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Journal of Applied Phycology 14: 343349, 2002. 2003 Kluwer Academic Publishers. Printed in the Netherlands.


Ecology of seaweed beds on two types of articial reef

Chang Geun Choi 1, Yashuo Takeuchi 2, Toshinobu Terawaki 3, Yukihiko Serisawa 4, Masao Ohno 5 and Chul Hyun Sohn 1,*
1 Department of Aquaculture, Pukyong National University, Pusan, 608-737, Korea; 2Shikoku Research Institute Inc., Yashima-nishimachi Takamatsu, Kagawa, 761-0192, Japan; 3National Research Institute of Fisheries and Environment of Inland Sea, Fisheries Agency, 2-17-5 Maruishi, Ohno, Saeki, Hiroshima, 739, Japan; 4Marine Biosystems Research Center, Chiba University, Amatu-kominato, Awa-gun, Chiba, 299-5502, Japan; 5Usa Marine Biological Institute, Kochi University, Inoshiri, Usa, Tosa, Kochi, 781-1164, Japan; *Author for correspondence (e-mail:; phone +82-51-620-6133; fax +82-51-628-7430)

Received 1 February 2002; accepted in revised form 13 May 2002

Key words: Articial reef, Community, Ecklonia karume, Enteromorpha intestinalis, Roughness, Sargassum, Settlement, Succession Abstract Articial reefs for the development of valuable shery resources were placed on sandy substrates at 8, 10 and 13 m depths in Muronohana, Ikata, Japan, and observed monthly or bimonthly from February 1999 to June 2001. Enteromorpha intestinalis and Colpomenia sinuosa were the primary dominants in spring. The number of seaweed species at all the sites gradually increased in winter. Each reef reached a climax stage of Sargassum spp., Ecklonia kurome and Padina arborescens within 18 months. Overall, 38 seaweed species were found during the study. The settlement of kelp, such as E. kurome, was promoted by reduced sand cover as a result of turbulence. Large-scale surface roughness could be important in maintaining the communities after initial establishment. Introduction Seaweeds provide a habitat and serve as spawningbeds, breeding and feeding grounds for coastal sh and shellsh, as well as other important sheries resources (Terawaki et al. 2001), which are commercially important in themselves (Watanuki and Yamamoto 1990). Seaweed beds of rocky bottoms are an important source of drifting plants that provide food and shelter for coastal marine life, including economically important sh (Largo and Ohno 1993). In order to achieve sustainable coastal development, seaweed forest formation has been emphasized as one method to conserve a healthy coastal environment (Choi 2001). An M type articial reef has been constructed for the purpose of increasing abalone numbers. The RF type of articial reef has become a popular technique for the enhancement of spiny lobster populations. These reefs are thought to improve sheries by concentrating sh and also increasing the natural production of biological resources. Many ecological studies have been undertaken regarding the re-establishment of kelp forests, with special attention to the use of articial reefs to recover lost seaweed beds (Choi et al. 2000a,b). Seaweed bed creation has become an important aspect of conservation of the marine ecosystem and of coastal zone management (Terawaki et al. 2001). The aims of the present study were to describe successional and seasonal trends of seaweed components on concrete reefs placed at different depths at a site in southern Japan.

Materials and methods The study was conducted on articial reefs placed at depths of 8, 10 and 13 m in Muronahana, Ikata, Shikoku islands, Japan on 16 February 1999. Two reefs of different sizes were placed at depths of 8 (site A), 10 (site B) and 13 m (site C) (Figure 1). The M


Figure 1. Schematic representation and arrangements of M type (I) and RF type reefs (II) placed at the different sites. III, IV: Growth of seaweeds on articial reefs.

type articial reef placed at sites A and B had a volume of 3.47 m 3 (2.5 1.5 1.25 m). The RF type reef at sites A and C was 4.8 m 3 (2 2 1.2 m) (Figure 1I, 1II). A large number of perennial seaweeds, such as Sargassum horneri, S. macrocarpum and Ecklonia kurome grow on natural substrata around the study site. The succession and growth of various marine algae were observed by scuba diving monthly or bimonthly over a 15-month period from March 1999 to June 2001. Photographs were taken by digital video camera, video camera and 35 mm camera, including ones of seaweeds on adjacent natural substrata. Water temperature, salinity and transparency (Secchi disk method) were measured at the study site. Sa-

linity was measured with a digital salinometer (Model 3-G, Tsurumi Seiki, Yokohama).

Results Water temperature, salinity and transparency are presented in Figure 2. Water temperature ranged between 10.0 C to 28.3 C and salinity between 32.96 and 35.49 during the study period. Transparency ranged from 6 to 8 m at site A, to 10 m at site B and to 12 m at site C. Table 1 shows the monthly abundance of algae on the articial reefs over the study period. A total of 38 species of seaweed (6 Chlorophyta, 18 Phaeophyta,


Figure 2. Seasonal changes in environmental factors at the study sites.

14 Rhodophyta) were identied during the study period. Diatom colonization on all reefs achieved 100% cover within one month of placement. Within three months after placement, Enteromorpha intestinalis and Colpomenia sinuosa appeared in abundance on the reefs, although they did not grow on the rocks in the study area. Thalli of E. intestinalis and C. sinuosa decayed over the summer, at which time coralline red algal species started to grow on the reefs. From August to October 1999, coralline algae covered almost 100% of the reefs at the study site. Diatoms and coralline algae had greatly increased their colonization of the reefs from October to December 1999. Brown algae such as Sargassum spp. and Ecklonia kurome covered over 20% of the reef during the winter season. The number of species tended to increase on the reefs during the winter and spring seasons (February to June 2000). In February 2000, C. sinuosa re-appeared and covered 2060% of the articial reefs until June 2000. The dominant and subdominant species were E. kurome, Padina arborescens, Sargas-

sum spp. and Jania adhaerens from June 2000 to June 2001. Table 2 shows the cover of Sargassum spp. and Ecklonia kurome attached to the tops and sides of M type and RF type reefs during June to August 2001. In June, Sargassum spp. covered approximately 50 60% of the top surfaces. In June and August, the Sargassum community was most abundant on the top surface of M type and RF type reefs than the sides. Ecklonia kurome covered approximately 50% of the side part of the reefs in June and August. However, coverage of the top surface did not exceed 23% within the same reefs. The cover of E. kurome tended to be the same on the sides during June to August. Within 5 months of reef placement, green algae were more abundant at the shallow site (A) than at deeper sites (B and C; both the M and RF type reefs). Brown algae at the shallower site (A) appeared earlier and were more abundant than at the deeper sites. In contrast, the cover ratio of red algae did not in-

Table 1. Monthly abundance of algae on the articial reefs at the study site. Species 1999 M M 2000 F M 2001 M A

Diatom Enteromorpha interstinalis Enteromorpha sp. Ulva pertusa Cladophora sp. Codium fragile C. tenue Ectocarpus sp. Colpomenia bullosa C. sinuosa Hydroclathrus clathratus Cutleria multida Ecklonia kurome Dictyota dichotoma D. linearis D. patens Pachydictyon coriaceum Padina arborescens Spatoglossum pacicum Sargassum hemiphyllum S. horneri S. macrocarpum S. patens S. piluliferum Sargassum sp. Scinaia japonica S. latifrons Asparagopsis taxiformis Gelidium sp. Amphiroa dilatata A. zonata Jania adhaerens Jania sp. Prionitis sp. Hypnea japonica Gigartina sp. Lomentaria catenata Ceramium kondoi Coralline algae

: Dominant (more than 40% cover), : Subdominant (1040% cover),

: Rare (less than 10% cover)

crease as fast and to such an extent at site A when compared to sites B and C. Algal biomass on the reefs averaged 1698 g wet wt m 2 at site A and averaged 620.6 g wet wt m 2 and 583.1 g wet wt m 2 at sites B and C, respectively in June, (Figure 3). The biomass at site A was mostly due to Sargassum spp. and Col-

pomenia sinuosa. The Sargassum community biomass was 1214 g wet wt m 2 at site A, 203.7 g wet wt m 2 at site B, and 74.1 g wet wt m 2 at site C in June 2000. Sargassum spp. biomass decreased remarkably in July and August, i.e. 439.1 g wet wt m 2 to 226.9 g wet wt m 2 at site A. In contrast Padina arborescens gradually increased on the

Table 2. Cover of Sargassum spp. and Ecklonia kurome attached to surface and side parts of M type and RF type reefs at the study site during June to August 2001. (Unit: %) June Sargassum spp. M Surface Side 60 40 RF 50 20 Ecklonia kurome M 20 50 RF 13 44 August Sargassum spp. M 30 5 RF 23 13 Ecklonia kurome M 23 50 RF 15 50

Discussion The succession on the articial reefs described here shows a similar pattern to that for other articial reefs (Serisawa and Ohno 1995a,b; Ohno 1993). In the initial stage, diatoms colonized all reefs at 100% cover. Commercially valuable shellsh such as turban shell and abalone appeared on the articial reefs after one month of establishment and grazed on the diatom and seaweed communities. Yamada et al. (1992) reported colonization of small annuals and crustaceous algae on articial concrete reefs. In this study, E. intestinalis and C. sinuosa were found to be the primary colonists. Early colonization by these species might be due to the annual release cycle of their reproductive cells. Several species of the Melobesioideae dominated in summer and autumn when water temperatures were higher. During autumn and winter, when water temperatures declined, the cover of brown algae was comparatively higher. The course of succession was clearly affected by season and temperature. Seaweeds present on the articial reefs in winter matured in May or June (Ohno et al. 1990). The total number of species found on all the reefs during the entire study period was 38, gradually increasing from February 2000, with the highest number in winter. Serisawa et al. (1998) reported that within a year following placement, Sargassum spp. was dominant on all plates. In our studies Sargassum spp., E. kurome and P. arborescens dominated in late successional (climax) stages of placement. The stability of the substrate is very important for maintaining seaweed beds (Watanuki and Yamamoto 1990) and in this study all reefs resisted waves and currents. Grazers, such as sea urchins, did not colonize the sites during the study. Foster (1975) reported that Macrocystis pyrifera colonization was greatest near the upper horizontal

Figure 3. Mean biomass of seaweed attached to the different types of articial reef at the study sites during July to August 2000. Other: Other algae, C. sin: Colpomenia sinuosa, E. kur: Ecklonia kurome, P. arb: Padina arborescens, S. spp.: Sargassum spp.

articial reefs from 103.5 to 206.6 g wet wt m 2. The biomass of seaweed was found to decay and decrease signicantly on the articial reefs at all study areas during the summer season (July to August 2000). At the early stage (March to July 1999), E. intestinalis and C. sinuosa covered 30100% of the available surface. In June and July, the cover of E. intestinalis at sites A and B was nearly 100% (Table 3). The thalli of E. intestinalis disappeared in July at site C, whereas at sites A and B it only disappeared in September and August, respectively. Coralline algae dominated from August to December 1999. Sargassum spp. appeared on the reefs at site A after 6 months, it only appeared after 16 months at sites B and C. In June 2000, P. arborescens, E. kurome and J. adhaerens gradually covered approximately 460% of the reefs. At site C, the cover of J. adhaerens tended to be comparatively greater than sites A and B (about 36100% on the reefs from June to December 2000).

Table 3. Dominant algal species attached to the articial reefs at sites A, B and C.

edges of blocks and concrete prisms. He mentioned that this edge effect may result from a combination of increased spore and larval settlement as well as from enhanced growth of plants associated with turbulent eddies formed around such obstructions. For similar reasons, E. kurome colonization might also be facilitated. We observed little sand cover on the edges and sides of the reefs, as a result of turbulence. The above results suggest that if settlement of kelp species such as E. kurome, on articial reefs is desired, then large-scale roughness and/or kelp knobs

(Terawaki et al. 2001) could help. Finally, depth is critical for achieving maximum algal biomass and in Muronohana, articial reefs should be established at depths of less than 10 m.

Acknowledgements The authors thank Dr Alan T. Critchley, Department of Botany, University of the Witwatersrand, Johan-

349 nesburg, South Africa, for useful suggestions in the preparation of this manuscript.
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