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Review Algal Resources (2015) 8:155-163

Harvest and cultivation of japonica in Northern Hokkaido, Japan, and southern Sakhalin and Primorye, Russia: A review

Tadashi KAWAI1 *, Dmitrii GALANIN2,TatianaKRUPNOVA3 and Norishige YOTSUKURA4

Abstract : History, catch, gear, propagation of resources in the wild, cultivation, and utilization of the in northern Hokkaido, Japan, and southern Sakhalin and Primorye, Russia are compared in this review of the fisheries in these areas. Sea currents (especially, warm currents) commonly affect the occurrence and production of this kelp in those regions. Techniques in propagation of S. japonica in the wild and by cultivation has been developed in both Hokkaido and Primorye. The total annual amount of harvest and season were regulated in fisheries in Sakhalin, though only season of the harvest is regulated in Hokkaido. Harvest and cultivation of kelp in three regions have many similarities.

Keywords : comparison, difference, Hokkaido, Primorye, Saccharina, Sakhalin, similarity

Introduction

Seaweeds of the Saccharina form kelp forests along the coasts of Hokkaido, Japan, and Sakhalin and Primorye in the Russian Far East (Sukhoveeva 1967; Krupnova 2013; Tokida 1954; Yotsukura et al. 1999; Kawai et al. 2014) (Fig. 1). They are the main constituents of rich marine forests (Klochkova 1996; Koroleva 2004, 2010; Saushkina 2006; Kusidi 2007), and the Saccharina are economically important in these areas (Krupnova 2004; Galanin and Repnikova 2014; Hatamiya 2014). The present status of kelp fisheries in Hokkaido has been well reported (e.g., Hatamiya 2014; Kuribayashi and Akaike 2014; Terai 2014; Watanuki 2014), and there are a few reports from southern Sakhalin (Galanin and Repnikova 2014). How- ever, information on kelp fisheries and aquac- Fig. 1. Regions of kelp fisheries in this review. A. ulture on the coast of Primorye is scarce. Pri- Primorye, Russian Far East; B. Northern morye region locate in eastern coast of main- Hokkaido,Japan;C,SouthernSakhalin.

1 Wakkanai Fisheries Research Institute,4-5-15 Suehiro, Wakkanai, Hokkaido 097-0001, Japan 2 Sakhalin Scientific Research Institute of Fisheries and Oceanography (SakhNIRO), 306, 196, Komsomolskaya Street, Yuzhno-Sakhalinsk, 693023, Russia 3 Pacific Scientific Research Fisheries Centre (TINRO), 4, Shevchenko Alley, Vladivostok, 690950, Russia. 4 Field Science Center for Northern Biosphere, Hokkaido University, North 9, West 9, Kita-ku, Sapporo, Hokkaido 060-0809, Japan *Corresponding author : Tel: +81-162-32-7188, fax: +81-162-32-7171, e-mail: [email protected] 155 Tadashi KAWAI, Dmitrii GALANIN, Tatiana KRUPNOVA and Norishige YOTSUKURA land of Russian Far East, terminal end of warm ever, tillage and weed control stopped in the sea current reach this area (Krupnova 1994, 2010, 1990s due to imbalance between costs and ben- 2013), so oceanographic condition might be efits. Kelp cultivation was initiated in 1975, and similar to Sea of Japan side of Southern cultivation technology has been developed and Sakhalin and Hokkaido. The present paper re- established with two-year cultivation cycles that views information on kelp fisheries and com- take total two years from seedling to harvest, pare them among northern Hokkaido, southern the cycle are continuing since the 1980s. Sakhalin, and Primorye to provide a better un- b. Sakhalin: The first scientific report on com- derstanding of the issues facing kelp fisheries. mercial fisheries of the kelp S. japonica along This comparison identifies the commonalities Sakhalin Island was published by Yu. P. and uniquenesses of the three localities with Schmidt (1905) stated that the total harvest from regard to the sustainable utilization of kelp kelp fisheries in Sakhalin was low until 1905, resources and promotion of the kelp industry. and the volume was not recorded at that time. To our knowledge, this is the first review to In order to explore the kelp resources, scientific introduce the current status of kelp in the studies were started during the 1940s and 1950s, Russian Far East in the English scientific liter- under the supervision of G. I. Gajl who is ature, which will aid in international research pioneer of specialist of algal resources. From the on kelp. 1950s to the 1970s, a researcher from TINRO Over 10 Saccharina species are distributed (The Pacific Scientific Research Fisheries Cen- along the coast of Hokkaido (Yotsukura et ter), V. F. Sarochan, studied the biology and dis- al. 1999). Saccharina japonica, S. religiosa, S. tribution of commercial species of Laminariales, ochotensis, and S. diabolica are now considered including S. japonica, on Sakhalin. SakhNIRO four varieties of one species, S. japonica, and the (The Sakhalin Fisheries and Oceanography In- presence of S. japonica var. ochotensis has been stitute) conducted further annual expeditions confirmed at northern Hokkaido (Yotsukura et and algologists others surveyed the coastal zone al. 2008). Another variety of S. japonica is found of Sakhalin Island (Schmidt 1905; Gajl 1949; in Primorye and southern Sakhalin, but the Zinova 1954; Sarochan 1963; Sukhoveeva 1969, scientific name of this variety is still unclear. 1971, Balkonskaya and Shpakova 1998; Galanin Basedonthistaxonomicsituation,wedefinethe and Repnikova 2014). kelp as S. japonica for the purposes of this c. Primorye: S. japonica is one of the most review. abundant species of brown along the coast of Primorye facing to the Sea of Japan. This A. History and structure of the kelp industry kelp was the dominant species of algae on the (production and marketing) seafloor along the coast of Primorye until ap- proximately the 2000s. The coast in this region a. Hokkaido: Statistics by the Hokkaido gov- was known for its rich fields of Saccharina and ernment show the kelp production in dry weight the production was estimated from 500,000 to in northern Hokkaido as follows: In 1935, the 700,000 t year-1 wet weight. The kelp resources annual production exceeded 26,000 t; from 1935 have contributed also as a food source for sea to 1990, annual production ranged from 2,000 to urchins. Since 1932, a fishing gear called“Kanza” 6,000 t; and following 2005, it dropped to less in Russian (see details in the following section than 2,000 t (Hatamiya 2014). After 1935, stone C) had been introduced in kelp fisheries along castings and concrete substratum were provid- the coast of the Primorye region, and the total ed to the seafloor in kelp fishery grounds to production had dropped to around 1000 t year-1 help propagate kelp resources. After 1980, sea- after 1933 due to change of oceanographic con- floor tillage and weed control were carried out dition. From 1960 to 1978, diving techniques were near the islands of Rebun and Rishiri; ca. 1985, adapted for collecting kelp, and harvest peaked a total of 500 kelp propagation machines, for at more than 200,000 t year-1. In this duration, example tillage using chain, were in use. How- kelp covered almost the whole substratum from

156 Harvest and cultivation of Saccharina japonica in Northern Hokkaido, Japan, and southern Sakhalin and Primorye, Russia: A review the shoreline to a depth of 20 m. However, in the 2004). Instead of kelp, the seafloor has been 1980s, the production of kelp sharply decreased, occupied by Corallinaceae algae, which is the to a low of 4,000 to 80,000 t year-1. It was sug- phenomenon known as barren ground or so- gested that change of oceanographic condition called “ Isoyake ” in South-western Hokkaido, occurred and water temperature in coast in- Japan and Southern Sakhalin (Krupnova 2008) creased, in general warmer temperature is not (Fig. 2). Changes in the oceanographic structure appropriate for growth of kelp (Krupnova 2002, of the Primorye Cold Current, as it is known in Russian (Liman Current in Japanese), are sug- gested as the major reason for this fluctuation ˝ in algal flora (Krupnova 1994). It is known from previous studies that the cold current flows along the Primorye coast and the conditions of the current affect the resources of S. japonica. Both water temperature and nutrients influence the growth of kelp (Krupnova 2002).

B. Kelp harvested

a. Hokkaido: In fisheries on natural kelp for- ests and cultivation, second-year kelp individuals are commonly harvested (Nabata and Takiya 2003). ˞ b. Sakhalin: In kelp fisheries 1.5-2-year-old in- dividuals are harvested, but cultivation of kelp is not developed at Sakhalin (Ministry of Justice of the Russian Federation 2011, 2012; Sukhoveeva 1969). c. Primorye: In fisheries on natural kelp forests and cultivation, second-year kelp individuals are commonly harvested.

C. Gear

a. Hokkaido: In general, fishermen catch kelp ˟ using poles that are several meters in length and equipped with a hook at the end (Nabata and Takiya 2003). b. Sakhalin: Fishermen catch kelp from fishery boats using poles with hooks or by trawling; other fishermen collect drifting kelp or directly harvest it from a kelp forest. Fishery divers also collect kelp (Gajl 1949; Sukhoveeva and Podkorytova 2006; Galanin and Repnikova 2014). c. Primorye: Fishermen catch kelp using poles that are several meters in length and equipped with a hook at the end, called“Kanza.” Fig. 2. Underwater photo at the kelp fisheries. So far coralline algae dominate the seafloor of D. Propagation of resources in the wild deeper area in fisheries ground and also the kelp forest has disappeared there. A. a. Hokkaido: To conserve the kelp forest, the Southern Primorye, B. Southern Sakhalin, C. manager of a fishing ground carried out weed Northern Hokkaido. control; this was conducted during winter, which

157 Tadashi KAWAI, Dmitrii GALANIN, Tatiana KRUPNOVA and Norishige YOTSUKURA is the reproductive season for kelp (Nabata and carried out during December, fishermen collect Takiya 2003). mother kelp (larger biennial plants that ap- b. Sakhalin: There is no artificial propagation in peared sori) for seeding and the kelp are set up this area (Gajl 1949; Sarochan 1963; Galanin et in indoor cultivation tanks, then zoospore re- al. 2010; Evseeva and Repnikova 2010). leased from the mother kelp and the zoospore c. Primorye: Thus far, artificial propagation attached the rope in the cultivation tank. After similar to the Japanese method has not been several hours, the ropes that attached zoospore developed in this area, but bottom-dwelling cul- in cultivation tank are took out and brought to tivation (see details in the following section F) sea. In December of the following year, 4 to 5 contributes to the restoration of kelp stock and kelp individuals on the cultivation rope are some kelp individuals drift out from the rope. thinned and unstable holdfasts bound with tape. These provide food for sea urchins on the sea- These are inserted into the cultivation rope. In floor. August of the second year, the cultivated kelp is harvested. E. Regulation of fishery season or total catch b. Sakhalin: Kelp is harvested only in fishing amount grounds, but the possibility of cultivation has a. Hokkaido: The Fisheries Cooperative Associ- been studied since the 1960s. In recent years, on ation prohibits harvest of first-year kelp and the southwest coast of Sakhalin, these studies permits only summer season harvest, generally were required because of insufficient volume of after July when the kelp is fully grown and sori are not present (Nabata and Takiya 2003). b. Sakhalin: Harvest of first-year kelp is pro- ˝ hibited, and is allowed only in the case of drifted first-year kelp on the beach. From May 1 to September 30, fishermen can harvest kelp on the southwest coast (Ministry of Justice of the Russian Federation 2011, 2012; Galanin and Repnikova 2014). c. Primorye: A protected area has been estab- lished in southwestern Great Peter Bay in the Primorye region. Only scientific research collec- tion of kelp is allowed in this area. ˞

F. Cultivation a. Hokkaido: The following information is based on Hasegawa (1971a, 1971b), Hasegawa and Sanbonsuga (1971), Kawashima (1984), and Ohno and Largo (1998). Three methods are employed for kelp cultivation and are used in the various cultivation grounds in Hokkaido: 1) Two-year cultivation; 2) Cultivation by transplanting (wild and small kelp individuals were knotted on cultivation rope); and 3) Forced cultivation (one year cultivation). All of the cultivation methods were in use in northern Hokkaido, but two-year Fig. 3. Hanging cultivation system for Saccharina cultivation was the most dominant. Two-year japonica in Primorye in the Russian Far cultivation is the traditional method, and it was East. A. Early-stage kelp on cultivation rope; applied as the general cultivation technique near B. Cultivated kelp just before harvest; these the islands of Rebun and Rishiri. Seeding is kelp were thinned to remove excess plants.

158 Harvest and cultivation of Saccharina japonica in Northern Hokkaido, Japan, and southern Sakhalin and Primorye, Russia: A review resources in fishing grounds off the west coast cultivation rope are thinned and holdfasts di- of Sakhalin Island (Sarochan and Vinokurova rectly inserted into the rope (Fig. 3). In the 1977; Evseeva and Repnikova 2010; Liu et al. other method, which is a bottom-dwelling culti- 2012; Galanin and Repnikova 2014). vation system, zoospores of kelp are attached to c. Primorye: Cultivation of kelp S. japonica along the net, then transported to sea and hung (Fig. the coast of Primorye started in 1974. From 1974 4). to 2000, hanging plantations were set up in three G. Utilization areas: 1) Southern Primorye, Great Peter Bay near Nahodka (approximate 30 ha); 2) Middle a. Hokkaido: Kelp is mainly used to make Primorye, near Valentin Bay (appox. 100 ha); “Dashi,”a Japanese traditional soup, its special and 3) Northern Primorye, near Oprichnik Bay flavor forms the base of many Japanese tradi- (approx. 60 ha). The crop from these planta- tional foods. Another, kelp is fed as stuffs of tions totals 50-100 t ha-1.Therearetwometh- Japanese popular foods such as“Oboro Kom- ods of cultivation used in hanging plantations. bu”and“Tororo ”(Nabata and Takiya In the first method, kelp individuals on the 2003).

˝ ˞

˟ ˠ

ˡ ˢ

Fig. 4. Bottom-dwelling cultivation system for kelp Saccharina japonica in Primorye in the Russian Far East. A. Mother plant; B. Net for seeding; C. Zoospore attaching to the net; D. Seeding net placed in the sea; E. Plastic buoys for the seeding net; F. Kelp growth on the net.

159 Tadashi KAWAI, Dmitrii GALANIN, Tatiana KRUPNOVA and Norishige YOTSUKURA

˝ ˞

˟ ˠ

ˡ ˢ

Fig. 5. Utilization of the kelp Saccharina japonica in Primorye in the Russian Far East. A. Kelp aquaculture facilities; B. Harvested kelp; C. Processing machine for harvested kelp; D. Processing of the kelp; E. Processed kelp; F. Shredded and dried kelp at market. b. Sakhalin: Kelp is used as food (Evseeva and is sharply decreasing due to decline of the kelp Repnikova 2010; Galanin and Repnikova 2014); forest. Since these three areas are located at the raw kelp fronds are chopped and sliced for terminal end of the warm current, oceano- seafood salads (Evseeva and Repnikova 2010; graphic conditions along the coasts where kelp Galanin and Repnikova 2014). forests are formed change dramatically. The c. Primorye: Raw or dried kelp is only used in scale of the forests fluctuate greatly and this seafood dishes (see Fig. 5). fluctuation corresponded with the path of the warm current, when the warm current comes Discussion near the coast, the forest can decline sharply. We suggest this environmental fluctuation af- It is common in northern Hokkaido, southern fects the occurrence of the kelp species and Sakhalin, and southern Primorye that the recent cause their annual volumes to fluctuate. Indeed, volume of S. japonica catch in the kelp industry the initial growth and maturation rates of young

160 Harvest and cultivation of Saccharina japonica in Northern Hokkaido, Japan, and southern Sakhalin and Primorye, Russia: A review kelp depend on water temperature and nutri- Gajl GI. Promyslovyje vodorosli Sakhalina I ent concentration under laboratory conditions Kurilskoi gryady. TINRO, Vladivostok. 1949 ; (Hasegawa et al. 1971). Attention should be given 1-87 (in Russian). to the conditions in oceanographic conditions Galanin D, Balkonskaya L, Prokhorova N. along the coasts which are affected by sea cur- Resources of (Saccharina) japonica rents, particularly, the water temperature and on the southwestern coast of Sakhalin is- nutrient levels in further kelp studies. land in recent years. Tasks of investiga- Various techniques in propagation of re- tions for the near period. Bull. Fish. Res. sources of S. japonica in the wild is developing Age. 2010 ; 32 : 3-46 (in Russian). in both Hokkaido and Primorye, but not in Galanin D, Repnikova A. Present situation of Sakhalin. However, regulation on the total catch fisheries research on kelp in southern Sa- and season in the kelp fisheries was present khalin, Russia. Fish. Engineer. 2014 ;51:65-69 only in Sakhalin. Japanese kelp fisheries regu- (in Japanese with English abstract). lated only the season, and the Primorye region Hasegawa Y. Cultivation of Laminaria in Japan. has a catch prohibition area. These three areas Bull. Hokkaido Reg. Fish. Res. Lab. 1971a ; commonly catch and harvest only second-year 37 :46-48. kelp individuals, and harvested kelp are used as Hasegawa Y. Forced cultivation of Laminaria in foods. Gear used to catch kelp in the wild is Japan. Bull. Hokkaido Reg. Fish. Res. Lab. similar in the three areas, but diving collection 1971b ; 37 : 49-52. is only carried out in Sakhalin. Cultivation is Hasegawa Y, Sanbonsuga Y. Laboratory rearing being conducted at Hokkaido and Primorye and of the laminariaceous plants. Bull. Hokkaido will be tried in the Sakhalin region. The culti- Reg. Fish. Res. Lab. 1971 ; 37 : 53-58. vation procedures in Hokkaido and Primorye Hatamiya T. Policy and direction of Hokkaido share many similarities. Catch, gear, propaga- government for suitable fisheries of kelp tion of resources in the wild, regulation of resource. Fish. Engineer. 2014 ; 51 : 75-77 (in fishery season or total catch amount, cultiva- Japanese with English abstract). tion, and utilization in these three areas show Kawai T, Galanin D, Tsukhay Z, Latokovskaya small differences, but have many similarities. E, Nagai N, Yotsukura N. Relationship be- tween occurrence of kelp species and water Acknowledgments temperature in northern Hokkaido, Japan, and southern Sakhalin, Russia. Algal Re- Our sincere thanks to Professor Nina G. sources 2014 ; 7 : 107-116. Klochkova of Kamchatka State Technical Uni- Kawashima S. Kombu cultivation in Japan for versity, who coordinated our interview research human foodstuff. Jap. J. Phycol. 1984 ;32:379- in Sakhalin and Vladivostok, Russia. This re- 394. search was supported by the Japan Society for Klochkova NG. Flora of of Tatar Strait the Promotion of Science (JSPS) Grant-in-Aid for (Sea of Japan) and features of its forma- Science research, 25304010. tion. Dalnauka, Vladivostok. 1996 (in Rus- sian). References Koroleva TN. Development of brown alga Laminaria bongardiana P. et R. in the wa- Balkonskaya LA, Shpakova TA. Change of veg- ters off Kamchatka. Dissertation of Candi- etative communities at southwest Sakhalin date of Biological Science. Moscow. 2004 (in (Nevelsk-Sadovniki). Problem hydrobiologi- Russian). cal researches of an island Sakhalin. Book Koroleva TN. Development of brown alga of VNIRO, 1998 ; 1-5 (in Russian). Saccharina bongardiana and adaptation to Evseeva NV, Repnikova AR. Resources com- anthropogenic pollution. KamchatSTU, mercial algae Sakhalin and Kuril region. Petropavlovsk-Kamchatsky. 2010 (in Rus- Rybprom 2010 ; 3 : 14-21 (in Russian). sian).

161 Tadashi KAWAI, Dmitrii GALANIN, Tatiana KRUPNOVA and Norishige YOTSUKURA

Krupnova TN. Regularity Laminaria japonica Ministry of Justice of the Russian Federation. reproduction as base of mariculture. Dis- Rules of fisheries for Far East fishery re- sertation of Candidate of Biological Science. gion, confirmed by the Order of Federal Vladivostok. 1994 ; 201 (in Russian with Agency on Fisheries on 6 th July 2011, Far English abstract). East, 2011 (in Russian) . Krupnova TN. Development of sporogenous tis- Ministry of Justice of the Russian Federation. sue in Laminaria japonica under the influ- The general admissible catch of water bio- ence of changing environment. Izv. TINRO. logical resources for 2012, confirmed by the 2002 ; 130 : 474-482 (in Russian with English Order of Federal Agency of Fisheries, on 5 th abstract). October 2011, Moscow, 2012 ; 983 (in Rus- Krupnova TN. Influence of oceanology-climatic sian). factors on dynamics of Laminaria japonica Nabata S, Takiya A. Laminaria ochotensis Miy- Aresch. fields in northwest part of the Sea abe. In: Ueda Y, Maeda K, Shimada K, of Japan. Sea Hydrology and Hydrochemis- Takami T (eds). Fisheries and aquatic life try 2004 ; 8 : 162-166 (in Russian with English in Hokkaido. The Hokkaido Shinbun Press, abstract). Sapporo. 2003 ; 428-431. Krupnova TN. Instruction on Cultivation and Ohno M, Largo DB. The seaweed resources of Recovery of Laminaria Fields. Pacific Japan. In: Critchley AT, Ohno M (eds). Research Fisheries Centre (TINROCentre), Seaweed Resources of the world. Japan In- Vladivostok. 2008 (in Russian with English ternational Cooperation Agency, Kanagawa abstract). International Fisheries Training Centre, 1998; Krupnova TN. Reorganization of Laminaria fields 1-14. under the influence of natural and anthro- Sarochan VF. Biology Japanese kelp at south- pogenic factors. TINRO archive 2010 ; 26962 : west coast of Sakhalin. Izv. TINRO. 1963;49: 56 (in Russian with English abstract). 115-136 (in Russian). Krupnova TN. On possible causes of lowering Sarochan VF, Vinokurova NV. Bioecological reproduction potential and growing small prerequisite and principles of cultivation of for laminaria Saccharina japonica at the luminaria in the soviet far east. Presented coast of Primorye. Izv. TINRO. 2013 ; 175 : at the 2nd Soviet-Japan Joint Symposium on 93-100 (in Russian with English abstract). Aquaculture, November 1973. Moscow. Vla- Kuribayashi T, Akaike S. Effects of nitrogen divostok. TINRO. 1977 ; 121-135. fertilization with density control of sea ur- Sauchkina LN. Developmental biology of some chins to reform community in the species of the genus Laminaria in Kam- southwest coast of Japan Sea off Hokkaido, chatka waters. Dissertation of Candidate using  15N signatures in algal tissue of of Biological Science. Petropavlovsk-Kam- Saccharina japonica var. religiosa (Miyabe). chatsky. 2006 (in Russian). Fish. Engineer. 2014 ; 51 : 47-54 (in Japanese Schmidt Yu. P. Sea crafts of island Sakhalin. with English abstract). St. -Petersburg Printing house of GP, Pozha- Kusidi AE. Developmental biology of some spe- rova. 1905 ; 1-458. cies of the genus Laminaria in Kamchatka Sukhoveeva MV. The algae distribution of along waters. Dissertation of Candidate of Biolog- the Primorye coast region. Izv. TINRO. 1967; ical Science. Petropavlovsk-Kamchatsky. 2007 61 : 255-260. (in Russian). Sukhoveeva MV. The distribution and resources Liu F, Yao J, Wang X, Repnikova A, Galanin condition of kelp and some other seaweeds DA, Duan D. Genetic diversity and struc- ofcoastofthemaritimeProvinceofSiberia. ture within and between wild and cultivat- Izv. TINRO. l969 ; 73 : 3-23. ed Saccharina japonica (Laminariales, Pha- Sukhoveeva MV. Laminaria japonica Aresch. eophyta) revealed by SSR markers. Aquac- and seaweed accompanying it. Izv. TINRO. ulture 2012 ; 358-359 : 39-45. 1971 ; 75 : 152-155 (in Russian).

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Sukhoveeva MV, Podkorytova AV. Harvesting 1999 ; 47 : 71-80. algae and grass Far Eastern seas: biology, Watanuki A. Development and maintain of kelp distribution, inventory, processing technolo- forest in barren ground using adaptive gy. TINRO, Vladivostok. 2006 ; 243. management process: a review. Fish. Engi- Terai M. Conservation of Kelp Saccharina neer. 2014 ; 51 : 33-37 (Japanese with English longissimima ground in eastern Hokkaido, abstract). Japan by removing weed control. Fish. En- Yotsukura, N, Kawashima S, Kawai T, Abe T, gineer. 2014 ; 51 : 55-58 (Japanese with English Druehl LD. A systematics re-examination of abstract). four Laminarian species: L. japonica, L. Tokida J. The marine algae of southern Sagha- religiosa, L. ochotensis and L. diabolica. J. lin. Mem. Fac. Fish. Hokkaido Univ. 1954 ; Jap. Bot. 2008 ; 83 : 165-176. 2 : 1-264. Zinova ES. Seaweed of Tatar strait. Works of Yotsukura N, Denboh T, Motomura T, Botanical institute AN USSR. 1954 ; 2-9 :311- Horiguchi T, Coleman AW, Ichimura T. 364. Little divergence in ribosomal DNA internal transcribed spacer-1 and -2 sequences among Received 25 November 2015 non-digitate species of Laminaria (Phaeo- Accepted 28 December 2015 phyceae) from Hokkaido, Japan. Phycol. Res.

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