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http://dx.doi.org/10.4490/algae.2021.36.5.31

Benthic dinoflagellates in Korean waters  

Lim, An Suk (Division of Life Science & Plant Molecular Biology and Biotechnology Research Center, Gyeongsang National University)
Jeong, Hae Jin (School of Earth and Environmental Sciences, College of Natural Sciences, Seoul National University)
Publication Information
ALGAE / v.36, no.2, 2021 , pp. 91-109 More about this Journal
Abstract
The occurrence of benthic dinoflagellates, many of which are known to be toxic, is a critical concern for scientists, government officers, and people in the aquaculture, dining, and tourism industries. The interest in these dinoflagellates in countries with temperate climate is increasing because tropical or subtropical species introduced into temperate waters by currents are able to survive the winter season in the new environment owing to global warming. Recently, several species from the benthic dinoflagellate genera Amphidinium, Coolia, Ostreopsis, Gambierdiscus, and Prorocentrum have been reported in the waters of the South and East Sea of Korea. The advent of the benthic dinoflagellates in Korean waters is especially important because raw or slightly cooked seaweeds, which may harbor these benthic dinoflagellates, as well as raw fish, which can be potentially intoxicated by phytotoxins produced by some of these benthic dinoflagellates, are part of the daily Korean diet. The recent increase in temperature of Korean coastal waters has allowed for the expansion of benthic dinoflagellate species into these regions. In the present study, we reviewed the species, distribution, and toxicity of the benthic dinoflagellates that have been reported in Korean waters. We also provided an insight into the ecological and socio-economic importance of the occurrence of benthic dinoflagellates in Korean waters.
Keywords
Amphidinium; Coolia; epiphytic; Gambierdiscus; Ostreopsis; Prorocentrum; warming;
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1 Lee, K. H., Jeong, H. J., Park, K., Kang, N. S., Yoo, Y. D., Lee, M. J., Lee, J. -W., Lee, S., Kim, T., Kim, H. S. & Noh, J. H. 2013. Morphology and molecular characterization of the epiphytic dinoflagellate Amphidinium massartii, isolated from the temperate waters off Jeju Island, Korea. Algae 28:213-231.   DOI
2 Jeong, H. J., Yoo, Y. D., Kim, J. S., Seong, K. A., Kang, N. S. & Kim, T. H. 2010. Growth, feeding, and ecological roles of the mixotrophic and heterotrophic dinoflagellates in marine planktonic food webs. Ocean Sci. J. 45:65-91.   DOI
3 Jeong, H. J., Yoo, Y. D., Lee, K., Kang, H. C., Kim, J. S. & Kim, K. Y. 2021b. Annual carbon retention of a marine-plankton community in the eutrophic Masan Bay, based on daily measurements. Mar. Biol. 168:69.   DOI
4 Kang, N. S., Jeong, H. J., Lee, S. Y., Lim, A. S., Lee, M. J., Kim, H. S. & Yih, W. 2013. Morphology and molecular characterization of the epiphytic benthic dinoflagellate Ostreopsis cf. ovata in the temperate waters off Jeju Island, Korea. Harmful Algae 27:98-112.   DOI
5 Kang, Y. Q. 2000. Warming trend of coastal waters of Korea during recent 60 years (1936-1995). Fish. Aquat. Sci. 3:173-179.
6 Karafas, S., Teng, S. T., Leaw, C. P. & Alves-de-Souza, C. 2017. An evaluation of the genus Amphidinium (Dinophyceae) combining evidence from morphology, phylogenetics, and toxin production, with the introduction of six novel species. Harmful Algae 68:128-151.   DOI
7 Accoroni, S., Percopo, I., Cerino, F., Romagnoli, T., Pichierri, S., Perrone, C. & Totti, C. 2015. Allelopathic interactions between the HAB dinoflagellate Ostreopsis cf. ovata and macroalgae. Harmful Algae 49:147-155.   DOI
8 Ministry of Oceans and Fisheries. 2020. Annual statistical report for oceans and fishery in Korea. Ministry of Oceans and Fisheries, Sejong, 502 pp.
9 Mohammad-Noor, N., Moestrup, O., Lundholm, N., Fraga, S., Adam, A., Holmes, M. J. & Saleh, E. 2013. Autecology and phylogeny of Coolia tropicalis and Coolia malayensis (Dinophyceae), with emphasis on taxonomy of C. tropicalis based on light microscopy, scanning electron microscopy and LSU rDNA. J. Phycol. 49:536-545.   DOI
10 Morton, S. L., Bomber, J. W. & Tindall, P. M. 1994. Environmental effects on the production of okadaic acid from Prorocentrum hoffmannianum Faust I. temperature, light, and salinity. J. Exp. Mar. Biol. Ecol. 178:67-77.   DOI
11 Morton, S. L., Norris, D. R. & Bomer, J. W. 1992. Effect of temperature, salinity and light intensity on the growth and seasonality of toxic dinofalgellates associated with ciguatera. J. Exp. Mar. Biol. Ecol. 157:79-90.   DOI
12 Murray, S. A., Garby, T., Hoppenrath, M. & Neilan, B. A. 2012. Genetic diversity, morphological uniformity and polyketide production in dinoflagellates (Amphidinium, Dinoflagellata). PLoS ONE 7:e38253.   DOI
13 Aligizaki, K. & Nikolaidis, G. 2006. The presence of the potentially toxic genera Ostreopsis and Coolia (Dinophyceae) in the North Aegean Sea, Greece. Harmful Algae 5:717-730.   DOI
14 Bomber, J. W., Rubio, M. G. & Norris, D. R. 1989. Epiphytism of dinoflagellates associated with the disease ciguatera: substrate specificity and nutrition. Phycologia 28:360-368.   DOI
15 Ciminiello, P., Dell'Aversano, C., Iacovo, E. D., Fattorusso, E., Forino, M., Tartaglione, L., Benedettini, G., Onorari, M., Serena, F., Battocchi, C., Casabianca, S. & Penna, A. 2014. First finding of Ostreopsis cf. ovata toxins in marine aerosols. Environ. Sci. Technol. 48:3532-3540.   DOI
16 Accoroni, S. & Totti, C. 2016. The toxic benthic dinoflagellates of the genus Ostreopsis in temperate areas: a review. Adv. Oceanogr. Limnol. 7:1-15.
17 Aquino-Cruz, A., Purdie, D. A. & Morris, S. 2018. Effect of increasing sea water temperature on the growth and toxin production of the benthic dinoflagellate Prorocentrum lima. Hydrobiologia 813:103-122.   DOI
18 Baek, S. H. 2012a. First report for appearance and distribution patterns of the epiphytic dinoflagellates in the Korean peninsula. Korean J. Environ. Biol. 30:355-361.   DOI
19 Ok, J. H., Jeong, H. J., Lim, A. S., Lee, S. Y. & Kim, S. J. 2018. Feeding by the heterotrophic nanoflagellate Katablepharis remigera on algal prey and its nationwide distribution in Korea. Harmful Algae 74:30-45.   DOI
20 Jeong, H. J., Lim, A. S., Jang, S. H., Yih, W. H., Kang, N. S., Lee, S. Y., Yoo, Y. D. & Kim, H. S. 2012a. First report of the epiphytic dinoflagellate Gambierdiscus caribaeus in the temperate waters off Jeju Island, Korea: morphology and molecular characterization. J. Eukaryot. Microbiol. 59:637-650.   DOI
21 Tester, P. A., Litaker, R. W. & Berdalet, E. 2020. Climate change and harmful benthic microalgae. Harmful Algae 91:101655.   DOI
22 Vila, M., Graces, E. & Maso, M. 2001. Potentially toxic epiphytic dinoflagellate assemblages on macroalgae in the NW Mediterranean. Aquat. Microb. Ecol. 26:51-60.   DOI
23 Baek, S. H. 2012b. Occurrence of the toxic benthic dinoflagellate Gambierdiscus spp. in the uninhabited Baekdo Islands off southern coast and Seopsom Island in the Vicinity of Seogwipo, Jeju Province, Korea. Ocean Polar Res. 34:65-71.   DOI
24 Ben Gharbia, H., Kefi-Daly Yahia, O., Cecchi, P., Masseret, E., Amzil, Z., Herve, F., Rovillon, G., Nouri, H., M'Rabet, C., Couet, D., Triki, H. Z. & Laabir, M. 2017. New insights on the species-specific allelopathic interactions between macrophytes and marine HAB dinoflagellates. PLoS ONE 12:e0187963.   DOI
25 Tichadou, L., Glaizal, M., Armengaud, A., Grossel, H., Lemee, R., Kantin, R., Lasalle, J. -L., Drouet, G., Rambaud, L., Malfait, P. & De Haro, L. 2010. Health impact of unicellular algae of the Ostreopsis genus blooms in the Mediterranean Sea: experience of the French Mediterranean coast surveillance network from 2006 to 2009. Clin. Toxicol. 48:839-844.   DOI
26 Tindall, D. R. & Morton, S. L. 1998. Community dynamics and physiology of epiphytic/benthic dinoflagellates associated with ciguatera. In Anderson, D. M., Cembella, A. D. & Hallegraeff, G. M. (Eds.) Physiological Ecology of Harmful Algal Blooms. NATO ASI Series. Springer, Heidelberg, pp. 293-313.
27 Tubaro, A., Durando, P., Del Favero, G., Ansaldi, F., Icardi, G., Deeds, J. R. & Sosa, S. 2011. Case definitions for human poisonings postulated to palytoxins exposure. Toxicon 57:478-495.   DOI
28 Yamaguchi, H., Yoshimatsu, T., Tanimoto, Y., Sato, S., Nishimura, T., Uehara, K. & Adachi, M. 2012. Effects of temperature, salinity and their interaction on growth of the benthic dinoflagellate Ostreopsis cf. ovata (Dinophyceae) from Japanese coastal waters. Phycol. Res. 60:297-304.   DOI
29 Yang, A. R., Lee, S., Yoo, Y. D., Kim, H. S., Jeong, E. J. & Rho, J. -R. 2017. Limaol: a polyketide from the benthic marine dinoflagellate Prorocentrum lima. J. Nat. Prod. 80:1688-1692.   DOI
30 Yoo, Y. D., Jeong, H. J., Lee, S. Y., Yoon, E. Y., Kang, N. S., Lim, A. S., Lee, K. H., Jang, S. H., Park, J. Y. & Kim, H. S. 2015. Feeding by heterotrophic protists on the toxic dinoflagellate Ostreopsis cf. ovata. Harmful Algae 49:1-9.   DOI
31 Cruz-Rivera, E. & Villareal, T. A. 2006. Macroalgal palatability and the flux of ciguatera toxins through marine food webs. Harmful Algae 5:497-525.   DOI
32 Bomber, J. W., Guillard, R. R. L. & Nelson, W. G. 1988. Role of temperature, salinity, and light in seasonality, growth, and toxicity of ciguatera-causing Gambierdiscus toxicus Adachi et Fukuyo (Dinophyceae). J. Exp. Mar. Biol. Ecol. 115:53-65.   DOI
33 Carlson, R. D., Morey-Gaines, G., Tindall, D. R. & Dickey, R. W. 1984. Ecology of toxic dinoflagellates from the Caribbean Sea: effects of macroalgal extracts on growth in culture. In Ragelis, E. P. (Ed.) Seafood Toxins. The American Chemical Society, Washington, D.C., pp. 271-287.
34 Carnicer, O., Garcia-Altares, M., Andree, K. B., Tartaglione, L., Dell'Aversano, C., Ciminiello, P., de la Iglesia, P., Diogene, J. & Fernandez-Tejedor, M. 2016. Ostreopsis cf. ovata from western Mediterranean Sea: physiological responses under different temperature and salinity conditions. Harmful Algae 57:98-108.   DOI
35 Fraga, S., Penna, A., Bianconi, I., Paz, B. & Zapata, M. 2008. Coolia canariensis sp. nov. (Dinophyceae), a new nontoxic epiphytic benthic dinflagellate from the Canary Islands. J. Phycol. 44:1060-1070.   DOI
36 Besada, E. G., Loeblich, L. A. & Loeblich, A. R. 3rd. 1982. Observations on tropical, benthic dinoflagellates from ciguatera-endemic areas: Coolia, Gambierdiscus, and Ostreopsis. Bull. Mar. Sci. 32:723-735.
37 Lim, A. S., Jeong, H. J., Jang, T. Y., Kang, N. S., Lee, S. Y., Yoo, Y. D. & Kim, H. S. 2013. Morphology and molecular characterization of the epiphytic dinoflagellate Prorocentrum cf. rhathymum in temperate waters off Jeju Island, Korea. Ocean Sci. J. 48:1-17.   DOI
38 Nishimura, T., Uchida, H., Noguchi, R., Oikawa, H., Suzuki, T., Funaki, H., Ihara, C., Hagino, K., Arimitsu, S., Tanii, Y., Abe, S., Hashimoto, K., Mimura, K., Tanaka, K., Yanagida, I. & Adachi, M. 2020. Abundance of the benthic dinoflagellate Prorocentrum and the diversity, distribution, and diarrhetic shellfish toxin production of Prorocentrum lima complex and P. caipirignum in Japan. Harmful Algae 96:101687.   DOI
39 Jeong, H. J., Kang, H. C., Lim, A. S., Jang, S. H., Lee, K., Lee, S. Y., Ok, J. H., You, J. H., Kim, J. H., Lee, K. H., Park, S. A., Eom, S. H., Yoo, Y. D. & Kim, K. Y. 2021a. Feeding diverse prey as an excellent strategy of mixotrophic dinoflagellates for global dominance. Sci. Adv. 7:eabe4214.   DOI
40 Fukuyo, Y. 1981. Taxonomical study on benthic dinoflagellates collected in coral reefs. Bull. Jpn. Soc. Sci. Fish. 47:967-978.   DOI
41 Loeblich, A. R., Sherley, J. L. & Schmidt, R. J. 1979. The correct position of flagellar insertion in Prorocentrum and description of Prorocentrum rhathymum sp. nov. (Phyrrhophyta). J. Plankton Res. 1:113-120.   DOI
42 Mustapa, N. I., Yong, H. L., Lee, L. K., Lim, Z. F., Lim, H. C., Teng, S. T., Luo, Z., Gu, H., Leaw, C. P. & Lim, P. T. 2019. Growth and epiphytic behavior of three Gambierdiscus species (Dinophyceae) associated with various macroalgal substrates. Harmful Algae 89:101671.   DOI
43 Rhodes, L. 2011. World-wide occurrence of the toxic dinoflagellate genus Ostreopsis Schmidt. Toxicon 57:400-407.   DOI
44 Ibrahim, A. M. M. 2007. Review of the impact of harmful algae blooms and toxins on the world economy and human health. Egypt. J. Aquat. Res. 33:210-223.
45 Park, J., Hwang, J., Hyung, J. -H. & Yoon, E. Y. 2020. Temporal and spatial distribution of the toxic epiphytic dinoflagellate Ostreopsis cf. ovata in the coastal waters off Jeju Island, Korea. Sustainability 12:5864.   DOI
46 Pavaux, A. -S., Berdalet, E. & Lemee, R. 2020. Chemical ecology of the benthic dinoflagellate genus Ostreopsis: review of progress and future directions. Front. Mar. Sci. 7:498.   DOI
47 Glibert, P. M., Burkholder, J. M. & Kana, T. M. 2012. Recent insights about relationships between nutrient availability, forms, and stoichiometry, and the distribution, ecophysiology, and food web effects of pelagic and benthic Prorocentrum species. Harmful Algae 14:231-259.   DOI
48 Gomez, F., Qiu, D. & Lin, S. 2017. The synonymy of the toxic dinoflagellates Prorocentrum mexicanum and P. rhathymum and the description of P. steidingerae sp. nov. (Prorocentrales, Dinophyceae). J. Eukaryot. Microbiol. 64:668-677.   DOI
49 Parsons, M. L., Aligizaki, K., Bottein, M. -Y. D., Fraga, S., Morton, S. L., Penna, A. & Rhodes, L. 2012. Gambierdiscus and Ostreopsis: reassessment of the state of knowledge of their taxonomy, geography, ecophysiology, and toxicology. Harmful Algae 14:107-129.   DOI
50 Parsons, M. L. & Preskitt, L. B. 2007. A survey of epiphytic dinoflagellates from the coastal waters of the island of Hawai'i. Harmful Algae 6:658-669.   DOI
51 Pezzolesi, L., Guerrini, F., Ciminiello, P., Dell'Aversano, C., Iacovo, E. D., Fattorusso, E., Forino, M., Tartaglione, L. & Pistocchi, R. 2012. Influence of temperature and salinity on Ostreopsis cf. ovata growth and evaluation of toxin content through HR LC-MS and biological assays. Water Res. 46:82-92.   DOI
52 Rhodes, L., Adamson, J., Suzuki, T., Briggs, L. & Garthwaite, I. 2000. Toxic marine epiphytic dinoflagellates, Ostreopsis siamensis and Coolia monotis (Dinophyceae), in New Zealand. N. Z. J. Mar. Freshw. Res. 34:371-383.   DOI
53 Lie, H. -J. & Cho, C. -H. 2016. Seasonal circulation patterns of the Yellow and East China Seas derived from satellitetracked drifter trajectories and hydrographic observations. Prog. Oceanogr. 146:121-141.   DOI
54 Lee, S., Yang, A. R., Yoo, Y. D., Jeong, E. J. & Rho, J. -R. 2019. Relative configurational assignment of 4-hydroxyprorocentrolide and prorocentrolide C isolated from a benthic Dinoflagellate (Prorocentrum lima). J. Nat. Prod. 82:1034-1039.   DOI
55 Lessard, E. J. 1991. The trophic role of heterotrophic dinoflagellates in diverse marine environments. Mar. Microb. Food Web 5:49-58.
56 Li, X., Yan, M., Gu, J., Lam, V. T. T., Wai, T. -C., Baker, D. M. Thompson, P. D., Yiu, S. K. F., Lam, P. K. S. & Leung, P. T. Y. 2020. The effect of temperature on physiology, toxicity and toxin content of the benthic dinoflagellate Coolia malayensis from a seasonal tropical region. Water Res. 185:116264.   DOI
57 Lim, A. S., Jeong, H. J., Ok, J. H. & Kim, S. J. 2018. Feeding by the harmful phototrophic dinoflagellate Takayama tasmanica (Family Kareniaceae). Harmful Algae 74:19-29.   DOI
58 Litaker, R. W., Vandersea, M. W., Faust, M. A., Kibler, S. R., Chinain, M., Holmes, M. J., Holland, W. C. & Tester, P. A. 2009. Taxonomy of Gambierdiscus including four new species, Gambierdiscus caribaeus, Gambierdiscus carolinianus, Gambierdiscus carpenteri, and Gambierdiscus ruetzleri (Gonyaulacales, Dinophyceae). Phycologia 48:344-390.   DOI
59 Schmidt, J. 1901. Flora of Koh Chang. Contributions to the knowledge of the vegetation in the Gulf of Siam. Peridiniales. Bot. Tidsskr. 24:212-221.
60 Rhodes, L. & Thomas, A. E. 1997. Coolia monotis (Dinophyceae): a toxic epiphytic microalgal species found in New Zealand. N. Z. J. Mar. Freshw. Res. 31:139-141.   DOI
61 Selina, M. S., Morozova, T. V., Vyshkvartsev, D. I. & Orlova, T. Y. 2014. Seasonal dynamics and spatial distribution of epiphytic dinoflagellates in Peter the Great Bay (Sea of Japan) with special emphasis on Ostreopsis species. Harmful Algae 32:1-10.   DOI
62 Yong, H. L., Mustapa, N. I., Lee, L. K., Lim, Z. F., Tan, T. H., Usup, G., Gu, H., Litaker, R. W., Tester, P. A., Lim, P. T. & Leaw, C. P. 2018. Habitat complexity affects benthic harmful dinoflagellate assemblages in the fringing reef of Rawa Island, Malaysia. Harmful Algae 78:56-68.   DOI
63 Litaker, R. W., Vandersea, M. W., Faust, M. A., Kibler, S. R., Nau, A. W., Holland, W. C. Chinain, M., Holmes, M. J. & Tester, P. A. 2010. Global distribution of ciguatera causing dinoflagellates in the genus Gambierdiscus. Toxicon 56:711-730.   DOI
64 Mangialajo, L., Ganzin, N., Accoroni, S., Asnaghi, V., Blanfune, A., Cabrini, M., Cattaneo-Vietti, R., Chavanon, F., Chiantore, M., Cohu, S., Costa, E., Fornasaro, D., Grossel, H., Marco-Miralles, F., Maso, M., Rene, A., Rossi, A. M., Sala, M. M., Thibaut, T., Totti, C., Vila, M. & Lemee, R. 2011. Trends in Ostreopsis proliferation along the Northern Mediterranean coasts. Toxicon 57:408-420.   DOI
65 You, J. H., Jeong, H. J., Kang, H. C., Ok, J. H., Park, S. A. & Lim, A. S. 2020. Feeding by common heterotrophic protist predators on seven Prorocentrum species. Algae 35:61-78.   DOI
66 Zingone, A., Siano, R., D'Alelio, D. & Sarno, D. 2006. Potentially toxic and harmful microalgae from coastal waters of the Campania region (Tyrrhenian Sea, Mediterranean Sea). Harmful Algae 5:321-337.   DOI
67 Jeong, H. J., Ok, J. H., Lim, A. S., Kwon, J. E., Kim, S. J. & Lee, S. Y. 2016. Mixotrophy in the phototrophic dinoflagellate Takayama helix (family Kareniaceae): predator of diverse toxic and harmful dinoflagellates. Harmful Algae 60:92-106.   DOI
68 Jeong, H. J., Yih, W., Kang, N. S., Lee, S. Y., Yoon, E. Y., Yoo, Y. D., Kim, H. S. & Kim, J. H. 2012b. First report of the epiphytic benthic dinoflagellates Coolia canariensis and Coolia malayensis in the waters off Jeju Island, Korea: morphology and rDNA sequences. J. Eukaryot. Microbiol. 59:114-133.   DOI
69 Sato, S., Nishimura, T., Uehara, K., Sakanari, H., Tawong, W., Hariganeya, N., Smith, K., Rhodes, L., Yasumoto, T., Taira, Y., Suda, S., Yamaguchi, H. & Adachi, M. 2011. Phylogeography of Ostreopsis along west Pacific coast, with special reference to a novel clade from Japan. PLoS ONE 6:e27983.   DOI
70 Sherr, E. B. & Sherr, B. F. 2007. Heterotrophic dinoflagellates: a significant component of microzooplankton biomass and major grazers of diatoms in the sea. Mar. Ecol. Prog. Ser. 352:187-197.   DOI
71 Shah, M. M. R., Samarakoon, K. W., Ko, J. -Y., Lakmal, H. H. C., Lee, J. -H., An, S. -J., Jeon, Y. -J. & Lee, J. -B. 2014b. Potentiality of benthic dinoflagellate cultures and screening of their bioactivities in Jeju Island, Korea. Afr. J. Biotechnol. 13:792-805.   DOI
72 Graneli, E., Vidyarathna, N. K., Funari, E., Cumaranatunga, P. R. T. & Scenati, R. 2011. Can increases in temperature stimulate blooms of the toxic benthic dinoflagellate Ostreopsis ovata? Harmful Algae 10:165-172.   DOI
73 Hallegraeff, G. M. 2010. Ocean climate change, phytoplankton community responses, and harmful algal blooms: a formidable predictive challenge. J. Phycol. 46:220-235.   DOI
74 Shah, M. M. R., An, S. -J. & Lee, J. -B. 2013a. Presence of benthic dinoflagellates around coastal waters of Jeju Island including newly recorded species. J. Ecol. Environ. 36:347-370.   DOI
75 Shah, M. M. R., An, S. -J. & Lee, J. -B. 2013b. Seasonal abundance of epiphytic dinoflagellates around coastal waters of Jeju Island, Korea. J. Mar. Sci. Technol. 21:156-165.
76 Shah, M. M. R., An, S. -J. & Lee, J. -B. 2014a. Occurrence of sand-dwelling and epiphytic dinoflagellates including potentially toxic species along the coast of Jeju Island, Korea. J. Fish. Aquat. Sci. 9:141-156.   DOI
77 Han, I. -S. & Lee, J. -S. 2020. Change the annual amplitude of sea surface temperature due to climate change in a recent decade around the Korean Peninsula. J. Korean Soc. Mar. Environ. Saf. 26:233-241.   DOI
78 Han, Y. B. 2010. Edible macroalgae 1. Component and bioactivity. Korea University Press, Seoul, 416 pp.
79 Holmes, M. J., Brust, A. & Lewis, R. J. 2014. Dinoflagellate toxins: an overview. In Botana, L. M. (Ed.) Seafood and Freshwater Toxins: Pharmacology, Physiology, and Detection. 3rd ed. Marcel Dekker, New York, pp. 3-38.
80 Yoshimatsu, T., Yamaguchi, H., Iwamoto, H., Nishimura, T. & Adachi, M. 2014. Effects of temperature, salinity and their interaction on growth of Japanese Gambierdiscus spp. (Dinophyceae). Harmful Algae 35:29-37.   DOI
81 Shears, N. T. & Ross, P. M. 2009. Blooms of benthic dinoflagellates of the genus Ostreopsis: an increasing and ecologically important phenomenon on temperate reefs in New Zealand and worldwide. Harmful Algae 8:916-925.   DOI
82 Kohli, G. S., Murray, S. A., Neilan, B. A., Rhodes, L. L., Harwood, D. T., Smith, K. F., Meyer, L., Capper, A., Brett, S. & Hallegraeff, G. M. 2014. High abundance of the potentially maitotoxic dinoflagellate Gambierdiscus carpenteri in temperate waters of New South Wales, Australia. Harmful Algae 39:134-145.   DOI
83 Laza-Martinez, A., Orive, E. & Miguel, I. 2011. Morphological and genetic characterization of benthic dinoflagellates of the genera Coolia, Ostreopsis and Prorocentrum from the south-eastern Bay of Biscay. Eur. J. Phycol. 46:45-65.   DOI
84 Kim, H. S., Yih, W., Kim, J. H., Myung, G. & Jeong, H. J. 2011. Abundance of epiphytic dinofalgellates from coastal waters off Jeju Island, Korea during Autumn 2009. Ocean Sci. J. 46:205-209.   DOI
85 Kim, S. & Seo, H. 2019. Distribution and molecular phylogeny of the toxic benthic dinoflagellate Ostreopsis sp. in the coastal waters off Jeju Island, Korea. The Sea 24:236-248.   DOI
86 Korea Fisheries Resources Agency. 2020. Marine forests white paper, Korea. Korea Fisheries Resources Agency, Busan, 368 pp.
87 Kwon, J. E., Jeong, H. J., Kim, S. J., Jang, S. H., Lee, K. H. & Seong, K. A. 2017. Newly discovered role of the heterotrophic nanoflagellate Katablepharis japonica, a predator of toxic or harmful dinoflagellates and raphidophytes. Harmful Algae 68:224-239.   DOI
88 Larsson, M. E., Smith, K. F. & Doblin, M. A. 2019. First description of the environmental niche of the epibenthic dinoflagellate species Coolia palmyrensis, C. malayensis, and C. tropicalis (Dinophyceae) from Eastern Australia. J. Phycol. 55:565-577.   DOI
89 Tawong, W., Yoshimatsu, T., Yamaguchi, H. & Adachi, M. 2015. Effects of temperature, salinity and their interaction on growth of benthic dinoflagellates Ostreopsis spp. from Thailand. Harmful Algae 44:37-45.   DOI
90 Tanimoto, Y., Yamaguchi, H., Yoshimatsu, T., Sato, S. & Adachi, M. 2013. Effects of temperature, salinity and their interaction on growth of toxic Ostreopsis sp. 1 and Ostreopsis sp. 6 (Dinophyceae) isolated from Japanese coastal waters. Fish. Sci. 79:285-291.   DOI
91 Tawong, W., Yoshimatsu, T., Yamaguchi, H. & Adachi, M. 2016. Temperature and salinity effects and toxicity of Gambierdiscus caribaeus (Dinophyceae) from Thailand. Phycologia 55:274-278.   DOI
92 Terrado, R., Vincent, W. F. & Lovejoy, C. 2009. Mesopelagic protists: diversity and succession in a coastal Arctic ecosystem. Aquat. Microb. Ecol. 56:25-39.   DOI
93 Lee, B. & Park, M. G. 2020. Distribution and genetic diversity of the toxic benthic dinoflagellate genus Ostreopsis in Korea. Harmful Algae 96:101820.   DOI
94 Leaw, C. -P., Lim, P. -T., Cheng, K. -W., Ng, B. -K. & Usup, G. 2010. Morphology and molecular characterization of new species of thecate benthic dinoflagellate, Coolia malayensis sp. nov. (Dinophyceae). J. Phycol. 46:162-171.   DOI
95 Leaw, C. P., Tan, T. H., Lim, H. C., Teng, S. T., Yong, H. L., Smith, K. F., Rhodes, L., Wolf, M., Holland, W. C., Vandersea, M. W., Litaker, R. W., Tester, P. A., Gu, H., Usup, G. & Lim, P. T. 2016. New scenario for speciation in the benthic dinoflagellate genus Coolia (Dinophyceae). Harmful Algae 55:137-149.   DOI
96 Lee, B. & Park, M. G. 2018. Genetic analyses of the rbcL and psaA genes from single cells demonstrate a Rhodophyte origin of the prey in the toxic benthic dinoflagellate Ostreopsis. Front. Mar. Sci. 5:217.   DOI
97 Kibler, S. R., Litaker, R. W., Holland, W. C., Vandersea, M. W. & Tester, P. A. 2012. Growth of eight Gambierdiscus (Dinophyceae) species: effects of temperature, salinity and irradiance. Harmful Algae 19:1-14.   DOI
98 Lee, K. -W., Kang, J. -H., Baek, S. -H., Choi, Y. -U., Lee, D. -W. & Park, H. -S. 2014. Toxicity of the dinoflagellate Gambierdiscus sp. toward the marine copepod Tigriopus japonicus. Harmful Algae 37:62-67.   DOI
99 Lee, L. K., Lim, Z. F., Gu, H., Chan, L. L., Litaker, R. W., Tester, P. A., Leaw, C. P. & Lim, P. T. 2020. Effects of substratum and depth on benthic harmful dinoflagellate assemblages. Sci. Rep. 10:11251.   DOI
100 Kwak, M. -T., Seo, G. -H., Cho, Y. -K., Kim, B. -G., You, S. -H. & Seo, J. -W. 2015. Long-term comparison of satellite and in-situ sea surface temperatures around the Korean Peninsula. Ocean Sci. J. 50:109-117.   DOI
101 Jang, S. H., Jeong, H. J. & Yoo, Y. D. 2018. Gambierdiscus jejuensis sp. nov., an epiphytic dinoflagellate from the waters of Jeju Island, Korea, effect of temperature on the growth, and its global distribution. Harmful Algae 80:149-157   DOI
102 Hwang, B. S., Yoon, E. Y., Jeong, E. J., Park, J., Kim, E. -H. & Rho, J. -R. 2018. Determination of the absolute configuration of polyhydroxy compound ostreol B isolated from the dinoflagellate Ostreopsis cf. ovata. J. Org. Chem. 83:194-202.   DOI
103 Hwang, B. S., Yoon, E. Y., Kim, H. S., Yih, W., Park, J. Y., Jeong, H. J. & Rho, J. -R. 2013. Ostreol A: a new cytotoxic compound isolated from the epiphytic dinoflagellate Ostreopsis cf. ovata from the coastal waters of Jeju Island, Korea. Bioorg. Med. Chem. Lett. 23:3023-3027.   DOI
104 Hwang, E. K. & Park, C. S. 2020. Seaweed cultivation and utilization of Korea. Algae 35:107-121.   DOI
105 Jeong, H. J. 1999. The ecological roles of heterotrophic dinoflagellates in marine planktonic community. J. Eukaryot. Microbiol. 46:390-396.   DOI
106 Jeong, H. J., Lim, A. S., Franks, P. J. S., Lee, K. H., Kim, J. H., Kang, N. S., Lee, M. J., Jang, S. H., Lee, S. Y., Yoon, E. Y., Park, J. Y., Yoo, Y. D., Seong, K. A., Kwon, J. E. & Jang, T. Y. 2015. A hierarchy of conceptual models of red-tide generation: nutrition, behavior, and biological interactions. Harmful Algae 47:97-115.   DOI
107 Sung, J. W. 2015. Phylogenetic position, morphological and physiological characterization of five benthic epiphytic dinoflagellates isolated from Korean coastal waters. M.S. thesis, Hanyang University, Seoul, Korea, 69 pp.