. Inset: spp. community (for details, see Titlyanov et al. 2014a)

© 2014 SCESAP - 31 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island intertidal area (cape site), a sandy beach and the upper Jaasund 1976, Reyes 1976, Abbott & Hollenberg 1976, intertidal fringed by offshore reef flat mostly comprising dead Vinogradova 1979, Perestenko 1980, Trono & Put 1982, corals and rare live colonies. In March 2012, during sampling Lewis & Norris 1987, Santelices, 1988, Trono & Ganzon- at Meixia, a green algal bloom (Monostroma nitidum and Fortes 1988, Kornmann 1989, Luan 1989, Price & Scott Ulva spp.) forming an extensive band on the beach was 1992, Wynne 1993, 1995, Lewis & Mei-Lan 1996, Draisma observed. At Haikou (very polluted area with green algal et al.,1998, Abbott 1999, Huang 1999, Littler & Littler blooms), algae were collected from intertidal rocks and 2000, 2003, Leliaert & Coppejians 2003, Skelton 2003, sandy beaches where seaweeds were washed ashore. Abbott & Huisman 2004, Saunders & Hommersand 2004, Skelton & South 2004, 2007, South & Skelton 2003, Tsuda Conservation and identification of marine plants 2006, Brodie, Maggs & John 2007, Dawes & Mathieson 2008, Tadashi et al. 2008, and also the above mentioned Herbarized specimens (from 1990 and 1992 collections) and publications of the Chinese authors (see Introduction). freshly collected algal samples (2008−2012) were identified The systematics and nomenclature followed Guiry & using monographic publications, floristic studies and Guiry (AlgaeBase, searched in 2014). The previously known systematic articles by Børgesen 1913, 1914, 1915−1920, and newly recorded species for Hainan and China were 1924, 1940, 1948, Setchell & Gardner 1930, Yamada 1934, verified using Algaebase, the Catalogue of Life China (2010), Tanaka 1938, Shen & Fan 1950, Taylor 1950, 1960, 1966, Annual Checklist and The checklist of marine biota of China Egerod 1952, Dawson 1954, 1956, 1957, 1961, 1962, seas (Liu 2008). Hierarchical classification of the Phylum Durairatnam 1961, Zinova 1967, Womersley 1967, 1981, Rhodophyta is from Saunders and Hommersand (2004). 1984, 1987, 1994, Hollenberg 1968a, b, Trono 1968, 1969, The classification system of the Phyla Chlorophyta and 1997, Pham 1969, Womersley & Bailey 1970, Itono 1972, Ochrophyta basically follows Tsuda (2006).

Results Discussion

The results of the present study are shown in Table 1, In terms of species richness, Hainan Island was close to which documents 506 species of red, brown and green Taiwan where 476 species of algae were found (Zhang marine algae in Hainan Island. The Phylum Rhodophyta 1996, Lewis & Norris 1987). We consider that the reason of comprised 4 classes, 19 orders, 37 families, 103 genera and such high similarity is the geographical closeness of these 289 species (57% of all species). Ochrophyta comprised islands, and the intensity of study of their marine floras in 2 classes, 8 orders, 10 families, 24 genera and 92 species comparison with other subtropical and tropical areas of (18%) and Chlorophyta comprised 5 classes, 8 orders, 20 China. families, 40 genera and 125 species (25%). The floristic composition of macrophytes in Hainan A total of 39 new taxa were found in 2008−2012 on Island (Rhodophyta 57%, Chlorophyta 25% and Ochrophyta Hainan Island (17 reds, 10 browns and 11 greens). Of these, 18%) was also similar to that of Taiwan (55%, 24% and 21%, 16 are new for China (8 reds, 3 browns and 5 greens): respectively) and the southern coasts of China (56%, 23% Acrochaetium chaetomorphae, Pneophyllum confervicola, and 19%, respectively) (Zhang 1996, Lewis & Norris 1987). Chondria minutula, Ch. pygmaea, Polysiphonia exilis, P. The first marine floristic study In Hainan was conducted pseudovillum, Peyssonnelia boergesenii, Peyssonnelia in 1933 and the subsequent studies were carried our in the inamoena (Rh); Chilionema ocellatum, Sargassum miyabei, 1950s−1970s, 1990/1992 and after 2000 up to now. Floristic Turbinaria ornata var. cordata (Oc); Ulothrix subflaccida, similarity between the periods of the 1930s–1970s and Ulva rigida, Cladophoropsis membranacea, Siphonocladus 2008−2012 amounted to 33% (Rh), 41% (Oc) and 53% (Ch), rigidus, Caulerpa fastigiata. while that between 1990/1992 and 2008−2012 was 65% In the 1990−2012 survey, high algal species richness (Rh), 55% (Oc) and 53% (Ch) (Titlyanov et al. 2014c). was recorded at Luhuitou (222 taxa), Xiaodong Hai (148), These results suggest that conspicuous changes in Yalong Wan (122 taxa), Dadong Hai (89 taxa), Shalao (83 algal species diversity and composition occurred from taxa), Wenchang (76 taxa), Ximao Zhou (63 taxa), while the 1930s to 2012, as exemplified by a decline in species other localities had much less species. Thus, more algal richness and the appearance of new algal species. The species occurred in coral reef localities. largest numbers of taxa losses were recorded for families

© 2014 SCESAP - 32 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island

; Lh ) ) ; Xincun, itou, ); 9 ); 10; 12 ) Xh G. veprecula) ) , ); 9 ; Luhu Lj G. pacifica, , 9; 14 ; 9 1 3 4 2 3 G. rudis 5 1;14 1; 14 1; 14 1; 14 7; 8; G. fasciculata This paper Galaxaura obtusata References Porphyra vietnamensis ; Xiaodong Hai, subfruticulosa (as 4; 5; Galaxaura arborea 5 Galaxaura apiculata Xn 5; 7 (as ; Lingao Jiao, 7; G. glabriuscula Lc 7; 9 (as 4 (as Galaxaura ventricosa 4; 5; 6 (as 9 (as 5; 7; 9 (as ; Xianhai, 4, (as Yw ; Linchang, Hk Lh Xh,

2 , Lh Dh, , Yw, Lh Yw, Lh Lh Lh Yw Yw, Lh Yw, ; Haikou, ; Yalong Wan, Wan, Yalong ; Xh, Lh Xh, Lh Yw, Lh Yw, Wc, Mx, Yw, Dh, Yw, Mx, Dh Wc, Wc, Wc 2008 - 201 Mx, Xh, Dh Mx, Wc, Mx,

Ty Ty ; Wenchang, ; Wenchang, ; Dadong Hai, Ty Ty , Lh, Ty , Lh, Tg Xz, Bh Qk, Sl, Yw, Yw, Qk, Sl, Xh , Lh, Xz Qk Dh Xh, Ty Xh, Dh Mx, Xc, Xh, , 1990, 1992 Sl Yw, Dh, Xh, Xz, Yw, , Yw, , Sl, Yw, Lh, Xz, Yw, Sl, Dh, Xh, ; Tielu Gang, Tielu ; Lc Lc, Nm, Lj, Mx, Qk, Sl, Xc, Lc, Nm, Ty + + + + + + + + + + 1930s - 1970s ; Tian Ya Hai Tiao, Tiao, Hai Ya Tian ; Sl ; Shalao, Qk . Algal species recorded from Hainan Island, frpom the1930s to 2010s. . ; Qukou, Qg Table 1 Table

(Tokida) Kikuchi (Tokida) (Kjellman) A. Kurihara & Huisman (Kjellman) ; Qinglan Gang, (J. Ellis & Solander) Lamarck Rosenvinge (C. Agardh) Basson (C. (J. Ellis & Solander) Lamarck (Kjellman) Kurihara & Masuda R. Chou in W.R. Taylor Taylor R. Chou in W.R. (J. Ellis & Solander) Huisman Borowitzka* (T. Tanaka & P.H. Ho) J.E. Sutherland & & P.H. Tanaka (T. Mx (Dillwyn) J. Agardh (Dillwyn) J. (Linnaeus) Huisman & R.A. Townsend Townsend (Linnaeus) Huisman & R.A. (Kjellman) Kurihara & Masuda (Rosenvinge) Kornmann (Sommerfelt) Meneghini (Forsskål) Børgesen . For the 1930s-70s, + denotes occurrece (locality not know n). * − New records for Hainan island; ** new fo r China. Species, varieties and forms (J. Ellis & Solander) J.V. Lamouroux (J. Ellis & Solander) J.V. Kjellman (Zanardini) K.M. Drew) Xz apiculata , Meixia, RHODOPHYTA Nm Family Bangiaceae

; Ximao Zhou, Stylonema alsidii CLASS COMSPOPOGONOPHYCEAE ORDER ERTHROPELTIDALES Family Erythrotrichiaceae Erythrotrichia carnea Erythrocladia irregularis Porphyrostromium japonicum Sahlingia subintegra CLASS BANGIOPHYCEAE ORDER BANGIALES Porphyra crispata Pyropia vietnamensis Monotilla CLASS FLORIDEOPHYCEAE ORDER HILDENBRANDIALES Family Hildenbraniaceae Hildenbrandia rubra ORDER NEMALIALES Family Galaxauraceae Actinotrichia fragilis Dichotomaria Dichotomaria falcata Dichotomaria marginata Dichotomaria obtusata Dichotomaria spathulata Galaxaura divaricata Galaxaura filamentosa Galaxaura rugosa cylindrica Tricleocarpa Division CLASS RHODELLOPHYCEAE ORDER STYLONEMATALES Family Stylonemataceae Chroodactylon ornatum Taxa recorded during three periods are shown. Locality abbreviations: Baishamen (Haikou fish pond), Taxa Nanmai, Xc

© 2014 SCESAP - 33 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ); 12 (as ) ); 13 ); 9 ); 9 ); 9 ; 9; 13 (as ); 9 ; 9 ); 14 ); 9; 13 5 ) ); ; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 5; 9 4; 9 4; 9 4; 9 9; 4 5 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 9 4; 5; 9 4; 9; 5 4; 9; 4 5; 9; 4 Liagora valida References Liagora formosana 4; 5; 9;13; 14 Liagora pinnata Liagora perennis Liagora orientalis Liagora samaensis Liagora farinosa Liagora papenfussii Liagora divaricata Tricleocarpa oblongata Tricleocarpa 4 (as 4 (as Galaxaura oblongata 4 (as 4 (as 13 (as Ganonema farinosa 4 (as 4 (as 4; 9 (as 4 (as 5; 7; 9 (as 2 Lh Lh Lh Wc, 2008 - 201 Sl Sl Xh Xh Xh Dh, Xh, Xz 1990, 1992 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s ) K.C. Fan & Y.C. Wang Y.C. ) K.C. Fan & (Yamada) I.A. Abbott & Doty I.A. (Yamada) Tseng & Li Tseng Yamada I.A. Abbott & Doty I.A. ( Tseng & Li Tseng Harvey Tseng & Li Tseng Tseng & Li Tseng & Li Tseng Tseng & Li Tseng (Grunow ex Holmes) Fan Tseng & Li Tseng Tseng & Li Tseng Li (I.A. Abbott) J.M. Huisman, I.A. Abbott, & Abbott) J.M. Huisman, I.A. (I.A. Tseng & Li Tseng (Yamada) S.-M. Lin, S.-Y. Yang & Huisman Yang S.-M. Lin, S.-Y. (Yamada) (I.A. Abbott) S.M. Lin, S.-Y. Yang & Huisman Yang Abbott) S.M. Lin, S.-Y. (I.A. Tseng & Li Tseng Tseng & Li Tseng (C.K. Tseng) Huisman Tseng) (C.K. (Lamouroux) Fan et Wang (Lamouroux) Fan et Wang C.K. Tseng & W. Li & W. Tseng C.K. (C.K.Tseng) S.M. Lin, S.-Y. Yang & Huisman Yang S.M. Lin, S.-Y. (C.K.Tseng) (Harvey) Huisman (Harvey) Huisman, G.W. Saunders & A.R. Saunders & (Harvey) Huisman, G.W. C. K. Tseng & Li Tseng C. K. Yamada Yamada (Linnaeus) Huisman & R.A. Townsend Townsend (Linnaeus) Huisman & R.A. J.V. Lamouroux J.V. Tseng & Li Tseng (J. Agardh) Pedroche & Ávila Ortíz (J. Tseng & Li Tseng Zeh Species, varieties and forms Tseng & Li Tseng (Yamada) S.M. Lin, S.-Y. Yang & Huisman Yang S.M. Lin, S.-Y. (Yamada) Yamada Yamada Tseng & Li Tseng (J. Agardh) Huisman & Schils (J. J.V. Lamouroux J.V. Fan & Li K.C. Fan, Y.C. Wang & K.Y. Pan & K.Y. Wang Y.C. K.C. Fan, Yamada Yamada Yamada Yamada C.K. Tseng & Li Tseng C.K. perennis Tseng & Li Tseng pinnatiramosa divaricata rocarpus izziella c o e Tricleocarpa fragilis Tricleocarpa Family Liagoraceae Dermonema pulvinatum Dermonema virens Dotyophycus hainanensis Ganonema farinosum Ganonema papenfussii A.R. Sherwood Ganonema Ganonema pinnatum Ganonema samaense Helminthocladia australis Helminthocladia hainanensis Helminthocladia pinnata Izziella formosana Izziella orientalis Liagora albicans Liagora boergesenii Liagora ceranoides Liagora clavata Liagora dongdaoensis Liagora fanii Liagora filiformis Liagora hainanensis Liagora paniculata Liagora qishuiwanensis Liagora rhizophora Liagora robusta Liagora rubra Liagora segawae Liagora sinensis Liagora subdichotoma Liagora Wcensis Liagora wilsoniana Ma N Sinocladia divergenscata Sinocladia dongjiaoensis Sinocladia flabelliformis Sinocladia hainanensis Sinocladia paniculata Sinocladia pinnata Sinocladia qionghaiensis Sinocladia ramosissima setchellii Titanophycus validus Titanophycus Sherwood hawaiiana Trichogloeopsis mucosissima Trichogloeopsis

© 2014 SCESAP - 34 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ); 4 ); 9 ) ) );9 ) ); 9 7 ); 9 ); 9 ); 9 ); ) ); 14 ); 9; 14 ) A. seriata , 5 5 3 3 3 3 3 3 5 5 4 S. cottonii ; 14 4; 9 4; 4; 1; 14 5 1; 14 1; 14 4; 14 1; 14 1; 14 1; 14 4; 9; 14 Liagora caenomyce 4; 5; 7; 9 4; 5; 7; 9 This paper Porolithon onkodes Amphiroa zonata References Gloiofloea chinensis Audouinella globosa 4; 5 (as Audouinella robusta Audouinella dictyotae (as Audouinella gracilis Audouinella netrocarpa (as Audouinella ryukyuensis Liagorophila endophytica Cheilosporum acutilobum 5 5 4; 5 (as 4 (as Audouinella hypneae (as Cheilosporum jungermannioides 4 (as 4 (as 4 (as 4 (as 4; 9; 13 (as 4 (as 4 (as (as 4 (as 5 2 Xh, Lh Lh

Lh , Xh, Lh Lh , , Lh , Lh Lh Lh Wc Xh, Lh Yw, Lh,Yw Xh Dh, Lh Yw, Lh Yw, Lh, Yw Lh, Wc Dh Mx, Yw Mx, Lh Yw, Mx, Wc, Wc, 2008 - 201 Wc, Wc, Yw, Xh, Lh Yw, Wc, Wc, Yw, Dh, Yw, Wc, Wc, Mx, Yw, Xh, Lh Yw, Wc, Mx, Xz Ty , Xh, , Lh , Lh Ty

, , Ty Lh Xh, Xz Xh, Yw, Dh, Yw, Xh, Yw Xh

Dh Xh , , , Sl Sl Sl , Ty Xz, Xh Xh Xh Xh Xh , Sl Sl, Lh Mx, Sl, Yw Mx, Sl, Sl Sl, Yw Sl, Yw Sl, Xh, Nm Yw Nm, Yw Lh, Xz Lh, Dh 1990, 1992 Qk, Xc, Dh, Mx, Sl, Nm, Lc, Sl, Mx, Qk, Qg, Sl, Yw, Dh, Xh, Yw, Qk, Qg, Sl, Nm, + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s (J.V. Lamouroux) Weber-van Lamouroux) Weber-van (J.V. Batters (Tanaka & Pham-Hoàng Hô) (Tanaka

(Nägeli ex Kützing) Nägeli Børgesen M.A. Howe Nägeli (Lyngbye)

Børgesen (Nakamura) Papenfuss Weber-van Bosse Weber-van (Harvey) Batters (Decaisne) I.A. Abbott (Decaisne) I.A. Børgesen (Yendo) Yendo (Yendo) Børgesen (Garbary) Y. Lee & I.K. Y. (Garbary) cyathifera (Collins) I.-K. Hwang & H.-S. Kim (Børgesen) A.A. Santos & C.W.N. Moura A.A. Santos & C.W.N. (Børgesen) (J.V. Lamouroux) D. Penrose & Y.M. Y.M. Lamouroux) D. Penrose & (J.V. (Rosenvinge) G. Hamel (Linnaeus) J.V. Lamouroux (Linnaeus) J.V. f. C.K. Tseng C.K. Tseng (Weber-van Bosse & Foslie) Foslie (Weber-van (Dillwyn) Stegenga (Heydrich) D. Penrose & Woelkerling (Heydrich) D. Penrose & Woelkerling J.V. Lamouroux J.V. (Børgesen) Ateweberhan & Prud’homme van (Børgesen) C.K. Tseng C.K. Tseng (Foslie) Y.M. Chamberlain Y.M. (Foslie) Species, varieties and forms J.V. Lamouroux J.V. (C.K. Tseng) Huisman Tseng) (C.K. brevior J.V. Lamouroux J.V. Harvey M.A. Howe Lamouroux J.V. f. (Decaisne) J.H. Kim, Guiry & H.-G. Choi J.V. Lamouroux J.V. J.V. Lamouroux J.V. Family Colaconemataceae Yamadaella caenomyce Yamadaella Family Scinaiaceae Scinaia boergesenii Scinaia chinensis Scinaia latifrons Scinaia tsinglanensis ACROCHAETIALES ORDER Acrochaetiaceae Family Acrochaetium catenulatum Acrochaetium chaetomorphae Heerebout** Acrochaetium globosum Acrochaetium macula Acrochaetium microscopicum Acrochaetium netrocarpum Acrochaetium robustum Acrochaetium ryukyuense Acrochaetium secundatum Acrochaetium subseriatum Acrochaetium virgatulum Acrochaetium yamadae ORDER COLACONEMATALES Colaconema bonnemaisoniae Colaconema daviesii Colaconema dictyotae Colaconema gracile Reine Colaconema hypneae ORDER CORALLINALES Family Corallinaceae Amphiroa anastomosans Amphiroa beauvoisii Amphiroa foliacea Amphiroa fragilissima Amphiroa fragilissima Bosse Hydrolithon boreale Hydrolithon farinosum Chamberlain Hydrolithon onkodes Hydrolithon reinboldii Jania acutiloba Jania adhaerens Jania capillacea Jania crassa Jania pumila Jania ungulata Jania verrucosa

© 2014 SCESAP - 35 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ) ) ) 5 5 5 2 4 5; 16 3 3 3 1 3 2 1 3 1 3 2 4; 4; 4; ; 17; 14 1 M. simulans 1; 14 1; 14 3; 14 1; 14 1; 14 1; 14 1; 14 1; 14 1; 14 5 4; This paper This paper This paper This paper This paper 4; 9; 10; 12 4; Lithoporella pacifica References (as 5 Lithophyllum moluccense 4; (as 5 (as 4; 5 4; Xh,

2 Xh, Lh

Lh Xh, Lh

Dh, Lh Lh Lh Lh Lh Lh Lh Lh Lh Yw, Dh, Yw, Yw Yw Wc Yw, Lh Yw,

Yw, Xh, Lh Yw, Xh, Lh Xh, Lh Xh, Lh Dh, Yw, Lh Yw, Lh Yw, Lh, Yw Lh, Wc, Lh Wc, Lh Mx, Yw, Lh Yw, Mx, Yw, Xh, Lh Yw, Xh, Lh Yw, Wc, Wc, Yw, Dh, Xh, Lh Yw, 2008 - 201 Xn, Wc, Wc, Wc, Wc, Mx, Wc, Mx,

, , Xh Xh

, Dh Dh, Xz , , Xz

, Ty , Ty Lh, Yw Yw , Sl Sl Sl Xz Xz , Xh Xh Xh Lh, Xz , , Sl Sl Nm, Xh

Xh, , Sl, Xh, Lh Lh Xh, 1990, 1992 Sl, Nm, Xn, Sl, Xh Nm Nm,

, Lc, Mx, Sl, Yw, Xh Yw, Lc, Mx, Sl, Lj, Sl, Qk, Yw, Dh, Xh Yw, Lj, Sl, Qk, Lc, Mx + + + + + + + + + + 1930s - 1970s

Asparagopsis taxiformis (Schiffner) E.M. Wollaston (Schiffner)

(Weber-van Bosse & Foslie) Setchell (Weber-van Foslie (Foslie) Adey Kjellman* (Foslie) Foslie Unger Stage of (Kützing) Y.M. Chamberlain** Y.M. (Kützing) (Berthold) J.H. (Heydrich) Heydrich E.Y. Dawson E.Y. B.-M. Xia & Y.-Q. Zhang Y.-Q. B.-M. Xia & Børgesen (Delile) Trevisan de Saint-Léon Trevisan (Delile) Foslie* (C. Agardh) Montagne (C. (Rothpletz) Kylin Setchell & N.L. Gardner Itono J. Agardh (G. Martens) Okamura Weber-van Bosse Weber-van Weber-van Bosse Weber-van Yamada H.E. Petersen Weber-van Bosse Weber-van Nakamura Børgesen (Heydrich) Foslie Kützing E.Y. Dawson E.Y. Species, varieties and forms P.C. Silva P.C. (C. Agardh) Setchell (C. (Setchell & N.L.Gardner) T.O.Cho & S.M.Boo* T.O.Cho (Setchell & N.L.Gardner) (Harvey ex Kützing) T.O. Cho & L.J. McIvor T.O. (Harvey ex Kützing) (H.Richards) Mazoyer SPOROLITHALES Falkenbergia hillebrandii, Mastophora pacifica Mastophora rosea Lithophyllum kotschyanum Lithophyllum pygmaeum Lithophyllum tumidulum Lithoporella melobesioides Neogoniolithon megalocystum & L.R. Mason* Pneophyllum confervicola Pneophyllum fragile Family Hapalidiaceae Lithothamnion intermedium Lithothamnion phymatodeum Mesophyllum mesomorphum Order Family Sporolithaceae Sporolithon erythraeum AHNFELTIALES ORDER Ahnfeltiaceae Family Ahnfeltia yinggehaiensis ORDER BONNEMAISONIALES Family Bonnemaisoniaceae Asparagopsis taxiformis ( ORDER CERAMIALES Family Ceramiaceae Antithamnion antillanum Antithamnionella spirographidis Antithamnionella elegans Price & D.M. John Centroceras clavulatum Centroceras japonicum Centroceras minutum Ceramium aduncum Ceramium borneense Ceramium camouii Ceramium cimbricum Ceramium cingulatum Ceramium codii Ceramium comptum Ceramium macilentum Ceramium marshallense Ceramium procumbens Ceramium tenerrimum Ceramium vagans Corallophila kleiwegii Gayliella fimbriata Gayliella flaccida

© 2014 SCESAP - 36 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ) ) ) ); 19 19 tropica ; ); 19; 21 ) ); 14 P. papillosa P. var. var. ); 17 cartilaginea

5 ; 19 19 19 ); 14 (as A. orientalis 3 3 1 4 3 3 19 3 1 L. flexilis 5 19 19 19 3; 4 4; ; 19; 14 4; 5 19; 14 1; 14 1; 14 1; 14 4; 4; 4; 19 1; 14 4; 14 3; 19 4; 1; 14 1; 14 19; 23 5 4; 4; 1; 19; 14 18; 19; 14 This paper This paper This paper This paper This paper 4; A. venulosum References Chondrophycus undulata B. binderi, tenella Amansia glomerata

P. papillosa P. 4 (as Chondrophycus

4; 5; 19; 20 (as 4; 5 (as ; 19; 24; 25 (as 4; 19 (as 5 4 (as 4; 5; 12 (as 4; 1; 19 (as 2 Lh Lh Lh Xh, Lh Lh

Lh Yw, Yw, Xh, Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Dh

Yw, Xh, Lh Yw, Dh, Yw, Lh Yw, Lh Yw, Yw, Lh Yw, Yw, Wc,

Dh, Xh, Lh Dh, Yw, Xh, Lh Yw, Wc, 2008 - 201 Yw, Dh, Xh, Lh Yw, Yw, Mx, Yw, Xh, Lh Yw, Mx, Wc, , Xh , Ty Ty Xz Xz

Dh Lh Xz Lh,

Xz Xz , , , Ty , Yw Ty Yw, Lh, Xz Yw, Xh, Nm Xh , , Xz Xh, Lh Dh, Xh Dh, Xh Sl Sl, Dh, Xh, Xz Yw, Xh, Yw, , Mx Yw, Xh, Yw, Xh Yw, Nm, Dh, Xh 1990, 1992 Lj Nm, Sl, Xh, Lh Nm, Yw, Xh, Yw, Nm, , Sl, Yw, Xh, Lh, Yw, Sl, Sl, Dh, Yw, Xh, Lh, Xz, Yw, Nm, Mx, Yw, Dh, Xh, Xz Yw, Mx, Lc Nm, Qk, Sl, Yw, Xh, Xz Yw, Nm, Qk, Sl, + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s Tseng implicata Kim & I.K. Lee (J. Agardh) Hollenberg (Hariot) Yoon (Hariot) (C. Agardh) M.J. Wynne (C. M.S. villum (Yamada) Garbary & J.T. Harper Garbary & J.T. (Yamada) (K. Hering & G. Martens) Harvey (Børgesen) M.S. Kim & I.K. Lee (Yamada) Garbary & Harper (Yamada) (Suhr) Kylin savatieri Hollenberg** tenella (C. Agardh) R.E. Norris (C. (Linnaeus) Bory de Saint-Vincent Harvey var. Montagne (Harvey ex Kützing) M. S. Kim & I.K. Lee Yamada Yamada (C. Agardh) Ambronn f. Suhr ex J. Agardh f. Suhr ex J. (M. Vahl) Børgesen (M. Vahl) L. Ding, B. Huang, B. Xia & C.K. Tseng L. Ding, B. Huang, Xia & C.K. Falkenberg (C.K. Tseng) K.W. Nam K.W. Tseng) (C.K. (Suhr ex J. Agardh) S.M. Guimarães & M.T. Agardh) S.M. Guimarães & M.T. (Suhr ex J. (Harvey) Kützing (J. Agardh) J. Agardh var. var. (Harvey) Kylin Setchell (Harvey) A.H.S. Lucas (Harvey) Okamura Harvey Yamada Yamada (J. Bailey) M.-S. Kim, H.-G. Choi, Guiry & G.W. (J. Bailey) M.-S. Kim, H.-G. Choi, Guiry & G.W. Hollenberg (C. Agardh) J. Agardh Garbary & Vandermeulen** Species, varieties and forms (Bory de Saint-Vincent) K.W. Nam K.W. (Bory de Saint-Vincent) Weber-van Bosse** Weber-van Yamada* (J.V. Lamouroux) J. Agardh Lamouroux) J. (J.V. Yamada Yamada (Kützing) Okamura Børgesen Nam & Saito cartilagineus undulatus Zanardini

Family Callithamniaceae Crouania attenuata Family Delesseriaceae Acrosorium ciliolatum perpusillum Taenioma Family Rhodomelaceae Acanthophora muscoides Acanthophora spicifera Acrocystis nana Melanamansia glomerata Bostrychia tenella Chondria armata Chondria minutula Chondria pygmaea Chondria repens Chondrophycus Chondrophycus Herposiphonia parca Herposiphonia secunda Herposiphonia secunda Laurencia decumbens Laurencia majuscula Laurencia mariannensis Laurencia nanhaiense Laurencia okamurae Laurencia pinnata Laurencia similis Laurencia tropica Leveillea jungermannioides Lophosiphonia cristata Lophosiphonia reptabunda Neosiphonia ferulacea Fujii Neosiphonia harlandii Neosiphonia harveyi Saunders Neosiphonia sphaerocarpa Neosiphonia tongatensis Palisada parvipapillata Palisada perforata Polysiphonia blandii Polysiphonia crassa Polysiphonia exilis** Polysiphonia ferulacea Polysiphonia howei Polysiphonia japonica Polysiphonia pseudovillum Polysiphonia scopulorum Polysiphonia scopulorum Polysiphonia subtilissima

© 2014 SCESAP - 37 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island var. var. ); 14 ; 14 ), 26; 14 34 ) ; 35 33 ; 33; 35; 14 ;17

16 ; 33 26 32 3 3 3 3 3 3 3 19; 14 ; 32; 33 33 26; 27; 14 ; 16 ; ; 35; 37; 14 35; 36 ; ; 32, 1; 14 3; 14 3; 14 4; 1; 14 16 11 5 26; 14 31 4; 31 4; 16; 35 4; 16 Gelidium pusillum Griffithsia tenuis 29; 30; 14 31 This paper This paper 4; 5; 11; 26 4; 5; 11; 4; 4; 5; 26; 28 conchicola 4, References G. rhizophora 2; 4; 5; 4; 5; (as 4; 5; 11 4; 5; 33 4; 5; 29; 30; 4; 11 (as 4; 11 4; 5; 11 (as 4; 5; 11 4, 2 Lh Lh Xh, Lh

, Lh Lh Xh, Dh, Lh Lh

Lh , Lh Lh Lh Lh Lh Xh, Lh Yw Yw, Lh Yw, Lh Yw, Lh Yw, Yw, Wc, Wc, Wc, Wc, Lh Dh, Xh, Lh Yw, Xh, Lh Yw,

Mx, Yw Mx, Wc, Yw, Dh, Yw, Dh, Xh, Lh Yw, 2008 - 201 Yw, Dh, Yw, Mx, Wc, Wc, Wc, Wc, Ty Ty Sl , Xz Xh Xh Xh Xh Qk Qk Nm Mx Xh, Lh Dh, Xz Hk, Xh Nm, Xh Sl, Xh, Lh Lh, Xz, Xh, Xz, Lh Qk, Xh, Lh 1990, 1992 Nm, Mx, Yw, Lh Yw, Nm, Mx, Xh, Lh, Xz, , Lc, Nm, Dh, Xh Lc, Nm, Sl, Dh, Xh Qk, Dh, Xh, Lh, Xz Lc Lj, Mx, Xn, Sl, Dh, Xh, Lc, Nm, Qk, Sl, Yw, Dh, Yw, Lc, Nm, Qk, Sl, Mx, Qk, Qg, Yw, Xh, Lh, Yw, Mx, Qk, Qg, + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s W.R. Taylor Taylor W.R. C.K. Tseng W.R. Taylor Taylor W.R. J. Agardh B.M. Xia & C.K. Tseng Tseng B.M. Xia & C.K. * (Kützing) Santelices & Hommersand (Yamada & T. Tanaka) Itono Tanaka) T. & (Yamada (C. Agardh) F. Schmitz Agardh) F. (C. esperi (Weber-van Bosse) P.C. Silva Bosse) P.C. (Weber-van (S.G. Gmelin) Kützing Okamura C.K. Tseng C.K. Tseng (Feldmann) G. Furnar cylindricum pacificum G. Martens (E.Y. Dawson) B. Santelices (E.Y. subsecunda var. var. (Wulfen) Harvey (Wulfen) T. Tanaka Tanaka T. J. Agardh Species, varieties and forms C.K. Tseng C.K. Tseng f. J.V. Lamouroux J.V. (C. Agardh) Nägeli (C. (Forsskål) Feldmann & G. Hamel Bosse) Feldmann & C. Hamel (Weber-van (Stackhouse) Le Jolis var. var. J. Agardh Tanaka Tanaka Setchell* (Kützing) Feldmann & G. Hame (Montagne) Montagne (Hare ex Turner) Gaillon Turner) (Hare ex (C. Agardh) Kützing (C. (Kützing) J. Agardh (Kützing) J. Bory de Saint-Vincen Tolypiocladia condensata Tolypiocladia glomerulata Tolypiocladia Family Spyridiaceae Spyridia filamentosa Family Wrangeliaceae Anotrichium tenue Gordoniella yonakuniensis Griffithsia heteromorpha Griffithsia metcalfii Griffithsia metcalfii Griffithsia subcylindrica Wrangelia argus Wrangelia hainanensis ORDER GELIDIALES Family Gelidiaceae Gelidium crinale Gelidium divaricatum Gelidium pusillum Gelidium pusillum Gelidium pusillum Family Gelidiellaceae Gelidiella acerosa Gelidiella bornetii (Hainan) Gelidiella lubrica Parviphycus adnatus Parviphycus pannosus Family Pterocladiaceae Pterocladiella caerulescens Pterocladiella capillacea Santelices & Hommersand Pterocladiella yinggehaiensis ORDER GIGARTINALES Family Cystocloniaceae Hypnea boergesenii Hypnea cenomyce Hypnea charoides Hypnea cornuta Hypnea esperi Hypnea japonica Hypnea musciformis Hypnea nidulans Hypnea pannosa Hypnea spinella

© 2014 SCESAP - 38 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island 43; ); 32 16 ); ); ; 16 ); Polycavernosa 44 43 ; 47; 48 41 44 45; 46 ; G. minor ; ; 42 ) , 43 43 44 49 49 orientalis 49 42; 44 ; ; 16 41; ; 37 ; 38 ; 39 ; ; 41; 43 ; 41; 16 16 16 44 44 ; 49 3 3 ; 43; ); 5 (as 5; 41; ; ; 14 20 5 14 29; 14 var. var. ; 41; ; 4; 14 ; ; 16; 40 ; 41; 42; ; 41; 42, 43 ; 41; 16 16 16 ; 41; 43; 16; 1; 14 1; 14 4; 4; 16 4; 4; 4; 50 16 16 5 16 16 S. robusta 4; ; 16 16 16 G. crassa 32 4; 4; 4; 16 16 16 G. cacalia 16 5; 5 15; ); This paper This paper 4; ; ; 5; ; ; 4; 4; References 4; 5 5 4; 5 4; 5 4; (as 46; 12 5 P. rubra P. 4; 4; 4; Eucheuma gelatinae 4; 5 (as 5; 16; 43; 41; 4; ; 41 (as 4; (as (as ramulosa 5 16 5 G. bangmeiana ; 4; 5 4; 4 (as Lh 2 Lh Yw, Yw, Lh Lh Lh Xn Lh Lh Xn Xn Dh, Xh, Mx, Yw, Lh Yw, Yw, Yw, Mx, Wc, , 2008 - 201 Yw Mx, Xn , Sl Sl Sl Sl Ty Hk Xn Xn

Nm Mx, 1990, 1992 Sl, Yw, Dh, Xh, Lh, Yw, Sl, , Nm + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s Zhang & Xia (Suringar) Hommersand* liui (Harvey) Masuda Xia et Wang Weber-van Bosse** Weber-van B.-M. Xia & Y.-Q. Zhang Y.-Q. B.-M. Xia & Zhang & E.Z. Xia (Kützing) Millar (Gmelin) P.S. Silva (Gmelin) P.S. Piccone & Grunow Pilger (Weber-van Bosse) Cormaci & G. Furnari (Weber-van J. Agardh var. var. (Kützing) Sonder (J. Agardh) P.C. Silva & DeCew Agardh) P.C. (J. (Doty) Doty ex P.C. Silva (Doty) Doty ex P.C. C.F. Chang & B.M. Xia C.F. (Esper) Doty ex P.C. Silva (Esper) Doty ex P.C. (Sprengel) Feldmann & G. Hamel (Weber-van Bosse) Doty ex P.C. Silva Bosse) Doty ex P.C. (Weber-van (Lyngbye) P.C. Silva P.C. (Lyngbye) Zhang & B.M. Xia Zhang & Xia (Sonder) Kylin Harvey (Okamura) I.K. Lee & Kurogi (Okamura) Chang & B.M. Xia (C. Agardh) E.Y. Dawson Agardh) E.Y. (C. C.F. Chang & B.M. Xia C.F. (Greville) J. Agardh (Greville) J. Species, varieties and forms Zanardini (Kützing) Weber-van Bosse (Kützing) Weber-van (Turner) Montagne (Turner) (B.M. Xia & I.A. Abbott) I.A. Abbott, J. Zhang & Abbott) I.A. (B.M. Xia & I.A. Holmes (J. Agardh) J. Agardh I.A. Abbott, J. Zhang & B.M. Xia I.A. (Yamada) Yoshida Yoshida (Yamada) (C. Agardh) J. Agardh Hypnea valentiae Family Gigartinaceae Chondracanthus intermedius Family Peyssonneliaceae Peyssonnelia boergesenii Peyssonnelia conchicola Peyssonnelia inamoena Peyssonnelia orientalis Peyssonnelia rubra Family Phyllophoraceae Ahnfeltiopsis flabelliformis Ahnfeltiopsis hainanensis Ahnfeltiopsis pygmaea Family Rhizophyllidaceae Portieria hornemannii Family Solieriaceae Betaphycus gelatinum Eucheuma edule Eucheuma serra Kappaphycus alvarezii Kappaphycus cottonii Sarconema filiforme Sarconema gracilarioides Solieria pacifica Wurdemannia miniata ORDER GRACILARIALES Family Gracilariaceae Gracilaria arcuata Gracilaria articulata Gracilaria blodgettii Gracilaria bursa-pastoris Gracilaria canaliculata Gracilaria changii B.M. Xia Gracilaria chondracantha Gracilaria chorda Gracilaria coronopifolia Gracilaria cuneifolia Gracilaria glomerata Gracilaria hainanensis Gracilaria mixta Gracilaria rubra Gracilaria salicornia Gracilaria spinulosa Gracilaria tenuistipitata Gracilaria yamamotoi Gracilaria yinggehaiensis

© 2014 SCESAP - 39 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ) ; 16 ; ) 16 5 33 ) ); 16 16 ); ); ; 33 bailiniae ); 16 Gracilaria ); ) 33 );

); 14 ); 33 ; ) ; 47; 52, 53 16 29 (as ; ; 14 lariopsis i ); ; 54; 14 5 ; 14 16 ); ; 37 44 55 16 16 16 3 1 G. ligulata Grac 16 4; 33 4; 33 4; 33 1; 14 4; 33 4; 33 4; 33 4; 4; 1; 14 1; 14 4; 1; 14 4; This paper 4; 33; References 51 (as 51 4; 10; 12; Gracilaria confervoides Prionitis formosana Gelidiopsis intricata Sarcodia ceylonensis ), Gracilariophila infidelis Cryptonemia ovalifolia Cryptonemia luxurians 41; 42, 43; Polycavernosa fastigiata Gracilariophila setchellii 12; 20 (as Gracilariophila deformans 4; Gracialria eucheumatoides Gracilaria edulis (as 4 (as Gracilaria bailinae 5 (as 20 (as 4 (as 4 (as 4 (as 4; (as 4 (as 4 (as (as 4 (as 4 (as 16 heteroclada Lh Lh 2 Lh Xh, Lh

Xh, Xh, Xh, Lh Lh , , Dh, Xh, Lh Lh Lh Xn Wc Wc

Dh, Xn, Lh Dh, Dh Dh, Yw, Yw, Dh, Yw,

2008 - 201 Wc, Dh Wc, Wc, Mx, Xh,

Xz Yw, Sl, Xc, Tg, Sl, Xc, Xh, Xz ,

Xh , Sl Lh Lh Xz Hk Xn Qg Sl, Xh, Lh, Yw, Xh, Xz, Yw, 1990, 1992 Sl, Dh Yw, Lc, Nm, Mx Lc, Nm, Sl, Tg, Lc, Nm, Sl, + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s (Harvey) Gurgel & Fredericq (Weber-van Bosse) G.W. Saunders, Bosse) G.W. (Weber-van (Weber van Bosse) Gerung & Yamamoto van Bosse) Gerung & (Weber Zanardini Yendo Yendo (S.G. Gmelin) M. Seentoft, L.M. Irvine & B.-M. Xia & Y.-Q. Zhang Y.-Q. B.-M. Xia & (Montagne & Millardet) R.E. Norris (C. Agardh) R.E. Norris (C. Okamura B.M. Xia & Wang B.M. Xia & Wang W.-X. Li & Z.-F. Ding Li & Z.-F. W.-X. (Weber van Bosse) Gerung & Yamamoto Yamamoto van Bosse) Gerung & (Weber (C. Agardh) J. Agardh B.M. Xia & Wang B.M. Xia & Wang (J.D. Hooker & Harvey) J. Agardh (J.D. Hooker & Harvey) J. (Weber van Bosse) Gerung & Yamamoto Yamamoto van Bosse) Gerung & (Weber Zhang & B.M.Xia (Okamura) Kawaguchi & Masuda Børgesen (Montagne & Millardet) Chiang (Harvey) Børgesen J. Agardh S. Kawaguchi & H.W. Wang* S. Kawaguchi & H.W. (J. Agardh) P.G. Parkinson Agardh) P.G. (J. (Kylin) Balakrishnan Kawaguchi (S.G. Gmelin) Gurgel & Fredericq (J.V. Lamouroux) C. Agardh Lamouroux) C. (J.V. Species, varieties and forms Kützing (Harvey) Yamada Yamada (Harvey) (C. Agardh) Harvey (C. bailiniae lariopsis i Gracilariocolax deformans Gracilariocolax infidelis Gracilariocolax setchellii Grac Gracilariopsis longissima Farnham W.F. Hydropuntia edulis Hydropuntia eucheumatoides ORDER HALYMENIALES Family Halymeniaceae Carpopeltis maillardii Cryptonemia basinervis Cryptonemia seminervis Cryptonemia xinhaiensis Halymenia maculata Grateloupia asiatica Grateloupia filicina Grateloupia hainanensis Grateloupia kurogii Grateloupia livida Grateloupia ramosissima Isabbottia ovalifolia formosana Yonagunia ORDER SEBDENIALES Family Sebdeniaceae Sebdenia flabellata ORDER PLOCAMIALES Family Sarcodiaceae Sarcodia montagneana pygmaeus Trematocarpus ORDER RHODYMENIALES Family Champiaceae Champia parvula Champia vieillardii Coelothrix irregularis Family Hymenocladiaceae Asteromenia anastomosans Schneider & Kraft C.E. Lane, C.W. Family Lomentariaceae Ceratodictyon intricatum Ceratodictyon scoparium Ceratodictyon spongiosum Lomentaria corallicola Family Rhodymeniaceae Rhodymenia hainanensis

© 2014 SCESAP - 40 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ) ); 14 ); 14 ); 14 5 5 ; 14 ; 14 ; 14 2 22 22 5 5 5 56 4; 4; 5; 46 1; 14 1; 14 4; 14 1; 14 1; 14 1; 14 4; 14 4; 4; 4; This paper This paper This paper This paper References Giffordia conifera Hincksia mitchelliae Ectocarpus rhodochortonoides Ectocarpus elachistaeformis 22 (as 4 (as 22 (as 22 (as 2 Wc, Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Yw, Xh, Lh Yw, , Xh, Lh Xh, Lh Dh, Lh Yw, Xh, Lh Yw, Dh, Xh, Lh Wc Wc, 2008 - 201 Wc, Mx, Xn, Mx, , Xz, Dh , Xz Ty Lh, Yw , , Lh, Xz Xz Ty

Xh, Xc , Lh, , Ty , Xz Yw Xz Lh Xh Sl , , , Dh, Lc, Lj, Qk Lc, Lj, Dh 1990, 1992 Xh Dh, Xc, Yw, Xh, Lh Yw, Xc, Yw, Xh, Yw, Nm , Yw , , Lc, Nm, Lj, Sl, Dh, Xh, Lc Lc, Nm, Sl, Yw, Dh, Xh, Yw, Lc, Nm, Sl, Mx Lc, Nm, Mx, Qk, Qg, Sl, + + + + + + + + + + + + + + + 1930s - 1970s Tseng minor var. var.

(Børgesen) D.G. Müller & E.R. Sonder (Heydrich) M. Balakrishnan & Kinkar (J. Agardh) Ouriques & Bouzon (J. C. Agardh Trono (C. Agardh) M.A. Howe (C. Greville (Harvey) H.-S.Kim (J. Agardh) Børgesen (J. Meneghini (Kützing) G. Hamel (J. Agardh) Børgesen (J. (Kützing) Kornmann** Børgesen J. Agardh C.K. Tseng & Lu Tseng C.K. (Mertens ex Roth) Derbès & Solier in Castagne (K. Hering) Papenfuss (J. Agardh) P.-E. Lim & H. Kawai ex Kraft Agardh) P.-E. (J. Kützing Draisma, Keum,Prud’homme van Reine & Species, varieties and forms (Børgesen) I.A. Abbott (Børgesen) I.A. (Linnaeus)* SCYTOTHAMNALES SCYTOTHAMNALES ECTOCARPALES RALFSIALES SCYTOSIPHONALES SPHACELARIALES OCHROPHYTA Division CLASS XANTHOPHYCEAE ORDER VAUCHERIALES Family Vaucheriaceae Pseudodichotomosiphon constricta CLASS PHAEOPHYCEAE ORDER Family Scytothamnaceae Asteronema breviarticulatum Asteronema rhodochortonoides Parodi ORDER Acinetosporaceae Family Ectocarpus simpliciusculus Ectocarpus tamarinii Feldmannia irregularis Feldmannia mitchelliae Hincksia conifera Kuetzingiella elachistaeformis Family Chordariaceae Chilionema ocellatum Myrionema strangulans ORDER Family Neoralfsiaceae Neoralfsia expansa ORDER Family Scytosiphonaceae Chnoospora implexa Chnoospora minima Colpomenia sinuosa Hydroclathrus clathratus Hydroclathrus tenuis Rosenvingea intricata Rosenvingea orientalis Family Pylaiellaceae Pylaiella littoralis ORDER Family Sphacelariaceae Sphacelaria carolinensis Sphacelaria novae-hollandiae Sphacelaria rigidula Sphacelaria tribuloides Sphacelaria tsengii Lokhorst

© 2014 SCESAP - 41 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island S. S. ; 14 59 (as 5 ) ; 14 ); ), 61 (as 5 ); ) ); ; 61 61 61 62 , ; 61 ; 64 ; 65 ; 61 5 5 5 , 64; ; 14 ; 14 61 4 5 14 5 5 57 63 4; 4; 4; 4; 5 61 60 61 59; 61 61 60 58, 61 61 4; 14 4; 14 4; 5; 57 H. articulata S. crassifolium 4; 4; 4; 4; 64; 4; 4; 4; 4; 4; 59; 4; 4; 58; S. crassifolium 4; 4; , S. crassifolium This paper This paper This paper This paper This paper This paper This paper This paper This paper This paper , References 5 (as Dictyota linearis (as Dictyota cervicornis binderi 60

), 4 (as S. binderi 4 (as binderi 4 (as 2 Lh Wc , Lh Lh Dh, Xh, Lh Lh Lh Lh Lh Wc, Xn Xn Wc Xn Mx Wc Yw, Dh, Xh, Yw, , Wc, Mx, Mx, Xn, Hk, Yw Wc, 2008 - 201 Wc, Mx, Ty Ty Xh Ty Xz, Ty Sl Yw, Yw, Ty Lc Lc Xh Xh Dh Mx Nm Xz, Yw, Yw, Xh, Lh, 1990, 1992 Nm, Sl, Nm, Sl, Dh, Xz Mx, Dh, Xh, Lh, Lc, Sl, Dh, Xh, Lh, Xz Lc, Nm, Mx, Sl, Yw, Dh, Yw, Lc, Nm, Mx, Sl, Nm, Qk, Qg, Sl, Yw, Xh, Yw, Nm, Qk, Qg, Sl, + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s Grunow

(C. Agardh) Greville (C. parcespinosa C.K.Tseng & Lu* C.K.Tseng C Agardh (Kützing) De Paula & Clerck J. Agardh Montagne J. Agardh (J.F. Gmelin) P.C. Silva Gmelin) P.C. (J.F. (Turner) C. A. Agardh A. C. (Turner) var. var. (Turner) C. Agardh C. (Turner) C. Agardh Tseng & Lu Tseng (Grunow) Tseng & Lu Tseng (Grunow) intricata Tseng & Lu Tseng Yamada Yamada Hauck Tseng & Lu Tseng J.V. Lamouroux J.V. Greville (Turner) C.Agardh (Turner) Agardh (Mertens) C. Pham-Hoàng Hô Tseng & Lu Tseng Setchell (Harvey) Setchell* (J.V. Lamouroux) Womersley ex Oliveira Lamouroux) Womersley (J.V. Holmes* (Yendo) I.K.Hwang, H.-S.Kim & W.J.Lee (Yendo) J. Agardh (Okamura) Børgesen (Hudson) J.V. Lamouroux (Hudson) J.V. var. (Turner) C. Agardh* C. (Turner) Species, varieties and forms Trono Trono (Desfontaines) J.V.Lamouroux Hauck Thivy Setchell Tsuda DICTYOTALES DICTYOTALES Yamada Yamada aquifolium ORDER Family Dictyotaceae Canistrocarpus cervicornis Dictyopteris pacifica Dictyopteris repens Dictyota bartayresiana Dictyota dichotoma Dictyota dichotoma Dictyota friabilis Dictyota implexa Lobophora variegata Padina arborescens Padina australis Padina boryana Padina jonesii Padina minor Padina tetrastromatica Spatoglossum dichotomum ORDER Family Sargassaceae Hormophysa cuneiformis Sargassum agaviforme Sargassum angustifolium Sargassum Sargassum baccularia Sargassum carpophyllum Sargassum cervicorne Sargassum cinctum Sargassum crispifolium Sargassum cystophyllum Sargassum dazhouense Sargassum dotyi Sargassum erumpens Sargassum euryphyllum Sargassum feldmannii Sargassum fusiforme Sargasssum glaucescens Sargassum granuliferum Sargassum hainanense Sargassum hemiphyllum Sargassum henslowianum Sargassum herklotsii Sargassum heterocystum Sargassum horneri

© 2014 SCESAP - 42 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island , S. S.

, and . duplicatum , S 67 S. ilicifolium ; 14 61 ); 64; 14 ; 14 61 67 68 S. swartsii S. duplicatum, 61 61 61 ; ; 63 ; 64 ; , 61 61 ; 61 ; 66; 14 61 ); 58; 61 ; ; 61; 63; 64 71 4; 5 60 60 61 61 58; 61 4; (as 61; 64 59 61 61; 62 4; 5; 4; 5; 60 60 60 (as 4; 4; 4; 58; 61 4; 4; 59; 61 4; 4; 61; 63 4; 57; 4; 72; 4; 61; 68; 70; 14 This paper This paper This paper This paper This paper 4; 5; 59; 4; 5; ); 4; 4; References 57; wightii ); 61 cristaefolium S. cristaefolium S. acutifolium 5 (as S. ilicifolium duplicatum 4 (as 4; Hk 2 Hk , Lh, , Lh Lh Lh Lh Lh Lh Mx Mx Wc Wc Xh, Lh Yw, Lh Yw, Wc Dh, Xh, 2008 - 201 Xn, Wc, Lh Wc, Xn, Hk, Wc, Yw, Dh, Yw, Xn, Hk, Wc, , Ty Xz Ty Ty Dh Xh, Lh Xc, Dh Xh, Lh, Lh, Xz, Qg, Dh, Xh 1990, 1992 Sl, , Dh, Xh, Lh, Lc, Nm, Lj, Mx, Qk, X Yw Lc, Qg, Sl, Yw, Dh, Xh, Yw, Lc, Qg, Sl, Lc, Nm, Mx, Sl, Qg, Xc, + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s Grunow ** Tseng & Lu Tseng Tseng & Lu Tseng W.R. Taylor Taylor W.R. cordata Tseng & Lu Tseng Tseng & Lu Tseng Tseng & Lu Tseng conduplicatum Tseng & Lu Tseng J. Agardh* J. Tseng & Lu Tseng Montagne (C. Agardh) Tseng & Lu Tseng (Turner) C. Agardh C. (Turner) J. Agardh J. Agardh Tseng & Lu Tseng (Mertens ex Roth) Kuntze* Wrigh var. var. (Turner) C. Agardh C. (Turner) Tseng & Lu Tseng Wittrock Setchell C. Agardh Yendo** Yendo** (J. Agardh) Kützing (J. C. Agardh hainanensis (Lagerheim) Bornet & Flahault Womersley* Species, varieties and forms Tseng & Lu Tseng (Turner) J. Agardh J. (Turner) var. (Turner) f. Gomontiaceae CHLOROCOCCALES CHAETOPHORALES ULOTRICHALES CHLOROPHYTA Sargassum ilicifolium Sargassum ilicifolium Sargassum intermedium Sargassum mcclurei Sargassum megalocystum Sargassum microcystum Sargassum miyabe Sargassum oligocystum Sargassum parvifolium Sargassum phyllocystum Sargassum polycystum Sargassum primitivum Sargassum pseudolanceolatum Sargassum qionghaiense Sargassum sanyaense Sargassum siliquosum Sargassum silvae Sargassum spinifex Sargassum swartzii Sargassum symphyorhizoideum Sargassum tenerrimum Sargassum thunbergii Sargassum Wcense Sargassum yinggehaiense conoides Turbinaria ornata Turbinaria ornata Turbinaria ornata Turbinaria Division CLASS ORDER Family Chlorochytriaceae Chlorochytrium cohnii ORDER Family Chaetophoraceae Uronema marinum CLASS ORDER Family Gomontia polyrhiza Monostroma nitidum

© 2014 SCESAP - 43 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ); 14 68 ; 5; 14 ); 20 (as ); ); 72; 14 68, 72; 14 68 ), 68 68 68 71 68; 14 68, 72 68; 14 68 68; 70 68; 14 72; 68; 70 68; 70 68; 70 68; 70 68; 70 68; 70 46; 46; 68; 70 4; 68;70; 14 68; 70; 14 68; 70; 14 68; 70; 14 72; 46, 46; This paper This paper This paper This paper This paper 20; 46;, References C. crassa Enteromorpha prolifera Enteromorpha linza Enteromorpha clathrata Enteromorpha flexuosa 46 (as 4 (as 72 (as Enteromorpha tubulosa (as 4 (as 4 68; Hk Hk , 2 Hk Lh Lh Lh, Lh, Wc Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Lh Wc Wc Wc, Wc, Yw, Yw, Xh, Lh Xh, Lh Xh, Lh Yw, Lh Yw, Lh Yw, Xn, Dh, Xh, Mx, Xn, Lh Mx,

2008 - 201 Wc, Yw, Mx, Xn, , , Xz Qg Dh, Ty , , Xh Xz Sl,

Lh Bh , , Ty Dh Ty Dh, Mx Lh, Xh Mx , Lj, Ty Ty Ty Xz Qk Yw, Yw, Bh, Dh, Xh Lj, Qk, Qg, Yw, Xh Yw, Mx, Yw Mx, Dh Qg, , Mx, Qk, Tg, Yw, Yw, Tg, Mx, Qk, Sl, Xh, Lc, , Xh, Xz, Lj, Qg, Xh, Sl, 1990, 1992 Sl, Dh, Xh, Lh Lc, Bh, Dh, Xh Lj, Nm, Xn, Xh, Xz Nm Nm, Yw Lc, Lc, Yw, Dh, Xh, Lh Yw, Lc, , Qg, , Nm, Yw, Dh, Lh, Yw, Nm, Lc, Sl Lc, Nm, Lj, Mx, Qk, Bh, Lc Mx, Xn, Nm, Mx, Sl, Yw, Dh, Xh, Yw, Nm, Mx, Sl, + + + + + + + + + + + + + + 1930s - 1970s . J . M , Silva . C . P , Maggs ,

Blomster Harvey , (Bory de Saint-Vincent) Kützing (Bory de Saint-Vincent) (Börgesen) Hayden, Blomster, Maggs, P.C. Maggs, P.C. (Börgesen) Hayden, Blomster, (Kützing) Børgesen* Setchell Kützing (Kützing) Kützing (P.L. Crouan & H.M. Crouan) Batters (P.L. Kützing (N.L. Gardner) O’Kelly Harvey ex J.E. Gray Müller) Kützing (O.F. Hayden Wille** (Huber) R. Nielsen, C.J. O’Kelly & B. Wysor ) Species, varieties and forms Waaland (Kützing) Kützing Kjellman Linnaeus . Linnaeus (Reinke) R. Nielsen, C.J. O’Kelly & B. Wysor Ulvellaceae Ulvaceae Anadyomenaceae Cladophoraceae Forsskål* R (Dillwyn) Thuret* (Dillwyn) (Roth) C. Agardh (Roth) C. ULVALES CLADOPHORALES . Wulfen O.F. Müller O.F. (Reinke) R. Nielsen, C.J. O’Kelly & B. Wysor Linnaeus Kjellman J Bliding P.L. Crouan & H.M. P.L. (Harvey) Le Jolis C.Agardh* (

Linnaeus & kylinii

Family Ulotrichaceae Ulothrix flacca Ulothrix implexa Ulothrix subflaccida ORDER Family Phaeophilaceae Phaeophila dendroides Family Acrochaete geniculata Ulvella leptochaete Ulvella viridis Ulvella lens Ulvella scutata Family Ulva chaetomorphoides Silva, M.J. Stanhope & J.R. Waaland Ulva clathrata Ulva compressa Ulva conglobata Ulva flexuosa Ulva intestinalis Ulva Stanhope Ulva lactuca Ulva linza Ulva pertusa Ulva prolifera Ulva ralfsii Ulva reticulata Ulva rigida ORDER Family Anadyomene wrightii Microdictyon japonicum Family Chaetomorpha antennina Chaetomorpha brachygona Chaetomorpha capillaris Chaetomorpha gracilis Chaetomorpha javanica Chaetomorpha ligustica Chaetomorpha linum

© 2014 SCESAP - 44 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island 68 ); 14 D. 68; 14 , ) 46; ) , ) ), 68 68 ), 46; 68; 14 ); 46 ), ); ) 68 68 69 68; 14 68 C. torta 68 D. bokotensis 46; ; 68; 14 14 C. fasciculata 68; 46; 68 46; 68 4 68, 70 68; 70 4; 68; 70 68; 70 68; 70 68; 70 68; 70 68; 70 46; 68; 70 5; 46; 73; 68 68; 70 68; 70 Cladophora horii C. rugulosa 4; 4; 4; versluysii 68; 70; 14 46; 68; 70; 14 68; 70; 14 Valonia forbesii Valonia This paper 5; 4; 46; 73; References Cladophora fasciculata 46 (as R. kochianum 4; 72 (as 4; 5 (as Cladophora fuliginosa 5; 4; 46 (as 68; 68; 70 (as 46 (as 46; 68; 73 (as 4; Struvea anastomosans 5 (as 4; 4; 5 (as 68; 70; 72 (as 2 Xh, Lh Yw, Lh Yw, , Yw , Lh Lh Lh Lh Lh Lh Lh Lh Wc , Xh, Lh Xh, Lh Dh, Lh Dh, Lh Wc, Lh Yw, Lh Yw, Lh Yw, Lh Yw, Wc, Lh Wc Wc, Wc 2008 - 201 , Yw, Dh, Xh, Lh Yw, , Yw, Dh, Xh, Lh Yw, Dh, Xh, Lh Yw, Mx, Mx, Wc Ty Ty Ty Xz, Ty Sl Ty Ty Tg, Xh , Dh, Xh, Lh, Sl Sl Sl Xz, Ty Lh Xh Xh Qg Yw

Nm Sl, Lh Xz, Sl, Bh, Xh Yw Xh, Lh, Ty Xh, Lh, Dh, Bh, Xh, Lh Sl, Dh, Xh, Bh, Dh, Xh Nm, Mx, 1990, 1992 Lc, Sl, Yw, Dh, Xh Yw, Sl, Yw, Xh, Xz, Yw, Qk, Dh, Xh, Xz Nm, Mx, Sl, Yw Nm, Mx, Sl, Lj, Lc, Sl, Xh, Xz, Nm, Dh, Xh, Lh, Nm, Sl, + + + + + + + + + + + + + + + + + + + 1930s - 1970s Børgesen (Dillwyn) Rosenvinge a (Hofman Bang ex C. Agardh) Børgesen (Hofman Bang ex C. Reinbold (Kjellman) Wille (Harvey) Kraft et & M.J. Wynne (Harvey) Kraft et & M.J. Wynne implexum Weber-van Bosse Weber-van (Forsskål) Børgesen (Montagne) Kützing Weber-van Bosse Weber-van (G. Martens) (C. Hoek & Chihara) Boedeker Leliaert (Linnaeus) Hoek (Dillwyn) Kützing Okamur var. var. F.S. Collins & Hervey F.S. (Montagne) Kützing M.A. Howe** Kützing (Roth) Harvey Skottsberg & Levring in Børgesen (Linnaeus) Kützing (O.F. Müller) Kützing (O.F. (Harvey) Feldmann (Roth) Kützing Reinbold C. Agardh Kützing (Harvey) F. Brand (Harvey) F. ium Species, varieties and forms J. Agardh (Nees) Kützing Sonder (Roth) C. Agardh (Roth) C. Chihara J.V. Lamouroux J.V. Harvey ex J.Agardh* Yamada Boodleaceae Bryopsidaceae SIPHONOCLADALES BRYOPSIDALES Chaetomorpha minima Chaetomorpha spiralis Cladophora albida Cladophora aokii Cladophora catenata Cladophora coelothrix Cladophora flexuosa Cladophora herpestica Cladophora laetevirens Cladophora patentiramea Cladophora perpusilla Cladophora prolifera Cladophora sibogae Cladophora socialis Cladophora vagabunda Pseudocladophora horii Rhizoclonium grande Rhizoclonium ripar Rhizoclonium riparium CLASS SIPHONOCLADOPHYCEAE ORDER Family Boodlea composita Cladophoropsis fasciculatus Cladophoropsis membranacea Cladophoropsis sundanensis Phyllodictyon anastomosans Struvea enomotoi Family Siphonocladaceae Boergesenia forbesii Dictyosphaeria cavernosa Dictyosphaeria intermedia Dictyosphaeria versluysii Siphonocladus rigidus Family Valoniaceae aegagropila Valonia fastigiata Valonia utricularis Valonia ventricosa Valonia pachynema Valoniopsis CLASS BRYOPSIDOPHYCEAE ORDER Family Bryopsis australis Bryopsis pennata

© 2014 SCESAP - 45 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island C. (as 68 f. ) ); 68 46; 68 5 (as ), 68 ); ); 72 68; 14 ); 68; 14 68 ; ) ); ), 68 . clavifera ) triloba ; typica 70 68 68 5; 46; var 74; f. C. coronatum ; ; 72 ; 72 ; 74 68 68 ); 5 ); 68 68 68 68 70 68 68 68 68 68; 14 68 68 var. var. 4 4 46; 46; 68; 71 46 5; ; ; 46 ; 68 ; 68 ; 68 ; 46; . clavifera 5; 4; 5; 4; 4; 68 ; 46; 46; 5 5 5 68; 5 46; 46; 5 4; 5; 5 4; 46; 72 (as 5; 46; 5 C. parvifolia 68; 70; 14 var This paper This paper 4; 4; 4; 4; 4; 4; 4; 5; 4; References macrophysa C. geppii H. opuntia 68; Codium elongatum Udotea javensis C. racemosa 46 (as Rhipidodesmis caespitosa 4 (as 5 (as Trichosolen hainanensis Trichosolen C. racemosa 5; 46, Udotea javensis 46 (as racemosa 4; 4; 46 (as 46 (as 4 (as 4 (as 4 (as Lh 2 , Lh Dh, Xh, Lh Lh Lh Lh Lh Yw Wc Wc , Dh, Lh Yw, Lh Yw, Wc Dh, Xh, Lh Dh, Xh, Lh Yw, Dh, Lh Yw, 2008 - 201 Wc, Yw Wc, Xh Sl Sl Xh Xn Qk Yw Yw Yw Yw Sl, Xh Qk, Sl Qk, Sl Xh, Xz Dh, Dh, Xh, Xz 1990, 1992 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 1930s - 1970s Weber-van Weber-van Kützing) W.R. Kützing) W.R. amicorum f. (Zonder ex (J. Agardh) Børgesen (J. C.K. Tseng C.K. Tseng (Bory) de Saint-Vincent) (Bory) de Saint-Vincent) (J. Agardh) Collins (Vahl) C. Agardh C. (Vahl) C.K. Tseng lycopodium typica A. Gepp & E.S. longipes longiseta J. Agardh macrophysa occidentalis (Vahl) C. Agardh (Vahl) var. hainanense (Gmelin) M.A. Howe f. f. Montagne var. var. C.K. Tseng & M.L. Dong Tseng C.K. (Woodward) M.A. Howe (Woodward) Harvey ex J. Agardh Harvey ex J. (Okamura) Prud’Homme van Reine, Lokhorst Bornet & Flahault Decaisne (J. Ellis) J.V. Lamouroux (J. Ellis) J.V. (Forsskål) J. Agardh var. Harvey Harvey ex J. Agardh Harvey ex J. J. Agardh var. var. O.C. Schmidt Montagne** Okamura var. var. (Forsskål) J. Agardh (Forsskål) J. Species, varieties and forms (Berkeley) A. Gepp & E.S. (Berkeley) A.Gepp and E.S. Gepp (Hudson) C. Agardh* (Hudson) C. Kützing (Linnaeus) J.V. Lamouroux (Linnaeus) J.V. (Ellis & Solander) M.A. Howe (M. Vahl) C. Agardh C. (M. Vahl) & M.L. Dong Tseng C.K. J.V. Lamouroux J.V. C.K. Tseng & W.J. Gilbert & W.J. Tseng C.K. P.L. Crouan & H.M. P.L. Caulerpaceae Udoteaceae Codiaceae Bryopsis plumosa Pseudobryopsis hainanensis Family Ostreobiaceae Ostreobium quekettii Family Caulerpa brachypus Caulerpa cupressoides Caulerpa cupressoides Bosse Caulerpa cupressoides Caulerpa fastigiata Caulerpa nummularia Caulerpa peltata Caulerpa racemosa Caulerpa racemosa Taylor Caulerpa racemosa Caulerpa serrulata Caulerpa sertularioides Caulerpa sertularioides Caulerpa sertularioides Svedelius Caulerpa taxifolia Caulerpa verticillata Caulerpella ambigua Family erecta Avrainvillea lacerata Avrainvillea Chlorodesmis caespitosa Chlorodesmis hildebrandtii Chlorodesmis sinensis Penicillus sibogae Rhipidosiphon javensis Udotea flabellum Udotea fragilifolia Family Codium arabicum Codium bartlettii Codium decorticatum Codium geppiorum Codium intricatum Codium papillatum Codium repens Halimeda incrassata Halimeda macroloba Halimeda opuntia

© 2014 SCESAP - 46 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island ) ); 14 68; 14 ), Acetabularia 75; 14 75; 14

68; 75 46 (as 75 14 75 75 68; 7; 14 ); 68; ); , 35 – Xia & Wang 1997, , 35 – Xia & Wang Acetabularia pusilla 68; 70 68; 46; a, 17 – Zheng et al. 2001, 18 5; 46; 4; 68; 75; 14 4; References Acetabularia exigua 4; 4; , 68 – Titlyanov et al. 2011a, 69 – et al. 2011a, Titlyanov , 68 – moebii Acetabularia clavata A. parvula 75 (as (as 46; 68; 75 (as 68; 46 (as 4 Zhang & Xia 1979 Xia & Wang 2000 Xia & Wang Tseng & Lu 1999 Tseng , 16 – 2 , Lh Lh , Yw, Xh, Lh Yw, , , Yw, Dh, Xh, Yw, , Xh, Lh Yw Lh Yw, Lh Yw, Lh Yw, Lh Yw, Wc Wc 2008 - 201 Xia & Zhang 1999 Mx, Mx, , 33 – Xia et al. 2004b, 34 41 – Xia 1985, 42 – Xia & Yamamoto 1985, 43 – Chang & Xia 1976, 44 Yamamoto 41 – Xia 1985, 42 & Xia 2004 Xia 1981, , Xh , 32 – Yw Lc Qk Xh, Xz Xh, Xz 1990, 1992 Nm, Yw, Dh, Xh, Xz Yw, Santelices 1988 1988, 40 – Zhang & Doty + + + + + + + + + + + b, 56 – Draisma et al. 1998, 57 – Tseng & Lu 1988, 58 – Tseng & Lu 1995a, 59 – Tseng & Lu 1995b, 60 – Tseng & Lu 1997, Tseng & Lu 1995b, 60 – Tseng & Lu 1995a, 59 – Tseng & Lu 1988, 58 – Tseng b, 56 – Draisma et al. 1998, 57 et al. 2002, 31 – 1930s - 1970s Xia Xia & Wang 1999 Xia & Wang b, 38 – Masuda, Zhang & Xia 1994a, 39 , 63 – Tseng & Lu 2002a, 64 – Tseng & Lu 2002b, 65 – Tseng & Lu 2002d, 66 – Chiang et al. 1992, 67 Tseng & Lu 2002b, 65 – Tseng & Lu 2002a, 64 – Tseng , 63 – . Wright) J. Agardh J. . Wright) E.P Xia & Wang 2000 Xia & Wang Tseng & Lu 1992 Tseng (Yamada & T. Tanaka) Kobara & Chihara Tanaka) T. & (Yamada M.A. Howe Lamouroux in Quoi & Gaimard J.V. Solms-Laubach (Yamada) S. Berger, U. Fettweiss, et al. S. Berger, (Yamada) (Solms-Laubach) S. Berger, U. Fettweiss, et al. (Solms-Laubach) S. Berger, (Solms-Laubach) S. Berger, U. Fettweiss, et al. (Solms-Laubach) S. Berger, (Zanardini) Solms-Laubach (M. Howe) S. Berger, U. Fettweiss, et al. (M. Howe) S. Berger, (Harvey ex , 62 – Dickie M.A. Howe Species, varieties and forms Munier-Chalmas ex Sonder Derbesiaceae Dasycladaceae Tseng & Lu 2000 Tseng Family Pedobesia ryukyuensis CLASS DASYCLADOPHYCEAE ORDER Family Bornetella capitata Bornetella nitida Bornetella oligospora Bornetella sphaerica Neomeris annulata Neomeris mucosa Neomeris vanbosseae Family Acetabularia calyculus Parvocaulis clavatus Parvocaulis exiguus Parvocaulis parvulus Parvocaulis pusillus – Titlyanov et al. 2011a, 19 – Tseng et al. 2011, 20 – Tseng 1935, 21 – Tseng 1943, 22 – Luan & 1995, 23 Ding et al. 2007, 24 Xia Zhang 1982, 25 1988b, 26 Zheng Tseng 1935, 21 – Tseng 20 – et al. 2011, Tseng 19 – et al. 2011a, Titlyanov – 1942b, 29 – Zhang & Xia 1988a, 30 Tseng 2001, 27 – Zheng 1980, 28 1987, 37 – 36 – Xia & Wang Abbott 2002, 51 – Zhang & Xia 1991, 52 Abbott et al. 1991, 50 – Xia & Wang 1936; 47 – Chang & Xia 1963, 48 Millar 1999, 49 Tseng & Xia 1999, 45 – Zhang 1980, 46 Tseng Abbott 1987, 54 – Chang & Xia 1962, 55 1985, 53 – Xia & 61 – 1985. Tseng & Gilbert 1942, 75 – Dong Tseng & Chang 1962, 74 – Tseng 1938, 73 – Tseng et al. 2012, 71 – Zheng 2009, 72 Titlyanova Zhu & Liu 1980, 70 – Localities: Baishamen (Haikou fish pond), Bh; Dadong Hai, Dh; Ha ikou, Hk; Linchang, Lc; Lingao Jiao, Lj; Luhuitou, Lh; Nanmai, Nm, Meixia, Mx; Qinglan Gang, Qg; Qukou, Qk; Shalao, Yw; Xianhai, Xn; Xiaodong Hai, Xh; Xincun, Xc; Ximao Zhou, Xz. Wan, Yalong Wc; Tg; Wenchang, Gang, Tielu Ty; Tiao, Hai Ya Tian Sl; & Chang 2001, 9 – Zeng 1941b, 8 – Wang Tseng (ed) 1983, 6 – Zheng & Li 2009, 7 Tseng et al. 2014с, 4 – Liu 2008, 5 Titlyanov et al. 2014d; 3 - Titlyanov et al. 2014a, 2 - Titlyanov References: 1 - 2013, 15 – Titlyanova & Titlyanov 1941a, 14– Tseng et al. 1980, 13 – Tseng 1942a, 12 – –Tseng 1937, 11 Tseng et al. 2005, 10 –

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Liagoraceae, Gracilariaceae (Rh), Caulerpaceae, Codiaceae consequences of natural catastrophes, probably coral mass (Ch) and Sargassaceae (Oc). mortality after bleaching episodes in the South China Sea in The majority of lost species (not found in subsequent/ the summer of 1998. recent studies) were low productive, annual or perennial representatives inhabiting hard substrata (epilithic algae) with voluminous form of thalli (fleshy, leathery, coarse articulated, foliose, etc.). At the same time, the following Acknowledgements families contributed to taxa increases: Ceramiaceae (20 taxa), Rhodomelaceae (18 taxa) (Rh), Sphacelariaceae E. Titlyanov and T. Titlyanova are grateful to Prof. Christian (3 taxa), Dictyotaceae (5 taxa) (Oc), Cladophoraceae (14 Wiencke and Dr. Inka Bartsch for invitation to work at the taxa) and Ulvaceae (7 taxa) (Ch). Representatives of these Alfred-Wegener-Institute for Polar and Marine Research families are mainly high productive ephemeral and epiphytic and are also grateful to collaborators of the Botanical algae with fine filamentous forms and high surface area- Department for technical help, use facilities and hospitality. volume ratios (Titlyanov et al. 2011а, 2014b, с, Titlyanov & Special thanks to Prof. Huang Hui and Dr. Li Xiubao for Titlyanova 2012). invitation to work at the Marine Biological Station (South The present work, in combination with previous studies China Sea Institute of Oceanology, Chinese Academy dealing with the marine flora of Hainan Island (pointed of Sciences). This study was partially supported by the out above), suggests that serious changes occurred in Russian Foundation for Basic Research (grant numbers 08- the benthic flora, due probably to: (1) over-exploitation of 04-00735-a, 12-04-01424-a), by three research grants of the the reef ecosystems, especially in the 1950s−1970s; (2) Alfred-Wegener Institute of Polar and Marine Research to eutrophication of shallow waters (bays) in the 1990−2010s, T. Titlyanova and E. Titlyanov and the grant ‘China–Russia most notably sewage disposal (coastal urbanization, special funds 2009’ from the Chinese Academy of Sciences. agriculture, mariculture, and tourism industry, etc.); (3)

References Børgesen F (1915–1920) The marine algae of the Danish West Indies. Part 3. Rhodophyceae. Bd. I. Dansk Botansk Abbott IA (1999) Marine red algae of the Hawaiian Islands. Arkiv 3, 1-412. Bishop Museum Press, Hawaii, Honolulu. Børgesen F (1924) Marine algae from Easter Island. In: C Abbott IA & Hollenberg JH (1976) Marine algae of California. Skottsberg (ed.) The natural history of Juan Fernandez Stanford University Press, California, Stanford. and Easter Island. Upsala 2; 247-309. Abbott IA & Huisman JM (2004) Marine green and Børgesen, F. 1940. Some marine algae from Mauritius. 1. of the Hawaiian Islands. Bishop Museum Bulletin in Chlorophyceae. Det Kgl. Danske Videnskabernes Botany, 4. Bishop Museum Press, Honolulu, Hawaii. Selskab Biologiske Meddelelser 15, 1-81. Abbott IA, Zhang JF & Xia BM (1991) Gracilaria mixta, sp. Børgesen F (1948) Some marine algal flora from Mauritius. An nov. and other western Pacific species of the additional list to part 1. Chlorophyceae. Det Kgl. Danske (Rhodophyta: Gracilariaceae). Pacific Science 45(1), 12- Videnskabernes Selskab Biologiske Meddelelser 20, 27. 1-64. Bischoff B, Karsten U, Daniel C, Kuck K, Xia B & Brodie J, Maggs CA & John DM (2007) Green seaweeds of Wiencke C (1994) Preliminary assessment of the Britain and Ireland. Dataplus Print & Design, Dummurry, B-dimethylsulfoniopropionate (DMSP) content of Northern Ireland. macroalgae from the tropical island of Hainan (People’s Chang CF & Xia BM (1962) A preliminary phytogeographical Republic of China). Australian Journal of Marine & study on Chinese species of Gracilaria. Oceanologia et Freshwater Research 45, 1329-1336. Limnologia Sinica 4(3-4), 189-198. Børgesen F (1913) The marine algae of the Danish West Indies. Chang CF & Xia BM (1963) Polycavernosa a new genus of the Part 1. Chlorophyceae. Bd. I. Dansk Botansk Arkiv 1, Gracilariaceae. Studia Marina Sinica 2(3), 119-127. 1-160. Chang CF (Zheng JF) & Xia BM (1976) Studies on Chinese Børgesen F (1914) The marine algae of the Danish West Indies. species of Gracilaria. Studia Marina Sinica 11, 91-163 Part 2. Phaeophyceae. Bd. 1. Dansk Botansk Arkiv 2, [in Chinese, Abstract and Diagnoses of new species in 1-68. English].

© 2014 SCESAP - 48 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island

Chiang Y-M, Yoshida T, Ajisaka T, Trono GJr, Tseng CK & Lu Hutchings PA & Wu BL (1987) Coral reefs of Hainan Island, B (1992) Distribution and variation in Sargassum South China Sea. Marine Pollution Bulletin 18, 25-26. polycystum C.A. Agardh (Fucales, Phaeophyta). In: IA Itono H (1972) The genus Ceramium (Ceramiaceae, Abbott (ed.) of economic seaweeds. With Rhodophyta) in southern Japan. Botanica Marina 15, reference to some Pacific and Western Atlantic species. 74-86. Volume III. La Jolla, California: California Sea Grant Jaasund E (1976) Intertidal seaweeds in Tanzania. A field guide. College, pp. 35-42. University Tromsö, Norway. Dawes CJ & Mathieson AC (2008) The seaweeds of Florida. Kornmann P (1989) Sahlingia nov. gen. based on Erythrocladia University Press of Florida, Gainesville, FL. subintegra (Erythropeltidales, Rhodophyta). British Dawson EY (1954) Marine plants of the vicinity of the Institut Phycological Journal 24, 223-228. Oceanographique de Nhatrang, Vietnam. Pacific Leliaert F & Coppejians E (2003) The marine species of Science 8, 373-469. Cladophora (Chlorophyta) from the South African East Dawson EY (1956) Some marine algae of the Southern Marshall coast. Nova Hedwigia 76, 45-82. Islands. Pacific Science 10, 25-66. Lewis JE & Norris JN (1987) A history and annotated account Dawson EY (1957) An annotated list of marine algae from of the benthic marine algae of Taiwan. Smithsonian Eniwetok Atoll, Marshall Islands. Pacific Science 11, 92- Contribution to Marine Science 29, 1-38. 132. Lewis JE & Mei-Lan C (1996) The species of Ceramium Dawson EY (1961) Plantas marinas de la zona de las mareas de (Ceramiaceae, Rhodophyta) around Taiwan. El Salvador. Pacific Naturalist 2, 389-461. Hydrobiologia 326/327, 149-157. Dawson EY (1962) The marine red algae of Pacific Mexico. Part Littler DS & Littler MM (2000) Caribbean reef plants. Offshore 7. Ceramiales: Ceramiaceae, Delesseriaceae. Allan Graphics, Inc., Washington, D.C. Hancock Pacific Expedition 26, 1-207. Littler DS & Littler MM (2003) South Pacific reef plants. A divers’ Ding L, Huang B, Xia BM & Tseng CK (2007) Laurencia guide to the plant life of South Pacific coral reefs. nanhaiense sp. nov., a new species of the genus Offshore Graphics, Inc., Washington, D.C. Laurencia (Rhodomelaceae, Rhodophyta) from China. Liu L (2008) Checklist of marine biota of China Seas. Scientific Acta Oceanologica Sinica 26(5), 136-143. Book Service Co. Ltd., Beijing. Dong ML & CK Tseng (1985) Studies on the Dasycladales Lu B & Tseng CK (1980). Studies on some marine brown algae (Chlorophyta) of China. Chinese Journal of Oceanology from the Xisha Islands, Guangdong Province, China. I. and Limnology 3(1), 1-20. Studia Marina Sinica 17, 21-35 [in Chinese, Abstract in Doty MS (1988) Prodromus ad systematica Eucheumatoideorum: English]. a tribe of commercial seaweeds related to Eucheuma Luan RX (1989) Dalian algae. Field research guidance. Dalian [in (Solieriaceae, Gigartinales). In: IA Abbott (ed.) Taxonomy Chinese]. of economic seaweeds. With reference to some Luan RX & Luan SJ (1995) New materials for Chinese Pacific and Caribbean species. Vollume II. University Ectocarpaceae. Acta Phytotaxonomica Sinica 33, 506- of California, La Jolla: California Sea Grant College 515. Program. Report No. T-CSGCP-018, pp. 159-207. Millar AJK & Xia BM (1999) The genera Gracilaria and Draisma SGA, Keum Y-S, Prud’homme van Reine WF & Lokhorst Gracilariopsis from Norfolk Island, southwestern Pacific. GM (1998) The species of Sphacelaria (Sphacelariales, In: IA Abbott (ed.) Taxonomy of economic seaweeds. Phaeophyceae) in China with a description of a new With reference to some Pacific species. Vollume species. Botanica Marina 41, 181-190. IIV. University of California, La Jolla: The California Durairatnam M (1961) Contribution to the study of the marine Sea Grant College System. Report No. T-044 ISBN algae of Ceylon. Fisheries Research Station, Ceylon 1-888691-08-5, pp. 113-119. Bulletin 10, 1-181. Masuda M, Zhang JF & Xia BM (1994a) Ahnfeltiopsis from the Egerod EL (1952) An analysis of the siphonous Chlorophycophyta western Pacific: key, description and distribution of with special reference to the Siphonocladales, the species. In: IA Abbott (ed.) Taxonomy of economic Siphonales and Dasycladales of Hawaii. University of seaweeds. With reference to some Pacific species. California Publications in Botany 25(1), 325-453. Vollume IV. University of California, La Jolla: The Fiege D, Neumann V & Jinhe L (1994) Observations on coral California Sea Grant College. Report No. T-CSGCP-031, reefs of Hainan Island, South China Sea. Marine pp. 159-183. Pollution Bulletin 29, 84-89. Masuda M, Zhang JF & Xia BM (1994b) Synopsis of the Chinese Gurianova EF (1959) Marine zoological expedition to Hainan species of Gelidium (Gelidiales, Rhodophyta). In: Abbott Island. Vestnik AN SSSR 3, 89-92 [in Russian]. IA (ed.) & KJ Mcdermid (co-ed.) Taxonomy of economic Guiry MD & Guiry GM (2014) AlgaeBase. World-wide electronic seaweeds with reference to some Pacific species. publication, National University of Ireland, Galway. http:// Volume VIII. University of California, La Jolla: California www.algaebase.org; searched on 2014. Sea Grant College Program, pp. 183-205. Hollenberg GJ (1968a) An account of the species of Polysiphonia Perestenko LP (1980) Algae of the Peter Great Bay (the Sea of of the central and western tropical Pacific Ocean. I. Japan). Nauka, Leningrad [in Russian]. Oligosiphonia. Pacific Science 22, 56-98. Pham-Hoang H (1969) Marine algae of South Vietnam. Study Hollenberg GJ (1968b) An account of the species of the red alga Center, Saigon [in Vietnamese]. Polysiphonia of the central and western tropical Pacific Price IR & Scott FJ (1992) The turf algal flora of the Great Barrier Ocean. II. Polysiphonia. Pacific Science 22, 198-207. Reef. Part I. Rhodophyta. James Cook University, Huang S-F (1999) Marine algae of Kuei-Shan Dao, Taiwan. Townsville, Queensland, Australia. Taiwania 44, 49-71.

© 2014 SCESAP - 49 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island

Reyes AY (1976) The littoral benthic algae of Siquijor Province. Titlyanov EA, Titlyanova TV, Huang H & Li XB (2014a) Seasonal I. Cyanophyta and Chlorophyta. Philippine Journal of changes in benthic algal communities of the upper Science 105, 133-191. subtidal zone in Sanya Bay (Hainan Island, China). Santelices B (1988) Taxonomic studies on Chinese Gelidiales Journal of the Marine Biological Association of the (Rhodophyta). In: IA Abbott (ed.) Taxonomy of United Kingdom 94(1), 51-64. economic seaweeds. With reference on some Pacific Titlyanov EA, Titlyanova TV, Li XB & Huang H (2014b) Recent and Caribbean species. Volume II. California, La Jolla: (2008−2012) flora of Hainan Island, South California Sea Grant College Program, University of China Sea. Marine Biology Research ‘in press’. California, pp. 91−107. Titlyanov EA, Titlyanova TV, Kalita T, Xia BM & Bartsch I (2014c). Saunders GW & Hommersand MH (2004) Assessing red algal Inventory and decadal changes (1930−1990s) in the supraordinal diversity and taxonomy in the context of marine flora of Hainan Island (South China Sea). Marine contemporary systematic data. American Journal of Biology ‘in press’. Botany 91, 1494-1507. Titlyanov EA, Titlyanova TV, Li XB, Hansen GI & Huang H Setchell WA & Gardner NL (1930) Marine algae of the (2014d) Seasonal changes in the benthic macroalgae Revillagigedo Islands expedition in 1925. Proceedings and Cyanobacteria of the intertidal zone in Sanya Bay of the California Academy of Science, Series 4 19, 109- (Hainan Island, China). Journal of the Marine Biological 215. Association of the United Kingdom 94(5), 879-893. Shen YF & Fan KC (1950) Marine algae of Formosa. Taiwania 1, Titlyanova T, Titlyanov E, Xia B & Bartsch I (2012) New records 317-345. of benthic marine green algae on Hainan Island, China. Skelton PA (2003) Seaweeds of American Samoa. Townsville, Nova Hedwigia 94(3-4), 441-470. Queensland 4810, Australia. Trono GC (1968) The marine benthic algae of the Caroline Skelton PA & South GR (2004) New records and notes on Islands, I. Introduction, Chlorophyta and Cyanophyta. marine benthic algae of American Samoa: Chlorophyta, Micronesica 4, 137-206. Phaeophyta. Cryptogamie Algologie 25, 291-312. Trono GC (1969) The marine benthic algae of the Caroline Skelton PA & South GR (2007) The benthic marine algae of the Islands, II. Phaeophyta and Rhodophyta. Micronesica 5, Samoan Archipelago, South Pacific, with emphasis on 25-119. the Apia District. Nova Hedwigia Beihefte 132, 1-350. Trono GCJr (1997) Field guide and atlas of the seaweed South GR & Skelton PA (2003) Catalogue of the marine benthic resources of the Philippines. Department of Agriculture macroalgae of the Fiji Islands, South Pacific. Australian Bureau of Agricultural Research Fisheries Sector Systematic Botany 16, 699-758. Program Diliman, Quezon City. Tadashi K, Dai CF, Park H-S, Huang H. & Ang PO (2008) Status Trono GCJr & Put Jr (1982) Marine benthic algae from Bugsuk of coral reefs in East and North Asia (China, Hong Kong, Island and vicinity, Palawan, Philippines. Kalikasan, Taiwan, South Korea and Japan). In: C Wilkinson ( ed.) Philippine Journal of Biology 11, 1-26. Status of Coral Reefs of the World. Global Coral Reef Trono GCJr & Ganzon-Fortes ET (1988) Philippine seaweeds. Monitoring, Townsville, Australia, pp. 145-158. National Book Store, INC. Tanaka T (1938) The genus Hypnea from Japan. Scientific Tseng CK (1935) Economic seaweeds of Kwangtung Province, S. Papers of the Institute of Algological Research, Faculty China. Lingnan Science Journal 14, 93-102. of Science, Hokkaido Imperial University 2, 227-250. Tseng CK (1936) Studies on the marine Chlorophyceae from Taylor WR (1950) Plants of Bikini and other northern Marshall Hainan. I. Chinese Marine Biology Bulletin 1, 129-200. Islands. University of Michigan Press, Ann Arbor, MI. Tseng CK (1937) On marine algae new to China II. Bulletin of the Taylor WR (1960) Marine algae of the eastern tropical and Fan Memorial Institute of Biology 11, 83-118. subtropical coasts of the Americas. University of Tseng CK (1938) Studies on the marine Chlorophyceae from Michigan Press, Ann Arbor, MI. Hainan. II. Lingnan Science Journal 17, 141-149. Taylor WR (1966) Records of Asian and Western Pacific marine Tseng CK (1941a) Studies on the Chinese species of Liagora. algae, particularly from Indonesia and the Philippines. Bulletin of the Fan Memorial Institute of Biology 10, 265- Pacific Science 20, 342-359. 281. Titlyanov EA & Titlyanova TV (2012) Marine plants of Asian- Tseng CK (1941b) Studies on the Chaetangiaceae of China. Pacific Region, their use and cultivation. Vladivostok: Bulletin of the Fan Memorial Institute of Biology 11, 83- Dalnauka [in Russsian, Species descriptions in English]. 118. Titlyanov EA & Titlyanova TV (2013) Changes in the species Tseng CK (1942a) Studies on Chinese species of Griffithsia. composition of benthic macroalgal communities of the Papers of the Michigan Academy of Science, Arts and upper subtidal zone on a coral reef in Sanya Bay (Hainan Letters 27, 105-116. Island, China) during 2009–2012. Russian Journal of Tseng CK (1942b) Two new species of Wrangelia from China. Marine Biology 39(6), 413-419. Lingnan Science Journal 20, 261-270. Titlyanov EA, Titlyanova TV, Xia BM & Bartsch I (2011a) Tseng CK (ed.) (1983) Common seaweeds of China. Science Checklist of marine benthic green algae (Chlorophyta) Press Book, Beijing on Hainan, a subtropical island off the coast of China: Tseng CK (2004) The past, present and future of phycology in comparisons between the 1930s and 1990–2009 reveal China. Hydrobiologia 512, 11-20. environmental changes. Botanica Marina 54, 523-535. Tseng CK (2009) Seaweeds in Yellow Sea and Bohai Sea of Titlyanov EA, Kiyashko SI, Titlyanova TV, Yakovleva IM, Li XB China. Science Press, Beijing [in Chinese]. & Huang H (2011b) Nitrogen sources to macroalgal Tseng CK & Chang CF (1962) Studies on Chinese species of growth in Sanya Bay (Hainan Island, China). Current Dictyosphaeria. Acta Botanica Sinica 10, 120-132 [in Development in Oceanography 2, 65-84. Chinese, Abstract in English].

© 2014 SCESAP - 50 - Coastal Ecosystems, 2014, Vol 1, 28-53 Titlyanov E et al. Algal flora of Hainan Island

Tseng CK & Dong ML (1978) Studies on some marine algae from Tseng CK & Lu B (2002b) Studies on the biserrulic Sargassum the Xisha Islands, Guangdong Province, China. I. Studia of China: IV. The series Plagiophyllae Tseng et Lu. In: Marina Sinica 12, 41-50. IA Abbott (ed.) & K McDermid (co-ed.) Taxonomy of Tseng CK & Gilbert WJ (1942) On new algae of the genus economic seaweeds. With reference to some Pacific Codium from the South China Sea. Journal of the species. Vollume VIII. University of California, La Jolla: Washington Academy of Sciences 32, 291-296. California Sea Grant College, pp. 11-33. Tseng CK & Lu B (1983) Two new brown algae from the Tseng CK & Lu B (2002c) Studies on the biserrulic Sargassum Xisha Islands, South China Sea. Chinese Journal of of China: V. The series Ilicifoliae (J. Agardh) Tseng et Oceanology and Limnology 1, 185-188 [in Chinese]. Lu. In: IA Abbott (ed.) & K McDermid (co-ed.) Taxonomy Tseng CK & Lu B (1988) Studies on Chinese species of of economic seaweeds. With reference to some Pacific zygocarpic Sargassum. In: IA Abbott (ed.) Taxonomy of species. Vollume VIII. University of California, La Jolla: economic seaweeds. With reference to some Pacific and California Sea Grant College, pp. 35-76. Caribbean species. Vollume II. University of California, Tseng CK & Lu B (2002d) Some new species and records of La Jolla: California Sea Grant College Program. Report Sargassum (Sargassaceae, Phaeophyta) from the No. T-CSGCP-018, pp. 23-54. China Sea. In: IA Abbott (ed.) & K McDermid (co-ed.) Tseng CK & Lu B (1992) Studies on the malocarpic Sargassum Taxonomy of economic seaweeds. With reference of China: II. Racemosae J. Agardh. In: IA Abbott (ed.) to some Pacific species. Volume VIII. University of Taxonomy of economic seaweeds. With reference to California, La Jolla: California Sea Grant College, pp. some Pacific and Western Atlantic species. Vollume III. 135-148. University of California, La Jolla: California Sea Grant Tseng CK & Xia BM (1999) On the Gracilaria in the western College. Report No. T-CSGCP-023, pp. 11-34. Pacific and southeastern Asia region. Botanica Marina Tseng CK & Lu B (1995a) Studies on the glomerulate Sargassum 42, 209-218. of China: II. The series group Swartzia. In: IA Abbott Tseng CK, Chang TJ, Chang CF, Xia EZ, Xia BM, Dong ML & (ed.) Taxonomy of economic seaweeds. With reference Yang ZD 1962. Economic Seaweeds of China. Science to some Pacific species. Vollume V. University of Press, Beijing, China [in Chinese]. California, La Jolla: California Sea Grant College. Report Tseng CK, Chang CF, Xia E & Xia BM (1980) Studies on some No. T-CSGCP-035, pp. 67-74. marine red algae from Hong Kong. In: BS Norton & CK Tseng CK & Lu B (1995b) Studies on the glomerulate Sargassum Tseng (eds.) Proc. 1th Int. Mar. Biol. Workshop: The of China: I. The series Binderiana. In: IA Abbott (ed.) marine flora and fauna of Hong Kong and Southern Taxonomy of economic seaweeds. With reference to China, Hong Kong. University Press, Hong Kong, pp. some Pacific species. University of California, La Jolla: 58-84. California Sea Grant College. Report No. T-CSGCP-035, Tseng CK, Luan R, Chen Z & Li W (2005) Flora algarum pp. 75-92. marinarum sinicarum. Tomus II Rhodophyta. No. II Tseng CK & Lu B (1995c) Studies on the glomerulate Sargassum Acrochaetiales, Nemaliales, Bonnemaisoniales. Science of China: III. The species group Binderia. In: IA Abbott Press, Beijing. (ed.) Taxonomy of economic seaweeds. With reference Tseng CK, Wei TS, Yu YN, Wu PS & Bi L (2001) Flora to some Pacific species. Vollume V. University of algarum marinarum sinicarum. Tomus II Rhodophyta. California, La Jolla: California Sea Grant College. Report No. VI Ceramiales (I) Ceramiaceae, Dasyaceae, No. T-CSGCP-035, pp. 93-106. Delesseriaceae. Science Press, Beijing [in Chinese]. Tseng CK & Lu B (1997) Studies on the glomerulate Sargassum Tseng CK, Xia BM, Ding LP & Wang YQ (2011) Flora algarum of China: IV. Series of Platycarpae. In: IA Abbott (ed.) marinarum sinicarum. Tomus II. Rhodophyta. No. VII Taxonomy of economic seaweeds. With reference Ceramiales, Rhodomelaceae. Science Press, Beijing [in to some Pacific species. Vollume VI. University of Chinese, Key to the Genus of the Rhodomelaceae, to California, La Jolla: California Sea Grant College species and Diagnoses to new species in Latin]. System. Report No. T- 040, pp. 9-26. Tsuda RT (2006) Checklist and bibliography of the marine benthic Tseng CK & Lu B (1999) Studies on the biserrulic Sargassum algae within Chuuk, Pohnpei, and Kosrae States, of China: II. The series Coriifoliae J.G. Agardh. In: Federated States of Micronesia. Pacific Biological IA Abbott (ed.) Taxonomy of Economic Seaweeds Survey. Honolulu, USA: Bishop Museum, pp. 1-35. with reference to some Pacific species. Vollume VII. Vinogradova KL (1979) Seaweeds of the Far East Seas of University of California, La Jolla: California Sea Grant USSR. Chlorophyta. Leningrad, Nauka [in Russian]. College System. Report No. T-044, pp. 3-22. Wang W-L & Chang Y-M (2001) The reproductive development Tseng CK & Lu B (2000) Flora algarum marinarum sinicarum. of the red alga Actinotrichia fragilis (Galaxauraceae, Tomus III. Phaeophyta. No II. Fucales. Science Press, Nemaliales). European Journal of Phycology 36, 377- Beijing [in Chinese]. 384. Tseng CK & Lu B (2002a) Studies on the biserrulic Sargassum of Womersley HBS (1967) A critical survey of the marine algae of China: III. On Sargassum parvifolium (Turner) C. Agardh, southern Australia. II. Phaeophyta. Australian Journal of with a note on the classification of the Biserrulae. In: Botany 15, 189-270. IA Abbott (ed.) & K McDermid (co-ed.) Taxonomy of Womersley HBS (1981). Biogeography of Australasian marine economic seaweeds. With reference to some Pacific macroalgae. In: MN Clayton & RJ King (eds.) Marine species. Vollume VIII. University of California, La Jolla: Botany an Australasian perspective. Melbourne: California Sea Grant College, pp. 3-9. Longman Cheshire, pp. 292-307.

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Womersley HBS (1984) The marine benthic flora of Southern Xia BM & Wang Y-Q (2000a) Studies on some new red algae Australia. Part I. Government Printer, South Australia, from China (I). Acta Phytotaxonomica Sinica 38, 86-95. Adelaide. Xia BM & Wang Y-Q (2000b) Studies on some new red algae Womersley HBS (1987) The marine benthic flora of Southern from China (II). Acta Phytotaxonomica Sinica 38, 77-85. Australia. Part II. Government Printing Division, South Xia BM & Wang Y-Q (2002) A new species of Gracilaria from Australia, Adelaide. China. In: IA Abbott & KJ Mcdermid (eds.) Taxonomy Womersley HBS (1994) The marine benthic flora of of economic seaweeds with reference to some Pacific southern Australia. Part IIIA. Bangiophyceae and species. Volume VIII. La Jolla: California Sea Grant Florideophyceae (Acrochaetiales, Nemaliales, College, pp. 231-235. Gelidiales, Hildenbrandiales and Gigartinales sensu Xia BM & Yamamoto H (1985) Gracilaria species from both lato). Australian Biological Resources Study, Canberra. China and Japan: key, list and distribution of common Womersley HBS & Bailey A (1970) Marine algae of the Solomon and economically important species. In: IA Abbott & Islands. Philosophical Transactions of the Royal Society JN Norris (eds.) Taxonomy of Economic Seaweeds. B 259, 257-352. With reference to some Pacific and Caribbean species. Wynne MJ (1993) Benthic marine algae from the Maldives, Indian Vollume VII. University of California, La Jolla: California Ocean, collected during the R/V Te Vega Expedition. Sea Grant College Program, pp. 69-70. Contributions from the University of Michigan Herbarium Xia BM & Zhang JF (Chang CF) (1982) A new raw material 19, 5-30. for the manufacture of agar-agar from Hainan Island, Wynne MJ (1995) Benthic marine algae from the Seychelles China. Oceanologia et Limnologia Sinica 13(6), 538-543 collected during the R/V Te Vega Expedition. [in Chinese; Abstract in English]. Contributions from the University of Michigan Herbarium Xia BM & Zhang J (1999) Flora algarum marinarum sinicarum 20, 261-346. Tomus II. Rhodophyta No. V Ahnfeltiales, Gigartinales, Xia BM (1985) Gracilaria from China: key, list and distribution of Rhodymeniales. Science Press, Beijing [in Chinese; Key the species. In: IA Abbott & JN Norris (eds.) Taxonomy in English]. of Economic Seaweeds. With reference to some Xia BM, Tseng CK & Wang YQ (2002) Synopsis of the Chinese Pacific and Caribbean species. Vollume VII. University species of Gelidium (Gelidiales, Rhodophyta). In: IA of California, La Jolla: California Sea Grant College Abbott (ed.) & KJ Mcdermid (co-ed.) Taxonomy of Program, pp. 71−76. economic seaweeds with reference to some Pacific Xia B (ed.) (2004) Flora algarum marinarum sinicarum Tomus species. Vollume VIII. University of California, La Jolla: II. Rhodophyta No. III Gelidiales Cryptonemiales California Sea Grant College Program, pp. 183-205. Hildenbrandiales. Science Press, Beijing [in Chinese; Xia BM, Tseng CK & Wang YQ (2004) Studies on Chinese Diagnoses to new species in Latin]. species of Gelidiella and Pterocladiella (Gelidiales, Xia BM & Abbott IA (1985) The genus Polycavernosa Chang et Rhodophyta). Hydrobiologia 512, 201-207. Xia (Gracilariaceae, Rhodophyta): a comparison with Xia BM, Wang Y, Xia E, Li W & Ding Z (2004) Flora algarum Gracilaria Grev., and key to the species. In: IA Abbott marinarum sinicarum. Tomus II. Rhodohyta. No. III & JN Norris (eds.) Taxonomy of Economic Seaweeds. Gelidiales, Cryptonemiales, Hildenbrandiales. Science With reference to some Pacific and Caribbean species. Press, Beijing, China [in Chinese, Diagnoses to new Vollume VII. University of California, La Jolla: California species in Latin]. Sea Grant College Program, pp. 157-162. Yamada Y (1934) The marine Chlorophyceae from Ryukyu, Xia BM & Abbott IA (1987) New species of Polycavernosa Chang especially from the vicinity of Nawa. Journal of the & Xia (Gracilariaceae, Rhodophyta) from the western Faculty of Science, Hokkaido Imperial University Series Pacific. Phycologia 26(4), 405-418. 5, 3, 33-88. Xia BM & Wang YQ (1997) Some species of Hypnea Zeng CK (Tseng CK), Luan R, Chen Z (Zheng Y) & Li W (Gigartinales, Rhodophyta) from China. In: IA Abbott (2005) Flora algarum marinarum sinicarum. Tomus (ed.) Taxonomy of Economic Seaweeds. With reference II. Rhodophyta No. II Acrochaetiales, Nemaliales, to some Pacific species. Vollume VI. University of Bonnemaisoniales. Beijing: Science Press [in Chinese; California, La Jolla: California Sea Grant College, pp. Diagnoses to new species in Latin.] 1-42. Zhang JF (Chang CF) & Xia BM (1979) Studies of some marine Xia B & Wang YQ (1999a) Taxonomic studies on Pterocladiella red algae of the Xisha Islands, Guangdong Province, (Gelidiaceae, Gelidiales, Rhodophyta) from China. China. II. Studia Marina Sinica 15, 21-45 [in Chinese, In: IA Abbott (ed.) Taxonomy of Economic Seaweeds. with Abstract in English]. With reference to some Pacific species. Vollume VII. Zhang J-F & Xia BM (1980) Studies on some red algae of the California Sea Grant College System. University of Xisha Islands, Guangdong Province, China. III. Studia California, La Jolla, California, pp. 81-86. Marina Sinica 17, 49-68. Xia BM & Wang YQ (1999b) Taxonomic studies on Halymenia Zhang JF (Chang CF) & Xia BM (1983) Studies of some marine (Halymeniaceae, Halymeniales, Rhodophyta) from algae of the Xisha Islands, Guangdong Province, China. China. In: IA Abbott (ed.) Taxonomy of Economic IV. Studia Marina Sinica 5(20),123-139 [in Chinese; Seaweeds. With reference to some Pacific species. Diagnoses to new species in Latin]. Vollume VII. University of California, La Jolla: California Zhang JF (Chang CF) & Xia BM (1984) Some problems in the Sea Grant College System, pp. 173-176. taxonomy of Chinese species of Gracilaria (Rhodophyta). Hydrobiologia 116/117, 59-62.

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Zhang JF (Chang CF) & Xia BM (1988a) On two new Zhang G, Que H, Liu X. & Xu H (2004) Abalone mariculture in (Gigartinales, Rhodophyta) from South China. In: IA China. Journal of Shellfish Research 23, 947-950. Abbott (ed.) Taxonomy of Economic Seaweeds. With Zhang Q, Shi Q, Chen G, Fong TC, Wong DC, Huang H, Wang reference on some Pacific and Caribbean species. H & Zhao M (2006) Status monitoring and health Volume II. University of California, La Jolla: California assessment of Luhuitou fringing reef of Sanya, Hainan, Sea Grant College Program, pp. 131-136. China. Chinese Scientific Bulletin 51, 81-88. Zhang JF (Chang CF) & Xia BM (1988b) Laurencia from China: Zhang S (1996) The species distribution of the seaweeds in the Key, list and distribution of the species. In: IA Abbott coast of China seas. Chinese Biodiversity 4, 139-144. (ed.) Taxonomy of Economic Seaweeds. With reference Zheng B (1980) Studies on the Ceramiaceae from the Xisha on some Pacific and Caribbean species. Volume II. Islands, Guangdong Province, China. I. Studia Marina University of California, La Jolla: California Sea Grant Sinica 12, 37-48 [in Chinese, Abstract in English]. College Program, pp. 249-252. Zheng BF & Li J (2009) Flora algarum marinarum sinicarum. Zhang, J.F. and Xia B.M. 1991. A new species name in the Tomus II Rhodohyta. No. I Porphyridiales, genus Gracilaria. Chinese Journal of Oceanology and Erythropeltidales, Goniotrichales, Bangiales. Science Limnology 9(3), 290. Press, Beijing [in Chinese, Key to the Genera Zhang JF (Chang CF) & Xia BM (1994) Three foliose species Rhodomelaceae in English, Diagnoses to species of Gracilaria from China. In: IA Abbott (ed.) Taxonomy description in Latin]. of Economic Seaweeds with reference to some Pacific Zheng B, Liu J & Chen Z (2001) Flora algarum marinarum species. Volume IV. University of California, La Jolla: sinicarum. Tomus II Rhodophyta No. VI Ceramiales (I). California Sea Grant College, pp. 103-110. Ceramiaceae, Dasyaceae, Delesseriaceae. Science Zhang JF (Chang CF) & Xia EZ (1981) A new species of Press, Beijing [in Chinese, Key to families and species Sarconema from the Hainan Island, China. Oceanologia in English]. et Limnologia Sinica 12(3), 259-265 [in Chinese, Zhu H-R & Liu X (1980) The marine Cladophoraceae of the Xisha Abstract in English]. Islands, Guangdong Province, China. Studies Marina Zhang JF (Chang CF) & Xia E (1988с) Species of Gelidium Sinica 17, 11-18. Lamouroux and other Gelidiales (Rhodophyta) with key, Zinova AD (1967) Green, brown and red algae of the southern list, and distribution of the common species. In: IA Abbott seas of USSR. Nauka, Moscow-Leningrad [in Russian]. (ed.) Taxonomy of Economic Seaweeds. With reference on some Pacific and Caribbean species. Volume II. University of California, La Jolla: California Sea Grant College Program, pp. 109-113.

Published: 18 December 2014

• • • Editor's note • • • This contribution is categorised as Data Paper, given the importance of its extensive checklist of species.

© 2014 SCESAP - 53 - Coastal Ecosystems, 2014, Vol 1, 28-53