Bull. Natl. Mus. Nat. Sci., Ser. B, 35(4), pp. 211–218, December 22, 2009

Distribution of Zostera Species in Japan. II
Zostera asiatica Miki, Z. caespitosa Mikiand Z. caulescens Miki (Zosteraceae)
Norio Tanaka1, *, Shoichi Akaike2, Mitsuhiro Ishii3, Ryoko Ishikawa4, Hiroshi Ito5,
Takahiro Kudo6, Daisuke Muraoka7, Akio Oshino8, Miho Saigusa9, Seiji Urabe10,
Kousuke Yatsuya11, Keizo Yoshimura12, Kenji Morita13 and Kenji Saitoh14
1

Department of Botany, National Museum of Nature and Science, Amakubo 4–1–1, Tsukuba, 305–0005 Japan
2
Hokkaido Hakodate Experiment Station, Yunokawa 1–2–66, Hakodate, Hokkaido, 042–0932 Japan
3
Chiba Prefectural Fisheries Research Center, Hiraiso 2492, Chikura-cho, Minami-Bousou, Chiba, 295–0024 Japan
4
Niigata Prefectural Fisheries and Marine Research Institute,
Ikarashi-sanno-cho 13098–8, Nishi-ku, Niigata, 950–2171 Japan
5
Hokkaido National Fisheries Research Institute, Coastal Fisheries and Aquaculture Division,
Fisheries Research Agency, Katsurakoi, Kushiro, 085–0802 Japan
6
Kanagawa Prefectural Fisheries Technology Center, Sea-farming Technique Division,
Jogashima, Misaki, Miura City, Kanagawa, 238–0237 Japan
7
Tohoku National Fisheries Research Institute, Coastal Fisheries and Aquaculture Division,
Fisheries Research Agency, Shinhama, Shiogama, 985–0001 Japan
8
Miyagi Prefecture Fisheries Technology Institute, Sodenohama 97–6, Ishinomaki, Miyagi, 986–2135 Japan
9
Agriculture, Forestry and Fisheries Policy Promotion Division of Agriculture, Forestry and Fisheries Department,
Miyagi Prefectural Government, Honcho 3–8–1, Aoba-ku, Sendai, Miyagi, 980–8570 Japan
10
Fisheries and Fishing Port Division, Toyama Prefectural Government,
Shinsogawa 1–7, Toyama, Toyama, 930–8501 Japan
11
Kyoto Institute of Oceanic and Fishery Science, Odashukuno, Miyazu, 626–0052 Japan(Present address: Coastal Fisheries and Aquaculture Division, Seikai National Fisheries Research Institute,Fisheries Research Agency, Taira 1551–8, Nagasaki, 851–2213 Japan)
12
Hokkaido Marine Culture Fisheries Experiment Station, Funami 1–156–3, Muroran, Hokkaido, 051–0013 Japan
13
Department of Research and Development, Tokyo Kyuei Co., Ltd.,
Shiba-tsurugamaru 6906–10, Kawaguchi, Saitama, 333–0866 Japan
14
National Research Institute of Fisheries Science, Aquatic Genomics Research Center,
Fisheries Research Agency, Fukuura, Kanazawa, 236–8648 Japan*Corresponding author: E-mail: ntanaka@kahaku.go.jp

Abstract The distribution of Zostera asiatica Miki, Z. caespitosa Miki and Z. caulescens Miki,
which are endemic to around Japanese archipelago, was assessed based on 396 herbarium specimens collected throughout Japan from 2004 to 2006. A total of 18 sites of Z. asiatica, 33 sites of
Z. caespitosa and 30 sites of Z. caulescens were recorded. Some new findings on distribution wereshown: Z. asiatica occurs in the western Hokkaido; Z. caespitosa occurs in Miyagi Prefecture as
southernmost population on the Pacific Ocean side; Z. caulescens occurs in Miyagi Prefecture andOki Islands, Shimane Prefecture.
Key words : Seagrass, Zostera asiatica, Zostera caespitosa, Zostera caulescens, Zosteraceae.

Introduction
The Japanese seagrass flora is noteworthy
since it contains several endemic species, namely

Zostera asiatica, Z. caespitosa, Z. caulescens, Z.
japonica, Phyllospadix iwatensis and P. japonicus (Zosteraceae), despite the fact that most seagrass species are widespread (Short et al., 2001).

212

Norio Tanaka et al.

However, as described by Tanaka et al. (2009),
reports on the distribution of seagrass species
around Japan are limited. Particularly the information about these endemic species is poor
(Miki, 1933, 1934; Omori, 1993, 1996; Environment Agency of Japan, 1994; Kudoh, 1999; Aioiet al., 2000; Shoji and Hasegawa, 2004; Akaike
and Goda, 2006; Tanaka et al., 2006; Yatsuya etal., 2007; Fujiwara et al., 2009; Ishikawa, 2009).
Based on a survey of over 6,000 herbarium
specimens collected from 2004 to 2006 in the research project being conducted by the Ministryof Agriculture, Forests and Fisheries and held in
the herbarium at the National Museum of Natureand Science, Japan (TNS), we have compiled
detailed locality data of all Zostera species inJapan. In this the second report resulting fromthis survey, we present distribution data for
Zostera asiatica, Z. caespitosa and Z. caulescens.
Materials and Methods
Surveys of Zostera species were conducted bymaking observations from the seashore, from aship, and by snorkeling and SCUBA diving excursions. Almost the entire Japanese coastline
was surveyed from May 2004 to December 2006.
Plants with several shoots connected by rhizomes
were collected at each population confirmed in
the surveys. Voucher specimens were made forall sites; these have been deposited in the herbarium of the National Museum of Nature and Science, Japan (TNS). Information on habitat parameters, including water depth, water temperature,transparency and bottom sediment, were also collected and recorded on the voucher specimenlabels.
Results and Discussion
A total of 18 sites of Zostera asiatica wereconfirmed only in Hokkaido Prefecture, based ona total of 148 herbarium specimens collected inthese surveys (Table 1, Fig. 1). The distributionof Z. asiatica has been known as the north of
Korean Peninsula, Japan (Hokkaido and Iwate

Prefectures), Sakhalin and the Kurile Islands(den Hartog, 1970, Aioi et al. 2000), however,
the western Hokkaido had been blank of distribution of the species (Omori, 2000). It became
clear for the first time in this survey that Z. asiatica inhabits in the western Hokkaido. Althoughsome of the specimens (No. 3, 4, 5, 7, 9, 10, 11,
13, 14, 15 in Table 1 and Fig.1) were collected as
drifting shoots (Akaike and Goda, 2006), it ispossible that some populations of Z. asiatica arepresent in their neighborhood besides confirmed
populations.
A total of 33 sites of Zostera caespitosa wereconfirmed in 9 prefectures, based on a total of
170 specimens (Table 1, Fig. 1). The distributionof Z. caespitosa has been reported as Korean
Peninsula, northeast China and Japan (Hokkaido
and north-central Honshu: Kyoto Prefecture on
the Sea of Japan side and Iwate Prefecture on the
Pacific Ocean side) (Omori, 2000). In this study,
it was revealed that many populations are occurred in Miyagi Prefecture. Particularly noteworthy of them is Samenoura Bay (No. 31 in
Table 1 and Fig. 1), which is southernmost population on the Pacific Ocean side. Also on the Sea
of Japan side where the density of Z. marina
populations is reported to be lower (Tanaka et al.,2009), some populations of Z. caespitosa are
confirmed.
A total of 30 sites of Zostera caulescens wereconfirmed in 5 prefectures, based on a total of 78
specimens (Table 1, Fig. 1). The distribution of
Z. caulescens has been known to be the south of
Korean Peninsula and Japan (north-central Honshu: Iwate, Chiba, Kanagawa, Niigata, Ishikawa
and Shimane Prefecture) (Omori, 2000). In thisstudy, it was revealed that many populations are
occurred in Miyagi Prefecture. Oki Islands (No.
81 in Table 1 and Fig. 1) is recorded to be southwesternmost population in the Sea of Japan for
the first time.
Z. asiatica, Z. caespitosa and Z. caulescens
were shown to be primarily occurred in the habitat similar to that of Z. marina: sheltered sites notopen coastline. The tendency is noticeable on the
Sea of Japan where the density of seagrass com-Samenoura Bay, Ojika, Ishinomaki, Miyagi Pref.

Katsuura Harbor, Tobi Is., Sakata, Yamagata Pref.

31

32

39°1032

38°2212

45°2544
45°2518
42°3031
40°5229
38°5139
38°4932
38°4721
38°4037
38°4034
38°4074
38°3841
38°3819

Zostera caespitosa Miki
19
Fujimi, Wakkanai, Hokkaido Pref.
20
Fujimi, Wakkanai, Hokkaido Pref.
21
Mouth of Usu Bay, Usu, Date, Hokkaido Pref.
22
Tofuga-ura, Noheji Bay, Noheji, Kitakami, Aomori Pref.
23
Iwatsuki Dainosawa, Kesennuma, Miyagi Pref.
24
Hajikami Akebo, Kesennuma, Miyagi Pref.
25
Tsuya Nagane, Motoyoshi, Motoyoshi, Miyagi Pref.
26
Arato Port, Minamisanriku, Motoyoshi, Miyagi Pref.
27
Fusubozaki, Minamisanriku, Motoyoshi, Miyagi Pref.
28
Omori, Minamisanriku, Motoyoshi, Miyagi Pref.
29
Kappazawa, Minamisanriku, Motoyoshi, Miyagi Pref.
30
Fujihama, Minamisanriku, Motoyoshi, Miyagi Pref.

Latitude45°0716
45°2420
45°2432
45°2643
45°2643
45°1331
45°0804
45°0650
45°0640
45°1058
45°1152
45°1221
44°5226
44°2003
44°1473
42°2918
42°2933
41°2744

Collection Site

139°3309

141°2928

141°3816
141°3804
140°4623
141°0835
141°3509
141°3536
141°3123
141°2920
141°2849
141°2734
141°2848
141°3015

142°2501
141°4543
141°4521
141°3837
141°3837
141°1719
141°1829
141°1719
141°1727
141°0819
141°0815
141°0813
141°4418
141°4045
141°3937
140°4810
140°4727
140°1454

Longitude

3.0

2.7

1.0
0.5
1.7
1.9
2.0
1.9
3.1
2.2

0.6
0.3
4.2

5.2
4.7
4.0

5.4

6.4

8.1

10.8
9.5

Water depth(m)

Herbarium specimens of Zostera species collected

Zostera asiatica Miki
1
Hama-tonbetsu, Hokkaido Pref.
2
Koetoioki, Wakkanai, Hokkaido Pref.
3
Koetoigyokounai, Wakkanai, Hokkaido Pref.
4
Esandomari, Wakkanai, Hokkaido Pref.
5
Keizanpaku, Wakkanai, Hokkaido Pref.
6
Esanuka, Sarufutsu, Hokkaido Pref.
7
Oniwakigyokounai, Rishirifuji, Hokkaido Pref.
8
Oniwakinumaura, Rishirifuji, Hokkaido Pref.
9
Oniwaki, Rishiri, Hokkaido Pref.
10
Izumi, Kutsugata, Rishiri, Hokkaido Pref.
11
Tanetomi, Kutsugata, Rishiri, Hokkaido Pref.
12
Funadomari Harbor, Rebun, Hokkaido Pref.
13
Teshiokounai, Tesio, Hokkaido Pref.
14
Okitsu, Tomamae, Hokkaido Pref.
15
Tomamae, Tomamae, Hokkaido Pref.
16
Cape Entomo, Minami-Usu, Date, Hokkaido Pref.
17
Minami-Usu, Date, Hokkaido Pref.
18
Fukushima, Hokkaido Pref.

No.*

Table 1.

2004. 6.15
2004.10.6
2005. 8.8
2006. 8.22
2004. 7.8
2005. 8.30
2005. 8.30
2006. 5.17
2006. 5.17
2006. 5.17
2006. 5.16
2006. 5.16
2006. 8.18
2006. 9.16
2006. 9.2

2005. 8.9
2006. 7.3
2006. 5.30
2005.11.9
2006.11.21
2005. 8.2
2006. 7.24
2006. 7.4
2004. 7.8
2006.11.21
2006.11.20
2005. 6.1
2006.11.17
2006.12.1
2004.10.29
2005. 8.8
2005. 8.8
2004. 6.24

Collection
Date9536754, 9541025
9535166, 9541023
9539404–9539406
9537267953504895350259535033
95336209533616
95336129533600
95335949536569–9536676
9536577–9536617
9535588–9535592

9539241–9539296
95333179533316
953676795333309539171–9539240
95333249533318–9533320
953517195333299533328
95367569533326953333195410279539407, 9539408
9539409–9539412
9539790

TNS accession No.**

Distribution of Zostera asiatica, Z. caespitosa and Z. caulescens
213

Collection Site

Uchiura, Awashimaura, Awashima Is. Niigata Pref.
Hiramatsu, Sado, Niigata Pref.
Shirose, Sado, Niigata Pref.
Futami, Sado, Niigata Pref.
Toyota, Sado, Niigata Pref.
Akadomari, Sado, Niigata Pref.
Nakanami, Himi, Toyama Pref.
Tagarasu, Obama, Fukui Pref.
Karubihama, Ine, Kyoto Pref.
Nagae, Miyazu, Kyoto Pref.
Nagae, Miyazu, Kyoto Perf.
Ejiri, Miyazu, Kyoto Pref.
Shimakage, Miyazu, Kyoto Pref.
Tai, Maizuru, Kyoto Pref.
Shimakage, Maizuru, Kyoto Pref.
Nariu, Maizuru, Kyoto Pref.
Nariu, Maizuru, Kyoto Pref.
Otsu Is. Maizuru, Kyoto Pref.
Takugi, Okinoshima, Oki, Shimane Pref.

No.*

33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51

Zostera caulescens Miki
52
Iwatsuki Senganda, Kesennuma, Miyagi Pref.
53
Iwatsuki Dainosawa, Kesennuma, Miyagi Pref.
54
Kodanohama Beach, Nagasaki, Kesennuma, Miyagi Pref.
55
Nagaisomori, Kesennuma, Miyagi Pref.
56
Nagaiso Shichihanzawa, Kesennuma, Miyagi Pref.
57
Hikado Port, Motoyoshi, Motoyoshi, Miyagi Pref.
58
Tanoura, Minamisanriku, Motoyoshi, Miyagi Pref.
59
Minato Port, Minamisanriku, Motoyoshi, Miyagi Pref.
60
Isatomae, Minamisanriku, Motoyoshi, Miyagi Pref.
61
Tatezaki, Minamisanriku, Motoyoshi, Miyagi Pref.
62
Yoriki, Minamisanriku, Motoyoshi, Miyagi Pref.
63
Hosoura, Minamisanriku, Motoyoshi, Miyagi Pref.
64
Shimizu, Minamisanriku, Motoyoshi, Miyagi Pref.
65
Fusubozaki, Minamisanriku, Motoyoshi, Miyagi Pref.
66
Hiraiso Port, Minamisanriku, Motoyoshi, Miyagi Pref.

—(Continued)—

Table 1.

38°5201
38°5131
38°5113
38°5050
38°5034
38°4832
38°4444
38°4435
38°4255
38°4230
38°4229
38°4140
38°4122
38°4034
38°4040

38°2734
38°1104
38°0823
37°5853
37°5632
37°5123
36°5700
35°3233
35°3930
35°3902
35°3815
35°3514
35°3521
35°3510
35°3435
35°3523
35°3515
35°2914
36°1010

Latitude141°3515
141°3512
141°3726
141°3549
141°3531
141°3322
141°3253
141°3212
141°3131
141°3208
141°3117
141°2957
141°2928
141°2849
141°2824

139°1518
138°2904
138°2719
138°1554
138°1935
138°2435
137°0224
135°4952
135°1717
135°1513
135°1504
135°1223
135°1502
135°1423
135°1532
135°2716
135°2707
135°2021
133°1448

Longitude

5.0
3.0
3.0
2.0
4.5
1.1
3.0
2.4
3.8
4.9
2.0
1.8
2.4
1.6–2.6
0.9

2.5–4.5
6.0–7.0
6.0
6.0
1.8

2.0
5.3
6.0
4.5

10.8
8.2
7.8
11.1
3.6
7.0
8.0

Water depth(m)2004. 7.8
2004. 7.8
2005. 1.6
2004. 7.8
2004. 7.8
2005. 8.30
2006. 8.23
2006. 8.23
2006. 8.23
2006. 8.23
2006. 8.23
2006. 5.17
2006. 5.17
2006. 5.17
2006. 5.17

2005. 7.22
2006. 8.3
2005. 8.18
2005. 7.13
2006.10.16
2006. 8.2
2005. 9.2
2005. 8.29
2006. 8.23
2005. 4.12
2006. 8.4
2006. 8.4
2006. 4.17
2006. 8.4
2006. 8.4
2006. 8.21
2006. 8.21
2006. 6.14
2005.10.8

CollectionDate9535050, 9535051
9535046, 9535047
9535053
953504395350429535030
95335809533583953358795335869533590953362595336229533617, 9533618
953361495400859535760
9540087
95400819535767, 9537656–9537690
9535755
9540765
9540088–9540105
9532149
9532064–9532066
953214495321399532076, 9532077
9532127
9532122953209195320889532113, 9532114
9540106–9540130

TNS accession No.**214
Norio Tanaka et al.

Collection Site

Mouth of Mizushiri River, Minamisanriku, Motoyoshi,
Miyagi Pref.
Hadenya, Minamisanriku, Motoyoshi, Miyagi Pref.
Oritate, Minamisanriku, Motoyoshi, Miyagi Pref.
Kappazawa, Minamisanriku, Motoyoshi, Miyagi Pref.
Nagashimizu, Minamisanriku, Motoyoshi, Miyagi Pref.
Samenoura Bay, Ojika, Ishinomaki, Miyagi Pref.
Oota, Sado, Niigata Pref.
Shintomi, Futtsu, Chiba Pref.
Futtsu, Futtsu, Chiba Pref.
Futtsu Tidal flat, Futtsu, Chiba Pref.
Tsuhama, Futtsu, Chiba Pref.
Chojagasaki, Hayama, Miura, Kanagawa Pref.
Sajima, Yokosuka, Kanagawa Pref.
Jogasaki, Miura, Kanagawa Pref.
Takugi, Okinoshima, Oki, Shimane Pref.

No.*

67
38°3852
38°3842
38°3841
38°3814
38°2216
35°5435
35°2018
35°1927
35°1916
35°1218
35°1515
35°1314
35°0815
36°0925

38°4019

Latitude*Numbers of sites are shown in Fig. 1.
**Accession numbers of National Museum of Nature and Science, Japan (TNS).

68
69
70
71
72
73
74
75
76
77
78
79
80
81

—(Continued)—

Table 1.

141°2845
141°2639
141°2848
141°3038
141°2908
138°2926
139°4964
139°4758
139°4735
139°5041
139°3440
139°3731
139°3719
133°1520

141°2641

Longitude

1.9
1.9
4.0
1.8
1.8
1.5
15.0

2.3
3.2
3.4
2.0
2.9
13.2

1.1

Water depth(m)2006. 5.16
2006. 5.16
2006. 5.16
2006. 5.16
2006. 8.18
2006. 8.2
2005. 6.27
2006. 6.26
2006. 6.26
2006.10.26
2004. 7.22
2004. 7.7
2004. 7.16
2005.10.20

2006. 5.17

CollectionDate9533601
95336079533598
95335939536539-9536568
9535756, 9535757
95410419534945-9534949
95410449534950-9534954, 9533566
9526486-9526489
9526520-9526523
95265629540016

9533610

TNS accession No.**

Distribution of Zostera asiatica, Z. caespitosa and Z. caulescens
215

216

Norio Tanaka et al.

Fig. 1. Distribtion of Zostera asiatica, Z. caespitosa and Z. caulescens in Japan. Numbers of the map correlate
to collection numbers in Table 1.

munities is lower. The few communities are Tobi
Island (No. 32 in Table 1 and Fig. 1), AwashimaIsland (33), Sado Island (34–38, 73), Toyama
Bay (39), Wakasa Bay (40–49) and Oki Islands(51, 81), in which Z. marina are also recorded(Tanaka et al. 2009).
Water depths of collection sites ranged from
4 m to 10.8 m in Z. asiatica, from 0.3 m to 11.1 m
in Z. caespitosa, and from 0.9 to 15 m in Z.
caulescens (Table 1). Z. caulescens denoted thesame tendency of the frequency distribution of Z.
marina, occurred primarily at depths of up to 3 m

and infrequently over 10 m (Tanaka et al., 2009).
In contrast, Z. asiatica and Z. caespitosa are
distributed with uniformity within each rangeof water depth. Z. asiatica was definitely foundonly at deep depth area. It was reported that Z.
asiatica produces thinner and larger leaves atdeeper bottoms with lower light condition, as insome shade-tolerant species in terrestrial plants(Watanabe et al., 2000). This species may adapt
to low light condition. The deep habitat of Z. asiatica may have disturbed discovery of their localities.

Distribution of Zostera asiatica, Z. caespitosa and Z. caulescens

217

hara, Hitoshi Iizumi, Masahiro Nakaoka, Toshinobu Terawaki, Hiroshi Sako, Kiyokazu Inoue,
Hirofuni Kawakami for conducting the research
project.
References

Fig. 2. Frequency distribution of water depths at
the Zostera asiatica, Z. caespitosa and Z.
caulescens collection sites.

The new findings on distributions of the three
species are important information for future
investigation about genetic structure, coveringwhole area of their distribution. It is highly possible that these endemic species and Z. marina
were differentiated in East Asia including Japanese Archipelago (Tanaka et al., 2003). Conducting studies on genetic structure, the occurrence
and diffusion of the species may be discussed inrelation with geological history, especially on
formation of the Sea of Japan.
Acknowledgements
The authors thank Yuriko Kawana, Yuki Ando,
Sho Nagata for mounting and organizing herbarium specimens, Keiko Aioi, Shogo Arai, Akihiro
Dazai, Masaaki Fukuda, Yumiko Kato, Nobuyasu Nakabayashi, Satoshi Ohata, Norihiko
Shoji, Ryuta Terada, Yozo Wada for sampling
plants, Masahiro Notoya, Yoko Satta, Shinji Kiri-

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Akaike, S. and Goda, H. 2006. Recent distribution of
Zosteraceae along the coast of northern Hokkaido.
Hokusuishi Dayori (73): 11–15 (in Japanese).
den Hartog, C. 1970. The sea-grasses of the world. NorthHolland, Amsterdam.
Environment Agency of Japan 1994. The report of themarine biotic environment survey in the 4th national
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Fujiwara, M., Yamaga, K., Kagawa, T. and Suenaga, Y.
2009. Horizontal distribution of seagrass and seasonalchange of the Zostera marina population off Shodoshima Island, Kagawa Prefecture, eastern Seto Inland
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Ishikawa, R. 2009. Distribution and flora of sea-grassbeds in coastal areas of Niigata Prefecture. Bulletin of
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Miki (Zosteraceae): Its distribution in Yamada Bay,
Iwate Prefecture, Japan. Otsuchi Marine Research Center Report (21): 32–37 (in Japanese).
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Shoji, Y. and Hasegawa, K. 2004. Distribution of eelgrass
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