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Identification of Ecklonian Kelps Which Have Similarity to Ecklonia Stolonifera Okamura (Laminariales, Phaeophyta), in Nishinoshima Coast of Oki Islands, Shimane Prefecture, Japan

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Identification of Ecklonian Kelps Which Have Similarity to Ecklonia Stolonifera Okamura (Laminariales, Phaeophyta), in Nishinoshima Coast of Oki Islands, Shimane Prefecture, Japan Algal Resources (2017) 10：1-16 Identification of ecklonian kelps which have similarity to Ecklonia storonifera Okamura (Laminariales, Phaeophyta), in Nishinoshima coast of Oki Islands, Shimane Prefecture, Japan Yuichi HAYASHI1 *, Masahiro NOTOYA2 and Norishige YOTSUKURA3 Abstract : The habitat, morphology, and propagation characteristics were examined on ecklonian kelps in Nishinoshima coast of Oki Islands that were difficult to identify. In addition, cultivation experiments and DNA sequencing on ITS-1 region were conducted. The data obtained were compared with those of Ecklonia stolonifera and E. kurome, and taxonomical discussion was done. The stolon of E. stolonifera was characterized by its small number of branches and cteno-rootlet on the back of the root. The root of E. kurome was characterized by multiple branching and rootlet at the tip of the root. The unidentifiable kelps were divided into two types (AK-1 and AK-2) according to the presence or absence of shoot at the tip of the stolon. Furthermore, plants without shoot (AK-2) were divided into two subtypes (AK-2(1) and AK-2(2)) based on the difference of morphology of holdfast. By morphological principal components analysis and sequence comparison, plants that have shoot (AK-1) and plants that have a root with the feature of E. stolonifera but have no shoot (AK-2(1)) were considered to be E. stolonifera. Furthermore, it was inferred that plants that have roots with the feature of both E. stolonifera and E. kurome and have no shoot (AK-2(2)) were sporophytes from natural crossbreeding between two ecklonian species. Keywords : asexual reproduction, ecklonian kelp, Ecklonia stolonifera, Oki Islands, propagation characteristic, sequence comparison Introduction Terawaki and Arai 2004). Ecklonia stolonifera is distributed along the coast of the Japan Sea in Ecklonia stolonifera Okamura is a kelp be- Japan and Korea, which is influenced by the longing to the family Lessoniaceae, in the or- Tsushima Warm Current (Taniguchi 1969, 1987; der Laminariales (Yoshida et al. 2010). The Notoya and Aruga 1992; Kawashima 1993a), and sporophyte of the species is similar to that of this is the primary habitat of seaweed beds at another Ecklonia species, E. kurome Okamura, adepthof2-35 m (Notoya 1995, 2003). Con- in that it has a blade with rugae. The former is versely, E. kurome grows along the coast of the distinguished from the latter by an asexual re- Pacific and the Seto Inland Sea, mainly in production ability that produces shoot at the tip Japan. However, it is also observed at a depth of the stolon (Okamura 1927, 1936; Yoshida and of 5-40 m in the Japan Sea in Japan (Ishida 1995, Terawaki 1990; Kawashima 1993a; Yoshida 1998; 1998), although not frequently, and therefore, the 1 Marine Bussiness Division, Okabe Company Limited, Oshiage 2-8-2, Sumida-ku, Tokyo 131-8505, Japan 2 Research Institute of Applied Phycology, Fukui 1237, Ama, Oki, Shimane 684-0404, Japan 3 Field Science Center for Northern Biosphere, Hokkaido Univercity, Kita9 Nishi9, Kita-ku, Sapporo, Hokkaido 060-0809, Japan ＊Corresponding author : E-mail: [email protected] 1 Yuichi HAYASHI, Masahiro NOTOYA and Norishige YOTSUKURA range of the two species partially overlaps. tween E. stolonifera and E. kurome based on the The two Ecklonia species can be discrimina- results of cultivation experiments and the se- ted by differences in the holdfast structure quence comparison of a specific DNA region. (Okamura 1936), but it is difficult to identify when the development of the holdfast is insuf- Materials and Methods ficient to enable these differences to be clearly visible (Arai et al. 1997ab). Incidentally, the for- Study sites mation of normal sporophytes by reciprocal The seaweed beds targeted in this study were crossing of the two species has been confirmed located along the Akanadaguchi coast of the with a high degree of interspecific cross-com- Nishinoshima Islands in Oki Islands, Shimane patibility, although the growth of hybrids and Prefecture (Fig. 1). Plants with shoot at the tip mode of asexual reproduction has not been of the stolon which were treated as AK-1 and examined in detail (Migita 1984). Furthermore, without shoot which were treated as AK-2 were the current status of natural crossing between observed here. Typical E. stolonifera plants with the two species in the coastal areas of the numerous shoots along the Hobomi coast of the Japan Sea, where both species coexist, is still Nakanoshima Islands which were treated as HO unclear. and typical E. kurome plants without shoot The decline and disappearance of ecklonian along the Sazae-jima coast of the Okinoshima seaweed beds has been reported in various re- Islands which were treated as SA were used for gions (Serisawa et al. 2004; Haraguchi et al. 2009; comparisons. The growth environment of each Fujita 2010; Hasegawa 2010; Kuwahara et al. seaweed bed was investigated and samples were 2010). However, notable expansions of seaweed collected to compare the morphology and nu- beds of E. stolonifera have occurred recently cleotide arrangement of a specific DNA region. along the northern coast of Japan (Kirihara et al. 2006). As such, more intensive conservation Substrata and growth conditions of Ecklonia of ecklonian seaweed beds is needed in order to Between July 9-10, 2005, the appearance rate protect the coastal ecosystem of the Japan Sea. of AK-1 and AK-2 plants was examined for 300 The development of techniques that will stimu- random individuals using scuba diving in a flat late the creation of seaweed beds will be nec- area (25 m×25 m) at a depth of 15 m at Aka- essary in the future to meet environmental nadaguchi (36233N, 13306E). In the growth changes. Therefore, it is necessary to under- environment, the substratum was divided into stand the component species of the target sea- bedrock, large boulders (diameter 100 cm <), weed beds as well as their growth and repro- boulders (diameter 30-100 cm), cobbles (diame- duction. The Oki Islands, of Shimane Prefec- ter 10-30 cm), pebbles (diameter 0.5-10 cm), and ture, are located in the Japan Sea and consist sand (diameter 0.5 cm >). Similar surveys were of four large islands (Nishinoshima Island, conducted at a depth of 15 m at Hobomi (365 Nakanoshima Island, Chiburishima Island, and 9N, 133812E) on July 3, 2005, and at a depth Okinoshima Island). Both E. stolonifera and E. of 15 m at Sazae-jima (36922N, 133149E) on kurome areknowntogrowinthewaterssur- July 1, 2005. The survey at Hobomi was con- rounding these islands (Hagiwara et al. 1970; ducted on a seabed slope in a 5 m×125 m area, Kajimura 1975; Kawashima 1993a). However, and the survey at Sazae-jima on a seabed slope there are local seaweed beds in this area that in an 8 m×75 m area. In each area, plant cover- are composed of ecklonian plants that are dif- age of sympatric seaweed was surveyed in a 5 ficult to identify. In this study, we developed m × 5 m square. The survey was repeated three criteria to distinguish Ecklonia species in sea- times, and the average was calculated. weed beds off the coast of Japan by compa- ring the habitat, morphology, and propagation Morphology of natural plants characteristics of E. stolonifera and E. kurome. Ten plants of 2-year-old or more were care- Furthermore, we discuss the relationship be- fully denuded using a spatula and removed 2 Identification of ecklonian kelps which have similarity to Ecklonia storonifera Okamura (Laminariales, Phaeophyta), in Nishinoshima coast of Oki Islands, Shimane Prefecture, Japan from each area. The distance between plants was more than 1 m. Subsequently, the samples were transported to the laboratory and the fol- lowing parameters were measured: (a) thallus length, (b) lamina length, (c) lamina width, (d) lamina thickness, (e) stipe length, (f) stipe di- ameter, (g) lateral length, (h) lateral width, (i) lateral number, (j) shoot number, (k) root (sto- lon) length, (l) root diameter, (m) holdfast size, (n) ratio of B to A, (o) ratio of C to A (Fig. 2). Fig. 2. Morphological parameters of ecklonian sporophyte in this study. Morphology of holdfast structure, stolon of Ecklonia stolonifera and root of Ecklonia kurome.(a) thallus length, (b) lamina length, (c) lamina width, (d) lamina thickness, (e) stipe length, (f) stipe diameter, (g) lateral length, (h) Fig. 1. Maps showing the locations of field investi- lateral width, (i) lateral numbers, (j) shoot gation and cultivation experiment (Dozen in numbers, (k) root (stolon) length, (l) root Map B) in Oki Islands (Map A), Shimane diameter, (m) holdfast size, (n) ratio of B to Prefecture. Map C is the magnification of A, (o) ratio of C to A. AK site in Map B. 3 Yuichi HAYASHI, Masahiro NOTOYA and Norishige YOTSUKURA Furthermore, (p) ratio of lamina length to lam- Tukey's-test and PCA (June 2007). ina width (b/c), (q) ratio of lamina length to stipe length (b/e), and (r) ratio of lateral length Analysis of rDNA ITS-1 sequences to lamina width (g/c) were calculated. The For the molecular characterization of plants, structure of the holdfasts was evaluated based a small piece of the blade (approximately 3 cm on the ratio of the diameter of the stolon and × 3 cm) of a sporophyte collected as shown in root to the diameter of the basal part of the the above-mentioned was cut and transported to branch point, according to the method of Arai the laboratory. Each specimen was washed in et al. (1997a). The statistical differences between sterile seawater and kept in silica gel prior to samples were assessed using analysis of vari- DNA sequencing. Genomic DNA was extracted ance (ANOVA), Tukey's-test, and principal and purified using the method described in components analysis (PCA), and pathognomonic Yotsukura et al.
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