Journal of Oceanography, Vol. 58, pp. 343 to 363, 2002
Review
Deep-Sea Chemoautosynthesis-Based Communities in
the Northwestern Pacific
SHIGEAKI K OJIMA*
Ocean Research Institute, University of Tokyo, 1-15-1 Minamidai, Nakano, Tokyo 164-8639, Japan
(Received 14 May 2001; in revised form 22 July 2001; accepted 24 July 2001)
I review chemoautosynthesis-based faunal communities which have been discovered
in the deep-sea areas in the Northwestern Pacific. Fauna of each community and further biological studies are summarized. Zoogeographical and evolutionary status of
communities in the Northwestern Pacific is discussed.
Keywords:
⋅ Northwestern
Pacific,
⋅ chemosynthetic
communities.
gressed. Unfortunately, not all of such results are globally
known, partly because not a few results were published
only in Japanese. This review summarizes information
on the various deep-sea chemosynthetic communities in
the Northwestern Pacific and recent achievements in the
studies of those communities made mainly by Japanese
deep-sea biologists.
1. Introduction
Since the first discovery at the Galápagos Spreading
Center in 1977, deep-sea chemoautosynthesis-based
faunal communities (chemosynthetic communities) established in dependence on the primary production by
chemoautosynthetic bacteria have been reported from
hydrothermal vent fields and cold seep areas in the world
(see Sibuet and Olu, 1998; Tunnicliffe et al., 1998; Van
Dover, 2000 for reviews). In the Northwestern Pacific,
many such communities have also been discovered from
sites with various geological settings (Figs. 1 and 2).
Tunnicliffe and Fowler (1996) pointed out the faunal
similarity between hydrothermal vent communities in the
Northwestern Pacific and those in the Manus, North Fiji
and Lau back-arc basins in the Southwestern Pacific. Generally speaking, communities around the Japanese Islands
in not only hydrothermal vent fields but also cold seep
areas are characterized by the dominance of clams of the
genus Calyptogena, mussels of the genus Bathymodiolus
and some groups of vestimentiferans. These animals house
chemoautotrophic and/or metanotrophic endosymbiontic
bacteria. In contrast, Calyptogena bivalves are absent in
the hydrothermal vent fields on the Ogasawara Islandarc and the Mariana Trough. In the vent fields in the
Mariana Trough, the niche of Calyptogena bivalves is
filled by the provannid gastropods of the genus
Alviniconcha.
With exhaustive surveys during the last two decades,
information on the chemosynthetic communities in the
Northwestern Pacific has increased and biological studies of such communities in the various fields have pro-
2.
Chemoautosynthetic Communities in Cold Seep
Areas
2.1 Sagami Bay
The first chemosynthetic community from Japanese
waters was discovered in 1984 in the cold seep area off
Hatsushima, Sagami Bay (34°57′–35°01′ N, 139°12–14′
E: 830–1,230 m depth) (Okutani and Egawa, 1985). Similar communities have been reported from other seep areas in Sagami Bay, namely, the Okinoyama Bank (34°58′
N, 139°31′ E: 1,180–1,450 m depth) and the Sagami Knoll
(35°06′ N, 139°20′ E: 1,390–1,460 m depth) (Hashimoto
et al., 1988; Hattori et al., 1993). Dominant benthic animals of these communities are Calyptogena clams,
Bathymodiolius mussels and vestimentiferans (Hashimoto
et al., 1989b). Although the existence of the communities was suggested at some other sites in Sagami Bay
(Tanaka and Hashimoto, 1991), no further survey has been
carried out.
A temperature anomaly of about 10°C warmer than
surrounding sea water was observed at the site off
Hatsushima, while no such phenomena were reported
from the Okinoyama Bank site (Naka et al., 1991).
Tsunogai et al. (1996) analyzed bottom water around
Calyptogena colonies at the site off Hatsushima and
showed that the cold seep fluid is a mixture of sea water,
pore water and land-derived groundwater with a tempera-
* E-mail address: [email protected]
Copyright © The Oceanographic Society of Japan.
343
Fig. 2. Distribution of chemosynthetic communities in the seep
areas off the central Japan. Only sites of which exact positions have been published are shown: a, the Sagami Knoll;
b, off Hatsushima; c, the Okinoyama Bank; d, off Toi;
e, the Kanesu-no-Se Bank; f, 1,200 m site off Omaezaki;
g, the Ryuyo Submarine Canyon; h, the Tokai Thrust; i, the
Daini Tenryu Knoll; j, the Yukie Ridge; k, the Daisan Tenryu
Submarine Canyon; l, the Tenryu Submarine Canyon;
m, the Zenisu Ridge; n, the Muroto Knoll; o, off the Muroto
Point; p, the Ashizuri Knoll; q, the Daiichi Minami Muroto
Knoll.
Fig. 1. Distribution of sites where chemosynthetic communities were reported, in the Northwestern Pacific. Open and
closed circles denote seep areas and hydrothermal vent
fields, respectively. Sites within the squire are shown in
Fig. 2. Only sites of which exact positions have been published are shown: A, the Okushiri Ridge; B, the Hiroo Submarine Canyon; C, the Kurile Trench; D, the Japan Trench;
E, the Daiichi Kashima Seamount; F, Kagoshima Bay;
G, off Kikaijima; H, the Minami-Ensei Knoll; I, the North
Iheya Knoll; J, the Iheya Ridge; K, the Izena Hole; L, the
Hatoma Knoll; M, the Kuroshima Knoll; N, the Myojin
Knoll; O, the Sumisu Caldera; P, the Suiyo Seamount;
Q, the Mokuyo Seamount; R, the Kaikata Seamount; S, the
Nikko Seamount; T, the Daini Kasuga Seamount; U, the
Central Mariana Trough; V, the South Mariana Ridge;
W, the 13°N Ridge; X, the TOTO Caldera.
ture range of 34 to 54°C. Species diversity of the communities at the site off Hatsushima is much greater than
that at the Okinoyama Bank site (Fujikura et al., 1995).
Cluster analysis allowed Fujikura et al. (1995) to conclude that fauna of the site off Hatsushima are more similar to those of the hydrothermal vent field on the Minami
Ensei Knoll, the Okinawa Trough, than to the Okinoyama
Bank site. These results may be explained by differences
in the origin of cold seep fluid and/or water temperature
between the two sites in Sagami Bay (Fujikura et al.,
1995).
Okutani (1957) described Calyptogena soyoae on the
basis of two dead shells collected from 750 m depth in
Sagami Bay by a deep-sea trawl. Molecular analysis re344
S. Kojima
vealed that two genetically distinct Calyptogena clams
inhabit the sites off Hatsushima and on the Okinoyama
Bank (Kojima et al., 1995). As there is a significant difference in shell shape between them, Kojima and Ohta
(1997a) described C. okutanii. While C. soyoae is endemic
to cold seep areas in Sagami Bay, C. okutanii also inbabits
a seep area in the Ryuyo Submarine Canyon, the Nankai
Trough, and hydrothermal areas in the Okinawa Trough.
The genetic differentiation of populations of C. okutanii
among the three sites was shown by analyses based on
nucleotide sequences of mitochondrial DNA and nuclear
DNA (Kojima et al., 1997a). No clear evidence has been
obtained to show the inhabitation of C. okutanii on the
Sagami Knoll, the most interior seep site in Sagami Bay,
from either molecular (unpublished data) or morphological (Ito and Hashimoto, 12th Meeting of Japanese Association of Benthology, 1998) analyses. In addition, it is
surprising that molecular phylogenetic analyses showed
that C. soyoae is more closely related to the Eastern Pacific species, C. kilmeri than to C. okutanii (Peek et al.,
1997; Kojima et al., 2000a).
Using an in situ pore water sampler, Hashimoto et
al. (1995b) measured hydrogen sulfide concentration in
bottom water and interstitial water in and around
Calyptogena colonies at the site off Hatsushima. The results indicated the concentrations from 0.05 to 0.6
mmol/kg are suitable for the habitat of Calyptogena
bivalves.
Since September 1993, a colony of Calyptogena
clams at the site off Hatsushima are being recorded by a
real time, long-term deep sea floor observatory (Momma
et al., 1993). During the first 1.5 years of this system’s
records, spawning behavior of Calyptogen bivalves was
observed on eleven occasions, when the ambient water
temperature increased more than 0.1°C (Fujiwara et al.,
1998a). In addition, Fujiwara et al. (1998a) succeeded in
inducing in situ spawning of clams by artificially increasing the ambient sea water temperature and concluded that
the increase of water temperature works as a cue to synchronize spawning.
Suzuki et al. (1989a, b) reported amino acid sequences of hemoglobin of a Calyptogena clam. Kim et
al. (1995) determined the nucleotide sequence of 16S
rDNA from endosymbionts of Calyptogena bivalves and
showed that their symbionts are chemoautotrophic. Occurrence of bacteria in the primary oocytes of Calyptogena
clams was also reported by Endow and Ohta (1990), but
Tsukahara (1994) reported the absence of endosymbionts
in the vitellogenic oocyte cytoplasm of Calyptogena individuals collected from the same site.
Phospholipids (Eguchi et al., 1994), trace metal concentrations (Masuzawa et al., 1988; Zhang and Ishii,
2000) and bioactive substance (Kamano et al., 2000) in
the tissues of Calyptogena bivalves have been reported.
Koike et al. (1988) measured the respiration rate of
Calyptogen bivalves. Koyama et al. (1999) succeeded in
the cultivation of mantle cells of a Calyptogena bivalve
for two weeks. Preliminary results of ecological studies
in parasitic polychaetes and copepods of Calyptogena
bivalves in Sagami Bay were summarized by Miura
(1988b).
Hashimoto and Okutani (1994) described three species of the genus Bathymodiolus, namely, B. platiformis,
B. japonicus and B. aduloides from Sagami Bay. These
three species are also collected from hydrothermal vent
fields in the Okinawa Trough. Growth rates of B.
platiformis and B. japonicus were estimated by the markand-recapture method (Yamamoto et al., 14th SHINKAI
Symposium, 1997).
Two undescribed vestimentiferan species inhabit seep
areas in Sagami Bay. The first species is relatively large
species in the genus Lamellibrachia, which was distinguished as Lamellibrachia sp. L1 from another by molecular phylogenetic analysis based on nucleotide sequences of mitochondrial DNA (Kojima et al., 1997b).
Suzuki et al. (1988, 1989c) reported amino acid sequences
of hemoglobin of this species. As the tube of the second
species is very slender and spiral, and similar to that of
the type species of the genus Alaysia (Southward, 1991),
this species is tentatively called Alaysia? sp. A1 (Kojima
et al., 2000a). Vestimentiferans with such tubes inhabit
seep areas in the Nankai Trough and hydrothermal vent
fields in the Okinawa Trough. As morphological exami-
nation is necessary to assess whether they belong to the
genus Alaysis, they are referred to as Alyasia? spp. in this
review.
The existence of sulfide-oxidizing bacteria and other
groups of endosymbiotic bacteria in trophosome of
Lamellibrachia sp. L1 was shown by molecular
(Naganuma et al., 1997; Kimura et al., 1999; Elsaied et
al., 2002) and morphological (Naganuma et al., 1997;
Barry et al., 2000) evidence. Barry et al. (2000) also reported the degradation of symbionts in some trophosome
regions of Lamellibrachia sp. L1. On the basis of the comparison of microflora between vestimentiferan
trophosome and that in the bottom sediment in the seep
area, Kimura et al. (1999) proposed a hypothesis that
endosymbionts of this species are squired from both the
ambient environment and by vertical transmission from
parents. Nucleotide sequences of the ribulose-1,5bisphosphate carboxylase/oxygenase (RuBisCO) largesubunit gene were determined for symbionts of
Lamellibrachia sp. L1 during a study of the phylogenetic
diversity of RuBisCO genes of deep-sea microorganisms
(Elsaied and Naganuma, 2001). Elemental distributions
in tubes of Lamellibrachia sp. L1 and Alaysia? sp. A1
from Sagami Bay as well as L. satsuma from Kagoshima
Bay were revealed by electron-probe X-ray microanalysis
(Naganuma et al., 1986).
Six gastropod (Okutani et al., 1992b, 1993b; Okutani
and Fujikura, 1992) and five polychaete species (Miura,
1988a; Miura and Laubier, 1990; Miura and Hashimoto,
1991a, 1996) were described from seep areas in Sagami
Bay. A detailed fauna list of the chemosynthetic communities in Sagami Bay is provided in Fujikura et al. (1996).
Sulfide-oxidizing bacteria-like symbionts were observed
in bacteriocytes of a solemyiid bivalve, Acharax sp. (Barry
et al., 2000). Endow et al. (1992) reported the ultrastructure of brachial cells of a turrid gastropod, Oenopota
sagamiana, collected near a Calyptogena colony.
Meiofauna and foraminiferal assemblages at the site off
Hatsushima was reported by Shirayama and Ohta (1990)
and Akimoto et al. (1990b), respectively. Benthopelagic
organisms distributing above the seep area at the site off
Hatsushima were also studied by Kikuchi et al. (1990)
and Lindsay et al. (1998).
Stable carbon and nitrogen isotope ratios of a
Calyptogena bivalve and Lamellibrachia sp. L1 collected
in Sagami Bay as well as C. phaseoliformis from the Japan Trench were reported by Saino and Ohta (1989). Stable and radiocarbon isotope ratios and stable oxygen isotope ratio of Calyptogena shells were reported by Yoshida
and Tsukahara (1997) and Hattori and Oba (1994).
Haemolymph sulfide binding properties were measured
for Bathymodiolus platiformis, a Calyptogena bivalve and
Lamellibrachia sp. L1 (Barry et al., 1997).
The composition of free-living bacteria in the bot-
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
345
tom sediment in the seep area on the Okinoyama Bank
was reported by Yoshida and Ishida (1989). On the basis
of the geochemical analyses of sediment samples inside
a colony of Calyptogena bivalves at the site off
Hatsushima, Masuzawa et al. (1992) suggested that sulfate
reduction and hydrogen sulfide production using methane are active within the sediments just beneath the
Calyptogena colony. The density and distribution of
sulfur-oxidizing bacteria in the sediment within the colony
were also estimated (Maki, 1994). Takasugi et al. (1998)
analyzed the microbial community in the seep area off
Hatsushima based on sediment fatty acids and compared
with those in the Nankai Trough.
2.2 Suruga Bay
To date, the chemosynthetic community in Suruga
Bay was reported only from the cold seep area off Toi,
the Izu Peninsula (34°55′ N, 138°39′ E: 1,490–1,500 m
depth), which was discovered during a test dive of the
submersible SHINKAI 2000 in 1992. The vesicomyid clam
collected from this site was described as a new species,
Calyptogena fausta (Okutani et al., 1993a). This species
was also collected from the Nankai Trough. No further
detailed survey has been curried out at this site. Tsunogai
et al. (1998) reported the methane-rich buoyant plumes
from the mouth of Suruga Bay, which suggests the existence of larger seep area there.
2.3 Nankai Trough
In the course of the French and Japanese Projects
KAIKO and KAIKO-NANKAI, surveys using the French
submersible Nautile were conducted in the Tenryu Submarine Canyon in 1985 and on the Yukie Ridge in 1989,
and chemosynthetic communities associated with cold
water seepages were discovered at both sites (Juniper and
Sibuet, 1987; Le Pichon et al., 1987, 1992; Ohta and
Laubier, 1987; Sibuet et al., 1988; Lallemand et al., 1992).
More than ten additional seep sites have been discovered
thereafter and surveyed by the Japanese submersibles
SHINKAI 2000, SHINKAI 6500 and the Japanese remote
operated vehicles (ROV) DOLPHIN-3K and KAIKO in
the Nankai Trough. Those sites can be divided into five
groups on the basis of the bathymetric zonation of the
primary components of the chemosynthetic communities
(Kojima and Ohta, 1997b; Fujikura et al., 2000).
The shallowest seep site in this area was discovered
on the Kanesu-no-Se Bank (34°17–18′ N, 138°15′ E: 290–
330 m depth) (Hashimoto et al., 1995c). The communities at this site were dominated by a lucinid bivalve,
Mesolinga soliditesta, which was reported only from this
site (Okutani and Hashimoto, 1997), and three
vestimentiferan species. Molecular analysis (Kojima et
al., 2001a) revealed the inhabitation of Lamellibrachia
satsuma, whose type locality is the hydrothermal vent sites
346
S. Kojima
between depths of 82 and 110 m in Kagoshima Bay (Miura
et al., 1997), at this site. Other species are Lamellibrachia
sp. L1 and Escarpia sp. E1 (Kojima et al., 1997b).
Colonies of vesicomyid clams were discovered by
surveys using a deep-tow TV system on the Muroto Knoll
(32°58′ N, 134°14′ E: 600 m depth) and the Ashizuri Knoll
(32°29′ N, 133°38′ E: 600 m depth) (Fujikura et al., 2000).
From dead shells, these clams were judged to be
Calyptogena solidissima, which had been described on
the basis of specimens from the hydrothermal vent field
at a corresponding depth on the Minami-Ensei Knoll, the
Okinawa Trough (Okutani et al., 1992a). Okabayashi et
al. (1986) collected vestimentiferans at a depth of about
700 m on the Ashizuri Knoll. Recently, C. solidissima
and Lamellibrachia sp. L1 ware collected from the Daini
(second) Tenryu Knoll (34°04′ N, 137°48′ E: 590–610 m
depth) (Kuramoto and Joshima, 1998; Fujikura et al.,
2000; unpublished data).
In the Ryuyo Submarine Canyon (34°12′ N, 137°46′
E: 1,090–1,100 m depth), two species of Calyptogena
bivalves and two vestimentiferan species, namely,
Lamellibrachia sp. L1 and Escarpia sp. E2, were collected
(Ashi et al., 1995; unpublished data). While a small
Calyptogena species was shown to be C. okutanii (Kojima
and Ohta, 1997b), the large one was described as a new
species, C. nankaiensis (Okutani et al., 1996). From seep
areas at the 1,200 m site on the southern slope of the
Omaezaki Spur (34°15′ N, 138°02′ E: 1,140–1,200 m
depth) (Ohta et al., 1995), Bathymodiolus aduloides,
Lamellibrachia sp. L1 and Escarpia sp. E2 (Kojima et
al., 1997b) were collected. Although dead shells of
vesicomyid clams were observed, no living individuals
were discovered at this site (Ohta et al., 1995). A fauna
list of this site was summarized by Ohta et al. (1995).
These two sites are thought to belong to the same
bathymetric zone as the sites in Sagami Bay (Kojima and
Ohta, 1997b).
From the seep area on the Yukie Ridge (33°50′ N,
137°54–55′ E: 1,940–2,180 m depth), C. fausta, C.
similaris and vestimentiferans of the genus
Lamellibrachia were collected. These vestimentiferans are
genetically distinct from congeneric individuals from
Sagami Bay and shallower sites in the Nankai Trough and
the Okinawa Trough (Lamellibrachia sp. L1) and tentatively thought to be another species, Lamellibrachia sp.
L2 (Kojima et al., 1997b, 2001a). In 1999, Alaysia-like
vestimentiferans were collected from this site. In contrast
to the case of the genus Lamellibrachia, our preliminary
molecular analysis showed that they and those in Sagami
Bay are more closely related to each other than to individuals from the Okinawa Trough (Kojima et al., 18th
SINKAI Symposium, 2001). The Tokai Thrust site
(34°06–07′ N, 138°07–08′ E: 2,080 m depth) (Ashi, 1997)
is the type locality of C. similaris (Okutani et al., 1997).
Calyptogena similaris and an undescribed vesicomyid
clam were collected on the summit of a mud volcano in
the Kumano Basin at a depth of 1,910 m (Kuramoto et
al., 2001). In addition, lamellibrachiid vestimentiferans
and Calyptogena bivalves were collected around a depth
of 2,500 m off Kumano in 2001 (unpublished data). These
sites are thought to belong to the same bathymetric zone
as the site off Toi, Suruga Bay (Kojima and Ohta, 1997b).
Into the fifth bathymetrical zone, some sites in the
deepest region of the Nankai Trough are classified. The
seep area in the Tenryu Submarine Canyon (33°36–37′
N, 137°32′ E: 3,760–3,840 m depth) is the first place
where the chemosynthetic community was discovered in
the Nankai Trough. On the basis of specimens collected
from this site during the Nautile dives, three species of
the genus Calyptogena were described, namely, C.
nautilei, C. laubieri and C. kaikoi (Okutani and Métivier,
1986). Sulfur oxidizing metabolisms of endosymbiotic
bacteria of C. laubieri and C. kaikoi as well as C. similaris
from the Yukie Ridge and C. phaseoliformis from the Japan Trench, were revealed in a series of anatomical, histological, ultrastructural, physiological and isotopic studies carried out during the KAIKO and KAIKO-NANKAI
Projects (Boulègue et al., 1987; Fiala-Médioni and Le
Pennec, 1988, 1989; Fiala-Médioni et al., 1993). Many
chemosynthetic communities of similar fauna are distributed around the Daisan (third) Tenryu Submarine Canyon (33°39′ N, 137°55′ E: 3,760 m depth). This site is
the type locality of the recently described species
Calyptogena tsubasa (Okutani et al., 2000a). Nucleotide
sequences of two genes encoding hemoglobin were determined for C. kaikoi, collected at this site (Suzuki et
al., 1998). Li et al. (1999a) estimated microbial diversity
in the seep area at this site by amplifying 16S rDNA in
the bottom sediment using the polymerase chain reaction
(PCR) method. Calyptogena nautilei was also collected
near the real-time, long-term deep sea floor observatory
placed in a seep area on the Daiichi (first) Minami Muroto
Knoll (32°21′ N, 134°32′ E: 3,620 m depth) (Iwai and
Momma, 1997; Fujikura et al., 2000). Using the observatory, demersal fishes (Endo et al., 2000), crustaceans
(Iwasaki et al., 2001) and moulting of a galatheid crab
(Iwasaki et al., 2000) were observed. The chemosynthetic
communities were discovered in some seep areas off the
Muroto Point (32°35′ N, 134°41–42′ E: 3,210–3,270 m
depth) (Saito et al., 17th SHINKAI Symposium, 2001),
where Lamellibrachia sp. L2 (Kojima et al., 2001a), C.
nautilei and undescribed species of the genus Calyptogena
as well as dead shell of C. tsubasa were collected.
Lamellibrachiid vestimentiferans and Calyptogena
bivalves were collected between depths of 3,120 m and
3,240 m off Kumano. A bacterial mat and a colony of
Calyptogena bivalves were observed at depths of 3,840
and 3,870 m, respectively, off Kumano (Ashi et al., 18th
SHINKAI Symposium, 2001). Colonies of C. tsubasa and
an unidentified Calyptogena species were also discovered on the Zenisu Ridge, where the chomosynthetic community had been discovered during the KAIKO Project
(Le Pichon et al., 1987), around a depth of 3,300 m (33°33′
N, 138°26′ E) (Ohta et al., 17th SHINKAI Symposium,
2001).
In the Nankai Trough, various chemosynthetic communities are distributed within a relatively narrow area
and there may probably be many undiscovered seep sites,
from which endemic species of deep-sea reducing environments will be described. For example, some new seep
sites were discovered in 2000 (Kuramoto et al. and
Tanahashi et al., 17th SHINKAI Symposium, 2001). Information on the detailed distribution and evolutionary
process of dominant groups as well as environments of
their habitats will provide us with an opportunity to understand the mechanism by which the chemosynthetic
communities are organized. On the other hand, habitats
of some endemic species, especially that of Calyptogena
bivalves, might be limited in very small areas within the
specific depth ranges. All three Calyptogena species that
inhabit the hydrothermal vent fields in the Okinawa
Trough, namely, C. solidissima, C. okutanii and C.
nankaiensis, were reported from seep areas in the Nankai
Trough. The vent areas should be treated as their temporal habitats, because venting activity in the Okinawa
Trough did not start long before (Lu et al., 1981) and
each vent site is too unstable for species with low dispersal ability, such as Calyptogena bivalves. Under the more
stable circumstances in the seep areas in the subduction
zone off the Japanese Islands, the remarkably high species diversity of this group might have been accomplished.
If limited habitats are lost by the mining of methane hydrate in the deep-sea sediment, those species will easily
become extinct. To preserve the high biodiversity in the
chemosynthetic communities in the Nankai Trough, we
should give the entire picture of the geological distribution of the communities.
2.4 Japan and Kurile Trenches
During a series of dives of the submersible Nautile
in 1985, colonies of a very slender vesicomyid clam,
Calyptogena phaseoliformis (Métivier et al., 1986), were
discovered from the Daiichi (first) Kashima Seamount
(35°54′ N, 142°31′ E: 5,640–5,700 m depth), the upper
region of the Sanriku Escarpment, the Japan Trench
(40°07′ N, 144°10′ E: 5,650–6,000 m depth) and the
southernmost part of the Kurile Trench (41°18′ N, 144°45′
E: 5,100–5,800 m depth) (Cadet et al., 1987; Juniper and
Sibuet, 1987; Ohta and Laubier, 1987; Pautot et al., 1987;
Sibuet et al., 1988). The use of new systems with improved diving ability, namely, the submersible SHINKAI
6500 and the ROV KAIKO, has revealed the bathymetric
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
347
zonation of the chemosynthetic communities in the deepest part of the landward slope of the Japan Trench (Kojima
et al., 2000b; Fujikura et al., 2002).
The largest escarpment on the landward slope of the
Japan Trench is the Sanriku Escarpment, which is situated off Sanriku, northeastern Japan, and between depths
of 5,300 and 6,400 m. Colonies of C. phaseoliformis were
also discovered on the lower part of the Sanriku Escarpment (40°06–08′ N, 144°11′ E: 6,180–6,470 m depth)
(Fujioka and Murayama, 1992; Ogawa et al., 1996;
Fujikura et al., 2002). The nucleotide sequence of
mitochondral DNA of this species and that of 16S rDNA
from the endosymbiotic bacteria were reported in Peek et
al. (1997, 1998). No genetic divergence along the depth
gradient on the Sanriku Escarpment was detected for the
populations of C. phaseoliformis as well as between
populations in the Japan Trench and those in the Kruil
Trench (Kojima et al., 18th SHINKAI Symposium, 2001).
The recent detailed surveys revealed the inhabitation of the other bivalve species with chemoautotrophic
endosymbiontic bacteria, namely, C. fossajaponica
(Okutani et al., 2000a), Parathyasira kairei (Okutani et
al., 1999) and Acharax johnsoni (Okutani and Fujikura,
2002), on this escarpment. Fujikura et al. (2002) summarized molluscan fauna and local distribution of the chemosynthetic communities on the Sanriku Escarpment, and
suggested that the two species of the genus Calyptogena
segregated their habitats according to the grain size of
bottom sediment. Fujiwara et al. (2000a) characterized
endosymbiontic bacteria of C. fossajaponica.
Foraminiferans (Akimoto and Kato, 1999) and free-living bacteria (Li et al., 1999b; Kato et al., 2000) within
the colonies of C. phaseoliformis were reported. In 2001,
bacterial mats were discovered at a depth of 6,200 m in
the Kruil Trench (Kojima et al., 18th SHINKAI Symposium, 2001).
On the foot of the relatively small escarpment between depths of 6,600 and 6,800 m off Sanriku (40°06′
N, 144°13–14′ E), only colonies of C. fassajaponica were
reported (Kojima et al., 2000b). No further survey has
been curried out on this escarpment.
The deepest chemosynthetic community in the world
was reported from the seep area on the landward slope
near the axis of the Japan Trench (40°03–04′ N, 144°17′
E: 7,330–7,430 m depth) (Fujikura et al., 1999). The community is dominated by a thyasirid clam with
chemoautotrophic endosymbiontic bacteria, Maorithyas
hadalis (Okutani et al., 1999). Fujiwara et al. (2001) characterized endosymbiontic bacteria of this species and discovered two distinct endosymbiotic sulfur-oxidizing bacteria, which shows different distribution patterns in the
host cells. The microbial diversity within the colony of
M. hadalis and the isolation of two novel bacteria were
reported by Kato et al. (1999).
348
S. Kojima
From the seep area on the landward slope of the Japan Trench, four gastropod species were described
(Okutani and Fujikura, 2002). On the basis of specimens
collected from the mantle cavity of C. phaseoliformis,
Miura and Laubier (1989) described Nautiliniella
calyptogenicola as a representative of a new polychaete
family Nautiliniellidae (preoccupied generic name,
Nautilina, has been replaced in their second paper (Miura
and Laubier, 1990)).
On the landward slope of the Japan Trench as well
as the Kurile Trench, there are many escarpments, where
new seep areas are expected to be discovered. Evidently,
two Calyptogena clams were collected by a beam-trawl
during the cruise of the research vessel (R/V) TanseiMaru, Ocean Research Institute, University of Tokyo,
from about 1,730 m depth off Sanriku. Their symbiotic
bacteria were observed using a transmission electron microscope during a series of histological studies on invertebrates inhabiting vents and seeps by Kim and Ohta
(2000). Preliminary molecular analysis suggested that
they are an undescribed species (Kojima, 1996).
2.5 Japan Sea
Low biodiversity in the Japan Sea have been attributed to severe anoxic conditions during the middle period of the last glacial period. This holds good for the
chemosynthetic communities. The first community in the
Japan Sea was discovered in the seep area on the northwestern slope of the Okushiri Ridge (44°14′ N, 139°06–
07′ E: 3,110 m depth) in 1991 (Takeuchi et al., 1992).
White bacteria mats, small snails, shrimps, sea cucumbers and anemones were observed in the open cracks
(Naganuma et al., 1999). Naganuma et al. (1999) analyzed
fatty acids in the bottom sediments collected from this
site as well as the Shiribeshi Seamount, where neither
open cracks nor bacteria mats were observed, and showed
the microfloral similarity between these two sites and the
seep area in Sagami Bay. This result suggests that the
community on the Okushiri Ridge is methane-driven.
Bacteria mats were also discovered on the slope off Cape
Motsuta (Takeuchi, 2000) and south to southwest of the
Matsumae Plateau (Takeuchi et al., 2000).
2.6 Around Ryukyu Islands
During surveys using a deep-sea TV system in 1996,
many dead shells of Calyptogana solidissima were observed on the Kuroshima Knoll, off the Yaeyama Islands
(24 °07–08′ N, 124°12–13′ E: 680–810 m depth)
(Machiyama et al., 2001), and a chemosynthetic community dominated by bivalves Vesicomyia kuroshimaena,
which was described on the basis of specimens from this
site (Okutani et al., 2000a), Bathymodiolus mussels and
two vestimetiferan species, namely, Lamellibrachia sp.
L1 and Lamellibrachia sp. L5 (Kojima et al., 2001a), were
discovered during the surveys using ROV DOLPHIN-3K
in 1997 (Matsumoto et al., 1998). Lamellibrachia sp. L5
has been known only from this site and no specimen was
available for taxonomic studies, because a sole specimen
had been used for isotopic analysis. A living individual
of C. solidissima was collected during the dive in 2000
(Maki et al., 17th SHINKAI Symposium, 2001).
From the seep area off Kikaijima Island (27°57′–
28° 26′ N, 130°13–19′ E: 1,440–1,650 m depth)
(Tsukahara et al., 15th SHINKAI Symposium, 1998), C.
similaris and Escarpia sp. E2 was collected (Fujikura et
al., 2000; Kojima et al., 18th SHINKAI Symposium,
2001).
Only two specimens of Calyptogena bivalves, which
were heavily damaged, were recovered from a depth of
5,800 m in the Ryukyu Trench in 1998 (Kato et al., 16th
SHINKAI Symposium, 1999). Although our preliminary
molecular analysis suggested that they are an undescribed
species (unpublished data), no specimen is available for
morphological examination and taxonomic description.
2.7 Off Pacific coast of Hokkaido Island
During the cruise of the R/V Oshoro-Maru, Hokkaido
University in 1997, undescribed Calyptogena bivalves
were collected by a trawl at a depth of about 800 m off
Kushiro. By surveys using a deep-tow TV system, the
chemosynthetic community dominated by Calyptogena
bivalves was discovered in the Hiroo Submarine Canyon
(42°10′ N, 144°10–11′ E: 1,200 m depth) (Hashimoto et
al., 14th SHINKAI Symposium, 1997). Our preliminary
molecular analysis also suggested that collected
Calyptogena bivalves are undescribed. The description
of this species is now in progress (Hashimoto, personal
communication).
2.8 Sea of Okhotsk
In 1932, a living specimen and some dead shells of
Calyptogena bivalve were collected from the east off
northern Sakhalin at a depth of 1,634 m (Scarlato, 1981).
Although they were described as subspecies of C. soyoae
(Scarlato, 1981), revision of this subspecies is necessary
because their habitat is deeper than the distribution range
of C. soyoae. Dense aggregation of a thyasirid bivalve
Conchocera bisecta and a polychaete of the family
Ampharetidae were found in the methane-rich area on the
submarine slope of Paramushir Island, Kuril Islands
(50°31′ N, 155°19′ E: 750–800 m depth) during a joint
expedition of R/V Akademik Mstislav Keldysh and R/V
Vulkanolog in 1986 (Kuznetsov et al., 1989). After then,
colonies of C. bisecta, which contains sulfur- and methane-oxidizing endosymbiotic bacteria (Strizhov et al.,
1990; Kuznetsov et al., 1991), were discovered from many
methane-rich areas in the Sea of Okhotsk (Kamenev et
al., 2001).
2.9 Whale bones and artificial reduced environments
During a dive of the submersible SHINKAI 6500 in
1992, whale bones were discovered on the Torishima
Seamount, Ogasawara Island-arc, at a depth of 4,037 m
(30°55′ N, 141°50′ E) (Fujioka et al., 1993). Many benthic
animals such as mytilid mussel of the genus Idasola and
Munidopsis crabs were observed around the bones (for
fauna list, see Naganuma et al., 1996). Analysis of fatty
acid in the bottom sediments around the bones suggested
the occurrence of methane-oxidizing bacteria and sulfurreducing bacteria (Naganuma et al., 1996).
As an analogue of a dead fall of a large marine vertebrate, Kitazato and Shirayama (1996) placed cattle
bones on the bathyal seafloor in Sagami Bay and reported
the creation of a reduced environment around the bones
within one year. They considered that the artificial reduced environment was at an early stage of a chemosynthetic community as thick microbial mat grew on the
bones and a galatheid crab and a lithoid crab were observed to aggregate. Naganuma et al. (2001) also placed
pig heads near the seep area off Hatsushima, Sagami Bay
and observed that small Calyptogena colony was newly
formed around the pig head within one month after the
placement.
3.
Chemosynthetic Communities in the Hydrothermal Vent Fields
3.1 Okinawa Trough
The first hydrothermal vent field in the Japanese
waters was discovered on the Natsushima 84-1 Knoll, the
Mid-Okinawa Trough in 1984, where the temperature of
venting fluid is relatively low and no hydrothermal ventendemic species were reported (Kimura et al., 1988). To
date, chemosynthetic communities have been reported
from four hydrothermal vent sites in the Mid-Okinawa
Trough, namely, the Izena Hole site, the Minami-Ensei
Knoll site, the Iheya Ridge site and the North Iheya Knoll
site. Unfortunately, further surveys using the submersibles
are not permitted for the Izena Hole and the Minami-Ensei
Knoll sites due to unrecoverable mooring implements for
fisheries.
The hydrothermal vent field in the Izena Hole (27°16′
N, 127°04–05′ E: 1,310–1,580 m depth), was discovered
in 1988 (Kimura et al., 1989). Galatheid crabs, a lithodid
crabs, shrimps, vestimentiferans, Bathymodiolus
platiformis and bivalves of the genus Calyptogena were
reported from this site (Kimura et al., 1989; Tanaka et
al., 1990). Detailed biological surveys have not yet been
carried out at this site.
Fauna and the microdistribution of the chemosynthetic communities in the hydrothermal vent field on the
Minami-Ensei Knoll (28°24′ N, 127°38–39′ E: 640–710
m depth) were summarized by Hashimoto et al. (1995a).
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
349
A vesicomyid bivalve, C. solidissima (Okutani et al.,
1992a), mussels B. japonicus and B. aduloides, and
vestimentiferans inhabit this site. Although Hashimoto et
al. (1995a) reported two vestimentiferan species from this
site, no specimens have been available for molecular systematic analysis and their taxonomic positions remain
unknown. Coupling behavior of red crab was observed at
this site (Ohta and Kim, 1992). Kim and Ohta (2000) observed methanotroph-like endosymbionts in the
bacteriocytes of Bathymodiolus bivalves and that some
part of the bacteriocytes left the endothelium of the gill.
A strain of hyperthermophilic sulfur-reducing
archaebacterium was isolated from this site (Kwak et al.,
1995).
On the Iheya Ridge (27°33′ N, 126°59′ E: 1,400–
1,430 m depth), chemosynthetic communities dominated
by Calyptogena okutanii (Kojima and Ohta, 1997b),
Lamellibrachia sp. L1, Escarpia sp. E2 (Kojima et al.,
1997b) and Alaysia? sp. A2 (Kojima et al., 2000a) were
discovered in 1988 (Ohta and Kim, 2001). In 1995, a new
vent field was discovered at the North Iheya Knoll site
(27°47′ N, 126°54′ E: 1,050 m depth), 30 km north of the
Iheya Ridge site. From the North Iheya Knoll site, C.
okitanii, C. nankaiensis (Fujikura et al., 2000),
Lamellibrachia sp. L1, Alaysia? sp. A2, B. japonicus and
B. platifrons were reported as dominant species
(Yamamoto et al., 1999). The preliminary fauna list of
the chemosynthetic communities on the Iheya Ridge and
North Iheya Knoll sites was published by Yamamoto et
al. (1999). Meiofauna in the Iheya Ridge site was reported
by Shirayama (1992). Foraminiferans within the
Bathymodiolus colonies at this site were also reported by
Akimoto and Hattori (2000). Microbial floras in the North
Iheya Knoll as well as the Izu-Ogasawara Island-arc were
reported by Kobayashi et al. (1999). Chemoautotrophic
sulfur bacteria were isolated from organisms and ambient seawater of the chemosynthetic communities on the
Iheya Ridge (Naganuma et al., 1990, 1991). Two strains
of hyperthermophilic bacteria were isolated from the vent
site on the Iheya Ridge (Kobayashi and Horikoshi, 1992)
and one of them was described as a new archaea species
(Kobayashi et al., 1994). Two additional
hyperthermophilic archaea were isolated and described
from the Iheya Ridge (Gonzalez et al., 1998; Grote et al.,
1999).
In the last three years, some hydrothermal vent fields
have been discovered from the Southern Okinawa Trough.
On the Hatoma Knoll (24°51′ N, 123°50–51′ E: 1,470–
1,520 m depth), the communities dominated by a mussel
B. platiformis, a galatheid crab Shinkaia crosnieri and a
bresiliid shrimp Alvinocaris longirostrus were discovered
(Tsuchida et al., 2000). Calyptogena bivalves and
vestimentiferans have not yet been reported from this site.
Fauna lists of the communities and mid-water animals
350
S. Kojima
over this site were provided by Tsuchida et al. (2000).
Decapods observed around this site were reported in
Watabe and Miyake (2000). From the hydrothermal vent
fields north of the Miyakojima Island, the chemosynthetic
communities were reported (Matsumoto et al., 2001).
Around 1,500 m depth on the seamount north of the
Yonaguni-jima Island, colonies of “tube worms” were also
observed (Shinjo et al., 2001).
In 2000, a new hydrothermal vent field was discovered between depths of 1,300 and 1,400 m on the Daiyon
(fourth) Yonaguni Knoll in SOTP (Southern Part of
Okinawa Trough) area, which is the westernmost vent site
within the Japanese waters (Matsumoto et al., 2001).
Bathymodiolus mussels, bythograeid crabs and the new
tentative species of Alaysia? were collected at this site
(Fujikura et al., 2001). At a depth of about 1,200 m off
northeastern Taiwan, the hydrothermal vent-endemic
galatheid crab Shinkaia crosniera was collected by oceanographic dredge (Chan et al., 2000).
From the hydrothermal vent fields in the Okinawa
Trough, seven species and one subspecies of gastropods
(Okutani et al., 1993b, 2000b; Okutani and Fujiwara,
2000), four caridean shrimp species (Kikuchi and Ohta,
1995; Kikuchi and Hashimoto, 2000), one lithodid crab
(Takeda and Hashimoto, 1990), one galatheid crab (Baba
and Williams, 1998) and four nautiliniellid polychaete
species (Miura and Hashimoto, 1993, 1996) were described. Foraminiferal assemblages in the hydrothermal
vent fields in the Izena Hole and the Iheya Ridge were
reported by Akimoto et al. (1990a). Using plankton nets
attached to the submersible, benthopelagic organisms
were collected above the vent fields on the Iheya Ridge
(Kikuchi et al., 1990).
3.2 Kagoshima Bay
Before the discovery of a colony of vestimentiferans
at a depth of 82 m in Kagoshima Bay (31°39′ N, 130°48′
E) (Hashimoto et al., 1993), this group had been thought
to be endemic to the deep-sea. The dense colonies of
vestimentiferans, Lamellibrachia satsuma (Miura et al.,
1997) were observed at this site using a deep-tow TV system in 1993. The colonies are distributed between depths
of 82 and 110 m. This species can be kept in an aquarium
at normal pressure for a long time and used for developmental and physiological studies (for example, Miura et
al., 2000). Some individuals have been kept for more than
five years. In addition, living individuals are put on view
at the Kagoshima City Aquarium. Miyake et al. (2000)
recently succeeded in introducing a vestimentiferan into
a transparent vinyl tube. This methodology is useful for
studies in their behavior and growth.
Miura and Hashimoto (1996) described a single species of nautiliniellid polychaete from this site. From bottom sediments at 200 m in the Wakamiko Proto-caldera,
where submarine volcanic activities are observed but
vestimentiferans have not been discovered. Naganuma et
al. (1991) isolated chemoautotrophic sulfur bacteria and
Yamanaka et al. (1999) reported petroleum-like products
from this site.
3.3 Izu-Ogasawara Island-arc
The chemosynthetic communities were reported from
some active submarine volcanoes of the Izu-Ogasawara
Island-arc, namely, the Myojin Knoll, the Sumisu Caldera,
the Suiyo Seamount, the Mokuyo Seamount, the Kaikata
Seamount and the Nikko Seamount. Common features of
the chemosynthetic communities in this sea area and those
in the Mariana Trough are absence of Calyptogena
bivalves and rarity of vestimentiferans, which have been
reported from the Sumisu Caldera, the Mokuyo Seamount,
the Nikko Seamount and the southernmost site of the
Mariana Trough. Vestimentiferans dominate the chemosynthetic community only at the Nikko Seamount site.
Low species diversity of the communities on the IzuOgasawara Island-arc might be attributed to the relatively
short history of the hydrothermal activities of this sea area
(Hashimoto and Fujikura, 9th SHINKAI Symposium,
1992).
At the northernmost hydrothermal site in this sea
area, the Myojin Knoll (North Bayonnaise Submarine
Caldera; 32°06′ N, 139°52′ E: 1,290 m depth), the chemosynthetic communities were discovered in 1996 (Iizasa
et al., 1999). The communities are dominated by a mussel Bathymodiolus septemdierum, a bythograeid crab
Austinograea yunohana and garatheid crabs. Fujiwara et
al. (1998b) maintained an individual of B. septemdierum
for 8 months alive without sulfide and observed disappear of symbiotic bacteria from epithelial cells. Recently,
Fujiwara et al. (2000b) characterized endosymbiont bacteria of B. septemdierum from the Myojin Knoll as well
as B. platiformis and B. japonicus from the North Iheya
Knoll, the Okinawa Trough, on the basis of nucleotide
sequences of 16S rRNA genes and showed that symbionts
of B. septemdierum are thioautotrophs and those of other
two host species are methaonotrophs. Yamanaka et al.
(2000) analyzed the sulfur isotope compositions of soft
tissues of four species of Bathymodiolus mussels collected
from both vent and seep sites and showed that B.
platiformis and B. japonicus assimilate seawater sulfatesulfur, while B. septemdierum and B. aduloides incorporate sulfide-sulfur. The interspecific difference of sulfur
sources might be attributed to differences in kinds of
endosymbiotic bacteria. Sulfur isotope ratios were also
reported for a Bathymodiolus mussel from the Kaikata
Seamount as well as other vent-endemic animals of the
Okinawa Trough, by Kim et al. (1989). Mineral distributions in free-living bacteria, barnacles, alvinellid
polychaetes and bythograeid crabs were examined by elec-
tron probe X-ray microanalysis and electron microscopic
imaging (Futaesaku et al., 2000).
In the Sumisu Caldera (31°28′ N, 140°04′ E: 670–
690 m depth), Bathymodiolus bivalves, tube worms, which
looked like vestimentiferans, and gastropods were observed in 1998 (Iwabuchi, 1999).
The chemosynthetic communities on the Suiyo
Seamount (28°34–35′ N, 140°39′ E: 1,370 m depth),
which were discovered in 1990 (Kasuga and Kato, 1992),
were dominated by a mussel, B. septemdierum, a
bythograeid crab, A. yunohana, a garatheid crab,
Alvinocaris shrimps and scale worms (Hashimoto, 1992;
Hashimoto and Okutani, 1994). A mytilid mussel, B.
septemdierum was described on the basis of specimens
from this site and the Mokuyo Seamount site (Hashimoto
and Okutani, 1994). A new species of extremely thermophilic archaebacteria was isolated and described from
this site (Gonzalez et al., 1995). The characteristics of
this species were reported in Gonzalez et al. (1996). Takai
and Horikoshi (1999) reported genetic diversity of
archaebacteria in the vent fields of this site, the Myojin
Seamount, and the Iheya Ridge and the North Iheya Knoll,
the Okinawa Trough.
The hydrothermal vent field on the Mokuyo
Seamount (28°19′ N, 140°34′ E: 1,260 m depth) was also
discovered in 1990 (Nagaoka et al., 1992). Dominant
benthic organisms of the chemosynthetic communities at
this site are shrimps of the genera Alvinocaris and
Lebbeus, Bathymodiolus septemdierum and
vestimentiferans (Hashimoto, 1992; Hashimoto and
Okutani, 1994).
The chemosynthetic communities on the Kaikata
Seamount (26°43′ N, 141°05′ E: 430 m depth), which was
discovered in 1989 (Mitsuzawa et al., 1989), were dominated by a bythograeid crab A. yunohana, Bathymodiolus
mussels and a tonguefish Symphurus sp. (see Ono et al.,
1996 for fauna list; Tsuchida et al., 2001).
The Nikko Seamount is situated in the southernmost
part of the Izu-Ogasawara Island-arc. Dense colonies of
vestimentiferans were observed on this seamount (23°05′
N, 142°20′ E: 430 m depth) (Hashimoto and Fujikura,
9th SHINKAI Symposium, 1992). The nucleotide sequence of mitochondrial DNA of a vestimentiferan collected from the Nikko Seamount was reported in Black et
al. (1997). By molecular phylogenetic analysis, it was
identified as Lamellibrachia satsuma (Kojima et al.,
2001a). Its symbiotic bacteria were characterized by Di
Meo et al. (2000). Other dominant animals of the communities are a tonguefish, a bythograeid crab and a primitive barnacle of Neolepas-type. A Bathymodiolus bivalve
was reported only from dead shells (Hashimoto and
Fujikura, 9th SHINKAI Symposium, 1992).
A bythograeid crab, A. yunohana (Takeda et al.,
2000) is distributed the Myojin Knoll Caldera, the Suiyo,
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
351
Kaikata and Nikko Seamounts. This species can be kept
in an aquarium at normal pressure for more than six years
and molting under laboratory condition has been observed
(Tsuchida et al., 1998). Fujikura et al. (1993) measured
the oxygen consumption rate of this species in an
aquarium and found almost same rates under in situ and
normal pressure conditions, as well as a higher thermal
tolerance than shallow water crustaceans. In addition,
Tanishima et al. (1996) showed that the locomotory activity pattern synchronized with the light-dark cycle of
this species in the aquarium. Such a pattern was lost by
the removal of eyestalks and this result suggests that this
species can use light for dial synchrony although it has
no compound eye and inhabits a low-light environment
between depths of 430 and 1,400 m (Tanishima et al.,
1996).
Two gastropod species (Okutani et al., 1993b) and
two polychaete species (Miura and Hashimoto, 1991b)
were described from the Izu-Ogasawara Island-arc.
3.4 Mariana Trough
The hydrothermal vent field was first discovered at
the Alice Springs Field site, the Central Mariana Trough
(18°13′ N, 144°43′ E: 3,600 m depth) during the dive of
the American submersible Alvin in 1987 (Craig et al.,
1987). The fauna of the chemosynthetic communities at
this site was summarized by Hessler and Lonsdale (1991),
Fujikura et al. (1997) and Hasegawa et al. (1997). The
second vent field was discovered at the Forecast Vent site,
the South Mariana Ridge (13°24′ N, 143°55′ E: 1,450 m
depth) (Johnson et al., 1993). A fauna list of the chemosynthetic communities at this site was provided by
Fujikura et al. (1997) and Hasegawa et al. (1997). In spite
of a large difference in water depth between two sites,
the faunas are quite similar to each other (Fujikura et al.,
1997).
The most dominant species of the communities in
hydrothermal vent fields in the Mariana Trough is a
provannid gastropod Alviniconcha hessleri (Okutani and
Ohta, 1988). Few genetic differentiations between the
Central Mariana Trough and the South Mariana Ridge
were revealed for populations of this species (Kojima et
al., 2001a). Endosymbiontic bacteria of this species were
reported in Stein et al. (1988) and Endow and Ohta
(1989a, b). Sulfur isotope ratios of this species as well as
a shrimp, Chorocaris vandoverae, were reported by Maki
and Mizota (1997). Other two gastropod species were described by Okutani (1990).
A bythograeid crab, Austinograea williamsi (Hessler
and Martin, 1989) is also dominant in the chemosynthetic
communities in the Mariana Trough. Tsuchida and
Fujikura (2000) reported heterochely, relative growth and
gonad morphology of this species.
On the basis of the nucleotide sequences of 16S
352
S. Kojima
rDNA and allozyme variations, Vrijenhoek et al. (1997)
analyzed the phylogenetic status of a limpet of the genus
Lepetodrilus from the Mariana Trough, which had been
thought to be conspecific to that in the Eastern Pacific,
and showed that species of the Mariana Trough is genetically distinct from East Pacific species.
From the summit of the Daini (second) Kasuga
Seamount, the Northern Mariana Trough (21°36′ N,
143°38′ E: 386 m depth), three vent-endemic animals,
namely, a bythograeid crab, a primitive barnacle and a
tonguefish were reported (Fujikura et al., 1998). Maki
(1998) reported microscopic structure of bacteria mats at
the vent sites on the Daini Kasuga Seamount as well as
those on the Suiyo Seamount.
In 1999, new vent sites were discovered in the
southernmost area of the Mariana Trough, namely, the
TOTO Caldera (12°43′ N, 143°32′ E) and the 13°N Ridge
(13°05′ N, 143°41′ E) (Mitsuzawa et al., 2000), where
the first vestimentiferans from the Mariana Trough were
discovered (Masuda et al., 2001).
4.
The Chemosynthetic Communities in the South
Pacific and the Indian Ocean, and Faunal Relationships to Those in the Northwestern Pacific
Using Japanese submersibles, the chemosynthetic
communities have been discovered and surveyed in the
other oceans than the Northwestern Pacific, such as the
hydrothermal vent fields in the Manus Basin (Hashimoto
et al., 1999) and the North Fiji Basin (Hashimoto et al.,
1989a; Desbruyères et al., 1994), the Southwestern Pacific and the vent site in the first segment of the Central
Indian Ridge, the Indian Ocean (Hashimoto et al., 2001).
During the sea bottom surveys using the ROV DOLPHIN
3K in the center of a M7.1 earthquake that occurred in
1998, the chemosynthetic community was discovered in
the seep areas off the Sissano Lagoon, northern coast of
the New Guinea Island (Matsumoto et al., 1999). At some
sites between depths of 600 and 2,200 m depth (02°48–
55′ S, 142°13–16′ E), Bathymodiolus bivalves, vestimentiferans and dead shells of white clams were observed
(Matsumoto et al., 17th SHINKAI Symposium, 2001).
Alviniconcha gastropods are one of the most dominant groups of the communities in all those vent sites as
well as vent sites in the Mariana Trough and the Lau Basin, the Southeastern Pacific (Desbruyères et al., 1994).
However, they have not yet reported from the seep areas
off the New Guinea Island. The sibling relationships between Alviniconcha gastropods of the Indian Ocean and
the Pacific species were shown by molecular phylogenetic
analyses (Hashimoto et al., 2nd International Symposium
on Deep-Sea Hydrothermal Vent Biology, 2001). On the
basis of molecular phylogenetic analyses, Kojima et al.
(2001b) concluded that Alviniconcha gastropods in the
Pacific should be classified into A. hessleri from the
Mariana Trough and at least two undescribed species from
the South Pacific, tentatively. Unfortunately neither a
specimen for molecular analysis nor DNA sequence data
were available for individuals of the Lau Basin.
Monotypic species of the closely related genus of
Alviniconcha, Ifremeria nautilei inhabits only vent sites
in the Manus Basin, the North Fiji Basin and the Lau
Basin. The relatively narrow distributional range of this
species might be attributed to its lower dispersal ability
than Alviniconcha spp. (Kojima et al., 2000c).
In the South Pacific, Calyptogena bivalves, which
have been not yet reported from the Izu-Ogasawara Island-arc and the Mariana Trough, were reported only from
the DESMOS Cauldron site, the Manus Basin (Hashimoto
et al., 1999) and the Edison Seamount, the New Ireland
Basin (Herzig et al., 1994). Those two sites are situated
off eastern and western coasts of the New Ireland Island,
Papua New Guinea. A close relationship between an
undescribed species from the Manus Basin and C.
similaris, which inhabits both the seep area in the Nankai
Trough and vent fields in the Okinawa Trough, was shown
by preliminary molecular analyses (Kojima et al., 2000a).
Although no information is available for a Calyptogena
bivalve of the New Ireland Basin, it may be the same species as that of the Manus Basin. In the Indian Ocean, only
dead shells of Calyptogena bivalves were observed and
collected.
A Lamellibrachia vestimentiferan collected from the
PACMANUS site, the Manus Basin, was shown to be
genetically distinct from those from the Northwestern
Pacific and L. column from the Lau Basin and called
Lamellibrachia sp. L4 (Kojima et al., 2001a). In addition, each of a lamellibrachiid of the DESMOS Basin and
the seep area off the New Guinea Island was suggested to
be endemic to its sampling site (Kojima et al., 18th
SHINKAI Symposium, 2001). On the other hand, our
preliminary molecular analysis suggested that Escarpia
vestimentiferans from the PACMANUS site and the site
off the New Guinea Island are Escarpia sp. E2. No genetic difference was detected for populations of Escarpia
sp. E2 between the Western Pacific and Bismarck Sea
(Kojima et al., 18th SHINKAI Symposium, 2001).
5. Discussion
During less than twenty years after the discovery of
the first chemosynthetic community in Japanese waters
in 1984, Japanese taxonomists have described many endemic species and subspecies of such communities in the
Northwestern Pacific (mentioned above) as well as the
South Pacific (Yamaguchi and Newman, 1990, 1997;
Okutani and Ohta, 1993; Von Cosel et al., 1994; Miura,
1994; Miura and Desbruyères, 1995; Newman and
Yamaguchi, 1995); Calyptogena bivalves and gastropods
by Dr. Okutani, emeritus professor of Tokyo University
of Fisheries and his collaborators, Bathymodiolus mussels by Dr. Hashimoto, Japan Marine Science and Technology Center, polychaetes by Professor Miura, Miyazaki
University, shrimps by Dr. Kikuchi, Yokohama National
University, barnacles by Professor Yamaguchi, Chiba University and others. A great portion of species of these
groups has already been described and I hope that remainder will be described soon. A Bathymodiolus bivalue of
the Indian Ocean was described by Hashimoto (2001).
Species of some other dominant groups from the Indian
Ocean are also now under description (Hashimoto, personal communication). On the other hand, taxonomic investigations of vestimentiferans, one of the most dominant groups of the chemosynthetic communities, in the
Northwestern Pacific as well as the Manus Basin, lag far
behind in other regions of the world. Only two species
(Miura et al., 1997; Southward and Galkin, 1997) of fourteen species suggested by the molecular analyses (Kojima
et al., 18th SHINKAI Symposium, 2001), have been described. As exact taxonomic information is indispensable for all biological studies, the earliest possible descriptions of remaining vestimentiferans and other minor
groups are urgently desired.
As mentioned above, fauna lists are available for
many vent and seep sites, though some lists are as yet
rather preliminary. On the basis of such information of
fauna, we can likely discuss the organization processes
of the chomosynthetic communities in the Northwestern
Pacific. On the basis of faunal similarity at the general
and familial levels, Tunnicliffe and Fowler (1996) pointed
out faunal similarity between hydrothermal vent communities in the Northwestern Pacific and those in the South
Pacific. Using the same methodology at the familial level,
Hashimoto et al. (2001) showed that the communities in
the Indian Ocean is more allied to those of both the Northwestern and the Southwestern Pacific than to other sea
areas. They thought that some groups might have colonized from the Indian Ocean into the Western Pacific
Ocean.
Information of phylogenies of dominant groups is
also useful for conjecturing the evolutionary histories of
the chemosynthetic communities. Recently, such information has been provided by the analyses based on DNA
sequences. Besides showing the derivation from annelids
(Kojima et al., 1993; McHugh, 1997; Kojima, 1998), the
molecular phylogenies of vestimentiferan species on the
basis of the mitochondrial DNA sequences, have been
reported (Kojima et al., 1997b, 2000a, 2001a; Black et
al., 1997). Recently, phylogenies of vent-endemic gastropods of the genera Alviniconcha and Ifremeria were
analyzed (Kojima et al., 2000c, 2001b). Molecular
phylogeny of Calyptogena bivalves in the Nothwestern
Pacific is still preliminary (Kojima et al., 1995, 2000a)
and it is difficult to assess the relationships between spe-
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
353
cies in the Western Pacific and those in the Eastern Pacific. Phylogenetic relationships among species of the
Northwestern Pacific as well as the South Pacific are now
analyzing for other dominant groups, Bathymodiolus
mussels (Kono et al., 18th SHINKAI Symposium, 2001)
and Austinograea crabs (Tsuchida et al., 18th SHINKAI
Symposium, 2001).
Vestimentiferans of each of the genera
Lamellibrachia and Escarpia inhabiting the Northwestern Pacific are thought to have two different origins
(Kojima et al., 2000a, 2001a). Species of each genus,
which inhabits sites shallower than 500 m are more closely
related to congeneric species of the Eastern Pacific than
to those of the deeper sites in the Northwestern Pacific.
Vestimentiferans, which can inhabit sites shallower than
500 m, have been known only from the Atlantic Ocean
and the Northwestern Pacific. The distribution of
methanotrophic symbionts of Bathymodiolus mussels are
also limited to the Atlantic and the Western Pacific
(Fujiwara et al., 2000b). Such patterns might be results
of the same evolutionary process.
On the other hand, Alaysia? spp. inhabiting the
Northwestern Pacific might have colonized into this sea
area along another route. The monophyletic group consisting of Alaysia? spp. and Arcovestia ivanovi from the
Manus Basin (Southward and Galkin, 1997) was shown
to have derived from vestimentiferans of the Northeastern Pacific by our preliminary analyses (Kojima et al.,
2000a).
While Alviniconcha gastropods inhabit the vent sites
not only in the South Pacific but also the Mariana Trough
as well as the Indian Ocean, gastropods of the closely
related genus Ifremeria have been known only from the
vent sites in the South Pacific. If differences in geographical distribution between these two genera are attributed
to the difference in dispersal ability (Kojima et al., 2000c),
Alviniconcha gastropods are thought to have dispersed
from the South Pacific to the Indian Ocean and the
Mariana Trough after the divergence from Ifremeria gastropods (Kojima et al., 1998).
The preliminary results of analyses on phylogenetic
relationships among Calyptogena species from both the
Northwestern and Northeastern Pacific oceans (Kojima
et al., 2000a) suggested that translocations between these
two regions have occurred many times. It has been suggested that although no genetic data are yet available,
species inhabit the Russian waters and the Aleutian Trench
are thought to be closely related to some Japanese species.
Molecular phylogenetic information for limited
groups has revealed that the chemosynthetic communities in the Northwestern Pacific have been constituted by
species with various origins. The dispersal abilities of
groups may be an important underlying factor. In order
354
S. Kojima
to understand the mechanisms by which such communities are organized, more accumulation of information of
more various groups is necessary.
The chemosynthetic communities established under
various geological settings in the Northwestern Pacific,
offer ideal subjects for studies in taxonomy, evolution and
ecology in the deep sea. Japanese deep-sea research systems are expected to bring great progress in studies of
these deep-sea chemosynthetic communities in the next
decade.
Acknowledgements
The author thanks Professor Y. Shirayama, Kyoto
University, for his kind invitation to prepare this review
article. Thanks are also extended to Drs. K. Fujikura and
J. Hashimoto, Japan Marine Science and Technology
Center, Dr. T. Okutani, emeritus professor of Tokyo University of Fisheries, and anonymous reviewers for useful
comments and information. Part of this work was supported by a grant from the Ministry of Education, Culture, Sports, Science and Technology, Japan (No.
12NP0201).
References
Akimoto, K. and M. Hattori (2000): Recent benthic
foraminiferal assemblage in the Bathymodiolus colony in
the northern Iheya Ridge, Okinawa Trough, Japan.
JAMSTEC J. Deep Sea Res., 16, 49–51 (in Japanese with
English abstract).
Akimoto, K. and C. Kato (1999): Recent benthic foraminiferal
assemblage in the Calyptogena colony on the landward slope
of the Japan Trench, Northwest Pacific Ocean. JAMSTEC
J. Deep Sea Res. I, 15, 5–9 (in Japanese with English abstract).
Akimoto, K., T. Tanaka, M. Hattori and H. Hotta (1990a): Recent benthic foraminiferal assemblages from the hydrothermal deposits in the Okinawa Trough, Ryukyu Islands, Japan. JAMSTEC Deepsea Res., 6, 99–114 (in Japanese with
English abstract).
Akimoto, K., T. Tanaka, M. Hattori and H. Hotta (1990b): Recent benthic foraminiferal assemblages around Calyptogena
colony off Hatsushima Island, Sagami Bay, Central Japan.
JAMSTEC Deepsea Res., 6, 205–215 (in Japanese with English abstract).
Ashi, J. (1997): Distribution of cold seepage at the fault scarp
of the eastern Nankai accretionary prism. JAMSTEC J. Deep
Sea Res., 13, 495–501.
Ashi, J., P. Huchon, J. Segawa, F. Thoué and K. Kobayashi
(1995): Geological structure and cold seepage at the prism
slope depression off Omaezaki: the 1994 KAIKO-TOKAI
“Shinkai 2000” dives. JAMSTEC J. Deep Sea Res., 11, 189–
195.
Baba, K. and A. B. Williams (1998): New Galatheoidea (Crustacea, Decapoda, Anomura) from hydrothermal systems in
the West Pacific Ocean: Bismark Archipelago and Okinawa
Trough. Zoosystema, 20, 143–156.
Barry, J. P., R. E. Kochevar, J. Hashimoto, Y. Fujiwara, K.
Fujikura and H. G. Greene (1997): Studies of the physiology of chemosynthetic fauna at cold seeps in Sagami Bay,
Japan. JAMSTEC J. Deep Sea Res., 13, 417–423.
Barry, J. P., K. R. Buck, S. K. Goffredi and J. Hashimoto (2000):
Ultrastructure studies of two chemosynthetic invertebratebacterial symbioses (Lamellibrachia sp. and Acharax sp.)
from the Hatsushima cold seeps in Sagami Bay, Japan.
JAMSTEC J. Deep Sea Res. I, 16, 91–99.
Black, M. B., K. M. Halanych, P. A. Y. Mass, W. R. Hoeh, J.
Hashimoto, D. Desbruyères, R. A. Lutz and R. C. Vrijenhoek
(1997): Molecular systematics of vestimentiferan
tubeworms from hydrothermal vents and cold-water seeps.
Mar. Biol., 130, 141–149.
Boulègue, J., E. L. Benedetti, D. Dron, A. Mariotti and R.
Létolle (1987): Geochemical and biogeochemical observations on the biological communities associated with fluid
venting in Nankai Trough and Japan Trench subduction
zones. Earth Planet. Sci. Lett., 83, 343–355.
Cadet, J. P., K. Kobayashi, S. Lallemand, L. Jolivet, J. Aubouin,
J. Boulègue, J. Dubois, H. Hotta, T. Ishii, K. Kinoshita, N.
Niitsuma and H. Shimamura (1987): Deep scientific dives
in the Japan and Kuril Trenches. Earth Planet. Sci. Lett.,
83, 313–328.
Chan, T., A. Lee and C. Lee (2000): The first deep-sea hydrothermal animal reported from Taiwan: Shinkaia crosnieri
Baba and Williams, 1998 (Crustacea: Decapoda:
Galatheidae). Bull. Mar. Sci., 67, 799–804.
Craig, H., Y. Hobibe, K. A. Farley, J. A. Welhan, K.-R. Kim
and R. N. Hey (1987): Hydrothermal vents in the Mariana
Trough: results of the first Alvin dives. EOS Trans. AGU,
68, 1531.
Desbruyères, D., A.-M. Alayse-Danet, S. Ohta and the Scientific Parties of BIOLAU and STARMER Cruises (1994):
Deep-sea hydrothermal communities in Southwestern Pacific back-arc basins (the North Fiji and Lau Basins): Composition, microdistribution and food web. Mar. Geol., 116,
227–242.
Di Meo, C. A., A. E. Wilbur, W. E. Holben, R. A. Feldman, R.
C. Vrijenhoek and S. C. Cary (2000): Genetic variation
among endosymbionts of widely distributed vestimentiferan
tubeworms. Appl. Environ. Microbiol., 66, 651–658.
Eguchi, E., M. Nakazawa, A. C. Suzuki and K. Nishimura
(1994): Comparative studies on phospholipids of deep-sea
and shallow-sea bivalves. JAMSTEC J. Deep Sea Res., 10,
481–490 (in Japanese with English abstract).
Elsaied, H. and T. Naganuma (2001): Phylogenetic diversity of
Ribulose-1,5-bisphosphate carboxykase/oxigenese largesubunit genes from deep-sea microorganisms. Appl.
Environ. Microbiol., 67, 1751–1765.
Elsaied, H., H. Kimura and T. Naganuma (2002): Molecular
characterization and endosymbiotic localization of the gene
encoding ribulose-1,5-bisphosphate carboxylase/oxygenase
(RuBisCO) form II in the deep-sea vestimentiferan
trophosome. Microbiology, UK (in press).
Endo, H., N. Iwasaki, Y. Machida and M. Iwai (2000):
Bathybenthic fauna of the Nankai Trough off Muroto observed using a deep seafloor observatory. I. Fishes.
JAMSTEC J. Deep Sea Res., 17, 13–17 (in Japanese with
English abstract).
Endow, K. and S. Ohta (1989a): Symbiotic gastropod from deep
sea hydrothermal vents. p. 28–32. In Recent Advances in
Microbial Ecology, ed. by T. Hattori, Y. Ishida, Y.
Maruyama, R. Y. Morita and A. Uchida, Japan Scientific
Societies Press, Tokyo.
Endow, K. and S. Ohta (1989b): The symbiotic relationship
between bacteria and a mesogastropod snail, Alviniconcha
hessleri, collected from hydrothermal vents of the Mariana
Back-Arc Basin. Bull. Jap. Soc. Microbial Ecol., 3, 73–82.
Endow, K. and S. Ohta (1990): Occurrence of bacteria in the
primary oocytes of vesicomyid clam Calyptogena soyoae.
Mar. Ecol. Prog. Ser., 64, 309–311.
Endow, K., J. Hashimoto, K. Fujikura and T. Uchida (1992):
Possible “talks” with environment: the branchial cells of a
deep-sea snail (Neogastropoda; Turridae) import “ligand(s)”
from the fluid of mantle cavity through coated pits. Proc.
JAMSTEC Symp. Deep Sea Res., 8, 327–333 (in Japanese
with English abstract).
Fiala-Médioni, A. and M. Le Pennec (1988): Structural adaptations in gill of the Japanese subduction zone bivalves
(Vesicomyidae) Calyptogena phaseoliformis and C.
laubieri. Oceanologica Acta, 11, 185–192.
Fiala-Médioni, A. and M. Le Pennec (1989): Adaptative features of the bivalve mollusks associated with fluid venting
in the subduction zones off Japan. Palaeogeogr.
Palaeoclimatol. Palaeoecol., 71, 161–167.
Fiala-Médioni, A., J. Boulègue, S. Ohta, H. Felbeck and A.
Mariotti (1993): Source of energy sustaining the
Calyptogena populations from deep trenches in subduction
zones off Japan. Deep-Sea Res. II, 40, 1241–1258.
Fujikura, K., J. Hashimoto, S. Segawa and Y. Fujiwara (1993):
Thermal tolerance of white blind crab (Bythograeidae) inhabited at hydrothermal vents. Proc. JAMSTEC Symp. Deep
Sea Res., 9, 383–391 (in Japanese with English abstract).
Fujikura, K., J. Hashimoto, Y. Fujiwara and T. Okutani (1995):
Community ecology of the chemosynthetic community at
Off Hatsushima site, Sagami Bay, Japan. JAMSTEC J. Deep
Sea Res., 11, 227–241 (in Japanese with English abstract).
Fujikura, K., J. Hashimoto, Y. Fujiwara and T. Okutani (1996):
Community ecology of the chemosynthetic community at
Off Hatsushima site, Sagami Bay, Japan—II: Comparisons
of faunal similaliy. JAMSTEC J. Deep Sea Res., 12, 133–
153 (in Japanese with English abstract).
Fujikura, K., T. Yamazaki, K. Hasegawa, U. Tsunogai, R. J.
Stein, H. Ueno, H. Yamamoto, Y. Maki, S. Tsuchida, T.
Kodera, H. Yamamoto, C.-H. Sun and T. Okutani (1997):
Biology and earth scientific investigation by the submersible “Shinkai 2000” system of deep-sea hydrothermalism
and lithosphore in the Mariana back-arc basin. JAMSTEC
J. Deep Sea Res., 13, 1–20.
Fujikura, K., S. Tsuchida, H. Ueno, J. Ishibashi, W. Gaze and
Y. Maki (1998): Investigation of the deep-sea chemosynthetic ecosystem and submarine volcano at the Kaguga 2
and 3 Seamounts in the Northern Mariana Trough, Western
Pacific. JAMSTEC J. Deep Sea Res., 14, 127–138.
Fujikura, K., S. Kojima, K. Tamaki, Y. Maki, J. Hunt and T.
Okutani (1999): The deepest chemosynthesis-based community yet discovered from the hadal zone, 7326 m depth,
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
355
in the Japan Trench. Mar. Ecol. Prog. Ser., 190, 17–26.
Fujikura, K., S. Kojima, Y. Fujiwara, J. Hashimoto and T.
Okutani (2000): New distribution records of vesicomyid
bivalves from deep-sea chemosynthesis-based communities
in Japanese waters. Venus (Jap. Jour. Malac.), 59, 103–121.
Fujikura, K., Y. Fujiwara, J. Ishibashi, S. Katayama, T. Komatsu,
Y. Maezawa, Y. Maki, J. Miyazaki, H. Miyake, K. Okamoto,
K. Okoshi, L. G. Toth, S. Tsuchida, T. Yamaguchi, T.
Yamanaka, H. Watabe, H. Watanabe, S. Zielinski and K.
Kato (2001): Report on investigation of hydrothermal vent
ecosystems by the crewed submersible ‘Shinkai 2000’ on
the Dai-yon (No. 4) Yonaguni Knoll and the Hatoma Knoll,
the Okinawa Trough. JAMSTEC J. Deep Sea Res., 19, 141–
154 (in Japanese with English abstract).
Fujikura, K., Y. Fujiwara, S. Kojima and T. Okutani (2002):
Micro-scale distribution of mollusks occurring in deep-sea
chemosynthesis-based communities in the Japan Trench.
Venus, 60, 225–236.
Fujioka, K. and M. Murayama (1992): Calyptogena community from landward slope of the Japan Trench and Mega
shear. Proc. JAMSTEC Symp. Deep Sea Res., 8, 17–27.
Fujioka, K., H. Wada and H. Okano (1993): Torishima whale
bone deep-sea animal community assemblage—new finding by Shinkai 6500. J. Geol., 102, 507–517.
Fujiwara, Y., J. Tsukahara, J. Hashimoto and K. Fujikura
(1998a): In situ spawnig of a deep-sea vesicomyid clam:
evidence for an environmental cue. Deep-Sea Res. I, 45,
1181–1189.
Fujiwara, Y., K. Uematsu, S. Tsuchida, T. Yamamoto, J.
Hashimoto, K. Fujikura, Y. Horii and M. Yuasa (1998b):
Nutritional biology of a deep-sea mussel from hydrothermal vents at the Myojin Knoll Caldera. JAMSTEC J. Deep
Sea Res., 14, 237–244 (in Japanese with English abstract).
Fujiwara, Y., S. Kojima, C. Mizota, Y. Maki and K. Fujikura
(2000a): Phylogenetic characterization of the
endosymbionts of the deepest-living vesicomyid clam,
Calyptogena fossajaponica, from the Japan Trench. Venus
(Jap. Jour. Malac.), 59, 307–316.
Fujiwara, Y., K. Takai, K. Uematsu, S. Tsuchida, J. C. Hunt
and J. Hashimoto (2000b): Phylogenetic characterization
of endosymbionts in three hydrothermal vent mussels: influence on host distributions. Mar. Ecol. Prog. Ser., 208,
147–155.
Fujiwara, Y., C. Kato, N. Masui, K. Fujikura and S. Kojima
(2001): Dual symbiosis in a cold seep thyasirid clam
Maorithyas hadalis from the hadal zone in the Japan Trench,
Western Pacific. Mar. Ecol. Prog. Ser., 214, 151–159.
Futaesaku, Y., Y. Kozuka, M. Ueno, M. Handa, T. Akagi, T.
Hashikawa, K. Uematu, Y. Fujiwara, S. Tsuchida, Y. Horii,
T. Yamaguchi and K. Iizasa (2000): Mineral distribution in
both chimney and organisms living in a deep sea on hydrothermal vents at Myojin Knoll Caldera. JAMSTEC J. Deep
Sea Res., 16, 53–67 (in Japanese with English abstract).
Gonzalez, J. M., C. Kato and K. Horikoshi (1995):
Thermococcus peptonophilus sp. nov., a fast-growing, extremely thermophilic archaebacterium isolated from deepsea hydrothermal vents. Arch. Microbiol., 164, 159–164.
Gonzalez, J. M., C. Kato and K. Horikoshi (1996): Culturability
and survival of an extreme thermophile isolated from deep-
356
S. Kojima
sea hydrothermal vents. Arch. Microbiol., 166, 64–67.
Gonzalez, J. M., F. T. Robb, J. W. Ammerman, D. L. Maeder,
M. Yanagibayashi, J. Tamaoka and C. Kato (1998):
Pyrococcus horikoshii sp. nov., a hyperthermophilic
archaeon isolated from a hydrothermal vent at the Okinawa
Trough. Extremophiles, 2, 123–130.
Grote, R., L. Li, J. Tamaoka, C. Kato, K. Horikoshi and G.
Antranikian (1999): Thermococcus siculi sp. nov., a novel
hyperthermophilic archaeon isolated from a deep-sea hydrothermal vent at the Mid-Okinawa Trough. Extremophiles,
3, 55–62.
Hasegawa, K., K. Fujikura and T. Okutani (1997): Gastropod
fauna associated with hydrothermal vents in the Mariana
Back-Arc Basin: Summary of the results of 1996 “Shinkai
6500” dives. JAMSTEC J. Deep Sea Res., 13, 69–83.
Hashimoto, J. (1992): Biological communities in the hydrothermal vent fields and cold seep areas around Japan. Kaiyo
Monthly, 24, 724–730 (in Japanese).
Hashimoto, J. (2001): A new species of Bathymodiolus
(Bivalvia: Mytilidae) from hydrothermal vent communities
in the Indian Ocean. Venus, 60, 141–149.
Hashimoto, J. and T. Okutani (1994): Four new mytilid mussels associated with deepsea chemosynthetic communities
around Japan. Venus (Jap. Jour. Malac.), 53, 61–83.
Hashimoto, J., S. Matsuzawa and H. Hotta (1988): Searching
for biological communities at the Okinoyama Bank site, the
Sagami Bay. Proc. JAMSTEC Symp. Deep Sea Res., 4, 177–
188 (in Japanese with English abstract).
Hashimoto, J., D. Jollivet and KAIYO 88 Shipboard Party
(1989a): The hydrothermal vent communities in the North
Fiji Basin: Result of Japan-France cooperative research on
board KAIYO 88. La Mer, 27, 62–71 (in Japanese with
English abstract).
Hashimoto, J., S. Ohta, T. Tanaka, H. Hattori, S. Matsuzawa
and H. Sakai (1989b): Deep-sea communities dominated by
the giant clam, Calyptogena soyoae, along the slope foot of
Hatsushima Island, Sagami Bay, central Japan. Palaeogeogr.
Palaeoclimatol. Palaeoecol., 71, 179–192.
Hashimoto, J., T. Miura, K. Fujikura and J. Ossaka (1993): Discovery of vestimentiferan tube-worms in the euphotic zone.
Zool. Sci., 10, 1063–1067.
Hashimoto, J., S. Ohta, K. Fujikura and T. Miura (1995a):
Microdistribution pattern and biogeography of the hydrothermal vent communities of the Minami-Ensei Knoll in the
Mid-Okinawa Trough, Western Pacific. Deep-Sea Res. I,
42, 577–598.
Hashimoto, J., S. Ohta, K. Fujikura, Y. Fujiwara and S. Sukizaki
(1995b): Life habit of vesicomyid clam, Calyptogena
soyoae, and hydrogen surfide concentration in interstitial
waters in Sagami Bay, Japan. J. Oceanogr., 51, 341–350.
Hashimoto, J., K. Fujikura, Y. Fujiwara, M. Tanishima, S. Ohta,
S. Kojima and S. Yieh (1995c): Observations of a deep-sea
biological community co-dominated by lucinid bivalve,
Lucinoma spectabilis (Yokoyama, 1920) and
vestimentiferans at the Kanesu-no-Se Bank, Enshu-Nada,
Centrl Japan. JAMSTEC J. Deep Sea Res., 11, 211–217 (in
Japanese with English abstract).
Hashimoto, J., S. Ohta, A. Fiala-Médione, J.-M. Auzende, S.
Kojima, M. Segonzac, Y. Fujiwara, J. C. Hunt, K. Gena, T.
Miura, T. Kikuchi, T. Yamaguchi, T. Toda, H. Chiba, S.
Tsuchida, J. Ishibashi, K. Henry, M. Zbinden, A. Pruski, A.
Inoue, H. Kobayashi, J.-L. Birrien, J. Naka, T. Yamanaka,
C. Laporte, K. Nishimura, C. Yeats, S. Malagun, P. Kia, M.
Oyaizu and T. Katayama (1999): Hydrothermal vent communities in the Manus Basin, Papua New Guinea: Results
of the BIOACCESS cruises ’96 and ’98. InterRidge News,
8(2), 12–18.
Hashimoto, J., S. Ohta, T. Gamo, H. Chiba, T. Yamaguchi, S.
Tsuchida, T. Okudaira, H. Watabe, T. Yamanaka and M.
Kitazawa (2001): First hydrothermal vent communities from
the Indian Ocean discovered. Zool. Sci., 18, 717–721.
Hattori, M. and T. Oba (1994): Oxygen and carbon stable isotope ratio of Calyptogena shells collected from Calyptogena
community associated with cold seeps off Hatsushima Island, Sagami Bay, Central Japan. JAMSTEC J. Deep Sea
Res., 10, 373–379 (in Japanese with English abstract).
Hattori, M., Y. Kanie, J. Hashimoto and K. Fujikura (1993):
Geological settings, mode of life and morphology of genus
Calyptogena along the subduction zone of the Sagami and
Suruga Trough, Central Japan. Proc. JAMSTEC Symp. Deep
Sea Res., 9, 237–251 (in Japanese with English abstract).
Herzig, P., M. Hannington, B. McInnes, P. Stoffers, H. Villinger,
R. Seifert, R. Binns and T. Liebe (1994): Submarine
volcanism and hydrothermal venting studies in Papua New
Guinea. EOS Trans. AGU, 75, 513–520.
Hessler, R. R. and P. F. Lonsdale (1991): Biogeography of
Mariana Trough hydrothermal vent communities. Deep-Sea
Res., 38, 185–199.
Hessler, R. R. and W. M. Martin (1989): Austinograea williamsi,
new genus, new species, a hydrothermal vent crab
(Decapoda: Bythograeidae) from the Mariana Back-Arc
Basin, western Pacific. J. Crust. Biol., 9, 645–661.
Iizasa, K., R. S. Fiske, O. Ishizuka, M. Yuasa, J. Hashimoto, J.
Ishibashi, J. Naka, Y. Fujiwara, A. Imai and S. Koyama
(1999): A Kuroko-type polymetalic sulfide deposit in a submarine silicic caldera. Science, 283, 975–977.
Iwabuchi, Y. (1999): Sumisu Caldera. JAMSTEC J. Deep Sea
Res., 15, 83–94 (in Japanese with English abstract).
Iwai, M. and H. Momma (1997): Deep-tow camera observation around the long-term deep sea floor observatory off
Muroto, Nankai Trough: Results from K96-09 cruise.
JAMSTEC J. Deep Sea Res., 13, 521–533 (in Japanese with
English abstract).
Iwasaki, N., T. Maeda and H. Momma (2000): Moulting of a
deep-sea galatheid crab (Anomura, Galatheidae) at a depth
of 3572 m. Hydrobiologia, 436, 237–239.
Iwasaki, N., M. Takeda, S. Nishida, M. Murano, C. Vallet and
M. Iwai (2001): Bathybenthic fauna of the Nankai Trough
off Muroto observed using a deep seafloor observatory 2.
Crustacea. JAMSTEC J. Deep Sea Res., 18, 217–224 (in
Japanese with English abstract).
Johnson, L., H. Fryer, H. Matsuda, T. Ishii and T. Gamo (1993):
Hydrothermal deposits and two magma sources for volcanoes near 13°20′ N in the Mariana backarc: a view from
Shinkai 6500. EOS Trans. AGU, 74, Fall Meet. Suppl., 381.
Juniper, S. K. and M. Sibuet (1987): Cold seep benthic communities in Japan subdustion zones: spatial organization,
trophic strategies and evidence for temporal evolution. Mar.
Ecol. Prog. Ser., 40, 115–126.
Kamano, Y., H. Hiraide, A. Yamashita, T. Watanabe, A.
Yamashita and J. Hashimoto (2000): Search for the chemical substances of deep-sea organisms Calyptogena soyoae.
JAMSTEC J. Deep Sea Res. I, 16, 1–6 (in Japanese with
English abstract).
Kamenev, G. M., V. A. Nadtochy and A. P. Kuznetsov (2001):
Conchocela bisecta (Conrad, 1849) (Bivalvia: Thyasiridae)
from cold-water methane-rich areas of the Sea of Okhotsk.
The Veliger, 44, 84–94.
Kasuga, S. and Y. Kato (1992): Discovery of hydrothermal ore
deposits in the crater of the Suiyo Smt. On the IzuOgasawara arc. Proc. JAMSTEC Symp. Deep Sea Res., 8,
249–256 (in Japanese with English abstract).
Kato, C., M. Yanagibayashi, Y. Nogi and K. Horikoshi (1999):
Analysis of microbial diversity in the sediment obtained
from Japan Trench at a depth of 7,326 m, and high pressure
cultivation. JAMSTEC J. Deep Sea Res., 15, 47–52 (in Japanese with English abstract).
Kato, C., Y. Nogi, T. Komatsu, T. I. Sato, T. Miura, E. Allen, F.
Lauro, S. Kurino and S. Noguchi (2000): Report of Japan
Trench microbial investigation cruises on YK99-05, leg. 4.
JAMSTEC J. Deep Sea Res., 17, 23–28 (in Japanese with
English abstract).
Kikuchi, T. and J. Hashimoto (2000): Two new caridean shrimps
of the family Alvinocarididae (Crustacea, Decapoda) from
a hydrothermal field at the Minami-Ensei Knoll in the MidOkinawa Trough, Japan. Species Diversity, 5, 135–148.
Kikuchi, T. and S. Ohta (1995): Two caridean shrimps of the
families Bresiliidae and Hippolytidae from a hydrothermal
field on the Iheya Ridge, off the Ryukyu Islands, Japan. J.
Crust. Biol., 15, 771–785.
Kikuchi, T., T. Toda, T. Nemoto and S. Ohta (1990): Preliminary survey on the deep-sea near bottom zooplankton by
means of deep-sea submersible “SHINKAI 2000”.
JAMSTEC Deepsea Res., 6, 115–122 (in Japanese with English abstract).
Kim, D. and S. Ohta (2000): TEM observation studies on the
chemoautotrophic symbiotic bacteria of invertebrates inhabiting at vents and seeps. Ocean Res., 22, 1–13.
Kim, E. S., H. Sakai, J. Hashimoto, F. Yanagisawa and S. Ohta
(1989): Sulfur isotopic ratios of hydrothermal vent-animals
at Ogasawara Arc and Mid-Okinawa Trough—evidence for
microbial origin of hydrogen sulfide at low-temperature
submarine hydrothermal areas. Geochem. J., 23, 195–208.
Kim, Y.-W., M. Yasuda, A. Yamagishi, T. Oshima and S. Ohta
(1995): Characterization of the endosymbiont of a deepsea bivalve, Calyptogena soyoae. Appl. Environ. Microbiol.,
61, 823–827.
Kimura, H., Y. Higashiide and T. Naganuma (1999):
Endosymbiotic and ambient microflorae of vestimentiferan
tube worm. JAMSTEC J. Deep Sea Res. I, 15, 25–33 (in
Japanese with English abstract).
Kimura, M., S. Uyeda, Y. Kato, T. Tanaka, M. Yamano, T. Gamo,
H. Sasaki, S. Kato, E. Izawa and T. Oomori (1988): Active
hydrothermal mounds in the Okinawa Trough backarc basin, Japan. Tectonophysics, 145, 319–324.
Kimura, M., T. Tanaka, M. Kyo, M. Ando, T. Oomori, E. Izawa
and I. Yoshikawa (1989): Study of topography, hydrother-
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
357
mal deposits and animal colonies in the Middle Okinawa
Trough hydrothermal areas using the submersible
“SHINKAI 2000” system. JAMSTEC Deepsea Res., 5, 223–
244 (in Japanese with English abstract).
Kitazato, H. and Shirayama, Y. (1996): Rapid creation of a reduced environment and an early stage of chemosynthetic
community on cattle bones at the deep-sea bottom in Sagami
Bay, Central Japan. An experimental approach to test the
stepping-stone hypothesis for chemosynthetic community
dispersion. Vie et Milieu, 46, 1–6.
Kobayashi, H., H. Takami, T. Nagahama, A. Inoue and K.
Horikoshi (1999): Microbial flora in deep-sea sites located
near the south part of Japan. JAMSTEC J. Deep Sea Res.,
15, 35–39 (in Japanese with English abstract).
Kobayashi, T. and K. Horikoshi (1992): Isolation and characterization of hyperthermophilic bacteria from a deep-sea hydrothermal vent system. Proc. JAMSTEC Symp. Deep Sea
Res., 8, 343–349 (in Japanese with English abstract).
Kobayashi, T., Y. S. Kwak, T. Akiba, T. Kudo and K. Horikoshi
(1994): Thermococcus profundus sp. nov., a new
hyperthermophilic archeon isolated from a deep-sea hydrothermal vent. Syst. Appl. Microbiol., 17, 232–236.
Koike, I., Y. Shirayama, T. Gamo and H. Sakai (1988): Respiration rate of Calyptogena soyoae obtained from the
Calyptogena communities at the Hatsushima site. JAMSTEC
Deepsea Res., 4, 233–237 (in Japanese with English abstract).
Kojima, S. (1996): Calyptogena bivalves around Japan. Kagaku,
66, 467–469 (in Japanese).
Kojima, S. (1998): Paraphyletic status of Polychaeta suggested
by phylogenetic analysis based on the amino acid sequences
of elongation factor-1α . Mol. Phylogenet. Evol., 9, 255–
261.
Kojima, S. and S. Ohta (1997a): Calyptogena okutanii n. sp.,
sibling species of Calyptogena soyoae Okutani, 1957
(Bivalvia: Vesicomyidae). Venus (Jap. Jour. Malac.), 56,
189–195.
Kojima, S. and S. Ohta (1997b): Bathymetrical distribution of
the species of the genus Calyptogena in the Nankai Trough,
Japan. Venus (Jap. Jour. Malac.), 56, 293–297.
Kojima, S., T. Hashimoto, M. Hasegawa, S. Murata, S. Ohta,
H. Seki and N. Okada (1993): Close phylogenetic relationship between Vestimentifera (tube worm) and Annelida revealed by the amino acid sequence of elongation factor-Iα .
J. Mol. Evol., 37, 66–70.
Kojima, S., R. Segawa, T. Kobayashi, T. Hashimoto, K.
Fujikura, J. Hashimoto and S. Ohta (1995): Phylogenetic
relationships among species of Calyptogena (Bivalvia:
Vesicomyidae) collected around Japan revealed by nucleotide sequences of mitochondrial genes. Mar. Biol., 122,
401–407.
Kojima, S., K. Fujikura, J. Hashimoto and S. Ohta (1997a):
Distribution and genetic structure of Calyptogena soyoae
complex. JAMSTEC J. Deep Sea Res., 13, 691–696 (in Japanese with English abstract).
Kojima, S., R. Segawa, J. Hashimoto and S. Ohta (1997b):
Molecular phylogeny of vestimentiferans collected around
Japan revealed by the nucleotide sequences of mitochondrial DNA. Mar. Biol., 127, 507–513.
358
S. Kojima
Kojima, S., S. Ohta, Y. Fujiwara, K. Fujikura and J. Hashimoto
(1998): Speciation of gastropods of the genus Alviniconcha
in the South Western Pacific. JAMSTEC J. Deep Sea Res.,
14, 501–505 (in Japanese with English abstract).
Kojima, S., S. Ohta, T. Miura, Y. Fujiwara and J. Hashimoto
(2000a):
Molecular
phylogenetic
study
of
chemoautosynthesis-based communities in the Manus Basin. JAMSTEC Deepsea Res., 16, 7–13 (in Japanese with
English abstract).
Kojima, S., T. Sasaki, K. Tamaki, Y. Fujiwara, K. Fujikura and
T. Okutani (2000b): Bathymetrical zonation of
chemoautosynthesis-based communities on the deepest area
of the landward slope of the Japan Trench. JAMSTEC Deep
Sea Res., 17, 89–93.
Kojima, S., R. Segawa, Y. Fujiwara, J. Hashimoto and S. Ohta
(2000c): Genetic differentiation of populations of a hydrothermal vent-endemic gastropod, Ifremeria nautilei, between the North Fiji Basin and the Manus Basin revealed
by nucleotide sequences of mitochondrial DNA. Zool. Sci.,
17, 1167–1174.
Kojima, S., S. Ohta, T. Yamamoto, T. Miura, Y. Fujiwara and J.
Hashimoto (2001a): Molecular taxonomy of
vestimentiferans of the western Pacific and their
phylogenetic relationship to species of the eastern Pacific.
I. Family Lamellibrachiidae. Mar. Biol., 139, 211–219.
Kojima, S., R. Segawa, Y. Fujiwara, K. Fujikura, S. Ohta and J.
Hashimoto (2001b): Phylogeny of hydrothermal vent-endemic gastropods Alviniconcha spp. from the Western Pacific revealed by mitochondrial DNA sequences. Biol. Bull.,
200, 298–304.
Koyama, S., T. Miwa and M. Aizawa (1999): First report of
primary culture for deep-sea clams. JAMSTEC J. Deep Sea
Res., 15, 41–46 (in Japanese with English abstract).
Kuramoto, S. and M. Joshima (1998): Precise gravity measurements for gas hydrate layer. JAMSTEC J. Deep Sea Res.,
14, 371–377 (in Japanese with English abstract).
Kuramoto, S., J. Ashi, J. Greinert, S. Gulick, T. Ishimura, S.
Morita, K. Nakamura, M. Okada, T. Okamoto, D. Rickert,
S. Saito, E. Suess, U. Tsunogai and T. Tomosugi (2001):
Surface observations of subduction related mud volcanoes
and large thrust sheets in the Nankai subduction margine;
Report on YK00-10 and YK01-04 Cruises. JAMSTEC J.
Deep Sea Res., 19, 131–139.
Kuznetsov, A. P., T. S. Rass and V. Galkin (1989): Benthic community of Paramushir gas hydrate source. Zoologicheskiy
Zhurnal, 58, 5–14 (in Russian with English abstract).
Kuznetsov, A. P., V. P. Strishov and S. I. Kijashko (1991): Respiratory and digestive systems in Conchocera sp. from the
Paramushir gas-hydrate vent (the Sea of Okhotsk) contained
by the bacterial biochemosynthesis. Izvestija Akademil Nauk
SSSR. Seria Biologicheskaya, 3, 445–457 (in Russian with
English abstract).
Kwak, Y. S., T. Kobayashi, T. Akiba, K. Horikoshi and Y. B.
Kim (1995): A hyperthermophilic sulfur-reducing
archaebacterium, Thermococcus sp. DT1331, isolated from
a deep-sea hydrothermal vent. Biosci. Biotech. Biochem.,
59, 1666–1669.
Lallemand, S. E., G. Glaçon, A. Lauriat-Rage, A. Fiala-Médioni,
J.-P. Cadet, C. Beck, M. Sibuet, J. T. Iiyama, H. Sakai and
A. Taira (1992): Seafloor manifestations of fluid seepage
at the top of a 2000-metre-deep ridge in the eastern Nankai
accretionary wedge: Long-lived venting and tectonic implications. Earth Planet. Sci. Lett., 109, 333–346.
Le Pichon, X., T. Iiyama, J. Boulègue, J. Charvet, M. Faure, K.
Kano, S. Lallemant, H. Okada, C. Rangin, A. Taira, T. Urabe
and S. Uyeda (1987): Nankai Trough and Zenisu Ridge: a
deep-sea submersible survey. Earth Planet. Sci. Lett., 83,
285–299.
Le Pichon, X., K. Kobayashi and Kaiko-Nankai Scientific Crew
(1992): Fluid venting activity within the eastern Nankai
Trough accretionary wedge: A summary of the 1989 KaikoNankai results. Earth Planet. Sci. Lett., 109, 303–318.
Li, L., J. Guenzennec, P. Nchols, P. Henry, M. Yanagibayashi
and C. Kato (1999a): Microbial diversity in Nankai Trough
sediments at a depth of 3,843 m. J. Oceanogr., 55, 635–
642.
Li, L., C. Kato and K. Horikoshi (1999b): Microbial diversity
in sediments collected form the deepest cold seep-area, the
Japan Trench at a depth of 6,400 m. J. Mar. Biotechnol., 1,
391–400.
Lindsay, D. J., J. C. Hunt, J. Hashimoto, K. Fujikura, Y.
Fujiwara, S. Tsuchida and K. Itoh (1998): The benthopelagic
community of Sagami Bay. JAMSTEC J. Deep Sea Res.,
14, 493–499.
Lu, R. S., J. J. Pan and T. C. Lee (1981): Heat flow in the southern Okinawa Trough. Earth Planet. Sci. Lett., 55, 299–310.
Machiyama, H., R. Shinjo, M. Hattori, M. Okano, T. Matsumoto,
M. Kimura, M. Nakamura, G. Toyama and T. Okada (2001):
Outline of Dolphin-3K dive surveys on the Kuroshima Knoll
and in the southern Okinawa Trough—Preliminary report
of the NT00-05 Cruise—. JAMSTEC J. Deep Sea Res., 18,
15–30 (in Japanese with English abstract).
Maki, Y. (1994): Numbers and distributions of sulfur-oxidizing bacteria in the sediments around the colony of clam
community at the bottom of the Sagami Bay. JAMSTEC J.
Deep Sea Res., 10, 389–393 (in Japanese with English abstract).
Maki, Y. (1998): Microscopic observations of microbial mats
at two hydrothermal vent areas: Suiyo Seamount and Kasuga
2 Seamount. JAMSTEC J. Deep Sea Res., 14, 593–607 (in
Japanese with English abstract).
Maki, Y. and C. Mizota (1997): Sulfur isotopic comparison of
Alviniconcha hessleri and Chorocaris vandoverae from
Mariana Back-Arc Basin. JAMSTEC J. Deep Sea Res., 13,
63–67 (in Japanese with English abstract).
Masuda, H., K. Mitsuzawa, N. Seama, S. Masumoto and YK99-11 Shipboard Scientific Party (2001): Bathymetry and
hydrothermal activities in the southern Mariana Trough—
reconnaissance results of YK-99-11—. JAMSTEC J. Deep
Sea Res., 18, 83–88 (in Japanese with English abstract).
Masuzawa, T., N. Handa and M. Koyama (1988): Trace element composition of Calyptogena soyoae from a giant clam
colony off Hatsushima Island, Sagami Bay, Japan.
JAMSTEC Deepsea Res., 4, 225–232 (in Japanese with English abstract).
Masuzawa, T., N. Handa, H. Kitagawa and M. Kusakabe (1992):
Sulfate reduction using methane in sediments beneath a
bathyal “cold seep” giant clam community off Hatsushima
Island, Sagami Bay, Japan. Earth Planet. Sci. Lett., 110,
39–50.
Matsumoto, T., M. Kimura, S. Nishida, T. Nakamura and T.
Ono (1998): Chemosynthetic communities and surface ruptures discovered on the Kuroshima Knoll south of Yaeyama
Islands (NT97-14 Cruise). JAMSTEC J. Deep Sea Res., 14,
477–491.
Matsumoto, T., D. R. Tappin, L. Anton, K. A. W. Crook, P. Hill,
Y. Iwabuchi, T. Kanamatsu, K. Kuratomi, Y. Lafoy, W. Lus,
Y. Matsumoto, M. Matsuyama, M. Hoihoi, Y. Murakami,
G. McMurtry, M. Nakamura, K. Satake, Y. Tsuji, J. P. Walsh,
II, P. Watts, H. Yeh, T. Kodera, M. Toizumi, H. Saitake, T.
Sugawara and K. Sagishima (1999): Precise topographic and
geological surveys off Sissana, Northern Papua New
Guinea—Preliminary results. JAMSTEC J. Deep Sea Res.
II, 15, 51–72.
Matsumoto, T., M. Kinoshita, M. Nakamura, J.-C. Sibuet, C.
Lee, S. Hsu, T. Oomori, R. Shinjo, Y. Hashimoto, S. Hosoya,
M. Imamura, M. Ito, K. Tukuda, H. Yagi, K. Takekawa, I.
Kagaya, S. Hokakubo, T. Okada and M. Kimura (2001):
Volcanic and hydrothermal activities and possible “segmentation” of the axial rifting in the westernmost part of the
Okinawa Trough—preliminary results from the
YOKOSUKA/SHINKAI 6500 Lequios Cruise—. JAMSTEC
J. Deep Sea Res., 19, 95–107 (in Japanese with English
abstract).
McHugh, D. (1997): Molecular evidence that echiurans and
pogonophorans are derived annelids. Proc. Natl. Acad. Sci.
USA, 94, 8006–8009.
Métivier, B., T. Okutani and S. Ohta (1986): Calyptogena
(Ectenagena) phaseoliformis n. sp., an usual vesicomyid
bivalve collected by the submersible Nautile from abyssal
depths of the Japan and Kurle Trenches. Venus (Jap. Jour.
Malac.), 45, 161–168.
Mitsuzawa, K., H. Momma, H. Hotta and Deep Sea Research
Group (1989): Measurements of water temperature and current around the submarine caldera in the KC Kaikata
Seamount. JAMSTEC Deepsea Res., 5, 47–55 (in Japanese
with English abstract).
Mitsuzawa, K., H. Matsuda, N. Seama, Y. Hasegawa, M.
Miyamoto, N. Togashi, A. So and H. Yamanobe (2000):
Preliminary report of Deep-Tow/Yokosuka Cruise at hydrothermal areas in the Mid and Southern Mariana. JAMSTEC
J. Deep Sea Res., 17, 73–87 (in Japanese with English abstract).
Miura, T. (1988a): A new species of the genus Protomystides
(Annelida, Polychaeta) associated with vestimentiferan
worm from the Hatsushima cold-seep site. Proc. Jap. Soc.
Syst. Zool., 38, 10–14.
Miura, T. (1988b): Parasitic animals collected in a Calyptogenadominant community developing off the Hatsushima, the
Sagami Bay. JAMSTEC Deepsea Res., 4, 239–244 (in Japanese with English abstract).
Miura, T. (1994): Two new scale-worms (Polynoidae:
Polychaeta) from the Lau Back-arc and North Fiji Basins,
South Pacific Ocean. Proc. Biol. Soc. Wash., 107, 532–543.
Miura, T. and D. Desbruyères (1995): Two new species of
Opisthotrachopodus
(Polychaeta:
Polynoidae:
Branchinotogluminae) from the Lau and the North Fiji Back-
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
359
arc Basins, southwestern Pacific Ocean. Proc. Biol. Soc.
Wash., 108, 583–595.
Miura, T. and J. Hashimoto (1991a): Nicomache otai, new species (Polychaeta: Maldanidae) collected from the
Hatsushima cold-seep in Sagami Bay. Proc. Biol. Soc.
Wash., 104, 159–165.
Miura, T. and J. Hashimoto (1991b): Two new branchiate scaleworms (Polynoidae: Polychaeta) from the hydrothermal vent
of the Okinawa Trough and the volcanic seamount off
Chichijima Island. Proc. Biol. Soc. Wash., 104, 166–174.
Miura, T. and J. Hashimoto (1993): Mytilidiphila, a new genus
of nautiliniellid polychaetes living in the mantle cavity of
deep-sea mytilid bivalve collected from the Okinawa
Trough. Zool. Sci., 10, 169–174.
Miura, T. and J. Hashimoto (1996): Nautiiliellid polychaetes
living in the mantle cavity of bivalve mollusks from cold
seeps and hydrothermal vents around Japan. Publ. Seto Mar.
Biol. Lab., 37, 257–274.
Miura, T. and L. Laubier (1989): Nautilina calyptogenicola, a
new genus and species of parasitic polychaete on a
vesicomyid bivalve from the Japan Trench, representative
of a new family Nautilidae. Zool. Sci., 6, 387–390.
Miura, T. and L. Laubier (1990): Nautiliniellid polychaetes
collected from the Hatsushima cold seep site in Sagami Bay,
with descriptions of new genera and species. Zool. Sci., 7,
319–325.
Miura, T., J. Tsukahara and J. Hashimoto (1997):
Lamellibrachia satsuma, a new species of vestimenitiferan
worms (Annelida: Pogonophora) from a shallow hydrothermal vent in Kagoshima Bay, Japan. Proc. Biol. Soc. Wash.,
110, 447–456.
Miura, T., J. Tsukahara and J. Hashimoto (2000): Larval development of a shallow-water vestimentiferan tube worm,
Lamellibrachia satsuma. Bull. Mar. Sci., 67, 669 (abstract).
Miyake, H., J. Hashimoto and S. Tsuchida (2000): Observation
method of behaviour of vestimentiferan tube-worm
(Lamellibrachia satsuma) in its tube. JAMSTEC J. Deep
Sea Res. I, 16, 35–39.
Momma, H., K. Mitsuzawa, Y. Kaiho and H. Hotta (1993):
“Hatsushima”—Real time long-term deep sea floor observatory. Proc. IEEE/Oceans ’93, 3, 473–477.
Naganuma, T. (1997): Morphological diversity of the microbial population in a vestimentiferan tubeworm. J. Mar.
Biotech., 5, 119–123.
Naganuma, T., M. Hattori, J. Hashimoto and Y. Kanie (1986):
Elemental distributions in the tubes of modern
vestimentiferan worms, and carbonate formation in their
habitats. Fossile, 60, 26–31 (in Japanese with English abstract).
Naganuma, T., E. Ikemoto, S. Sukizaki, Y. Tsuji and H. Hotta
(1990): Sulfur bacteria originating from the water and organisms in a hydrothermally active area of the Mid-Okinawa
Trough. J. Oceanogr. Soc. Japan, 46, 111–117.
Naganuma, T., Deep Sea Research Group and Deep Star Group
(1991): Collection of chemosynthetic sulfur bacteria from
a hydrothermal vent and submarine volcanic vents.
JAMSTEC J. Deep Sea Res., 13, 63–67.
Naganuma, T., H. Wada and K. Fujioka (1996): Biological community and sediment fatty acids associated with the deep-
360
S. Kojima
sea whale skeleton at Torishima Seamount. J. Oceanogr.,
52, 1–15.
Naganuma, T., C. Kato, H. Hirayama, N. Moriyama, J.
Hashimoto and K. Horikoshi (1997): Intracellular occurrence of ε -proteobacterial 16S rDNA sequences in the
vestimentiferan trophosome. J. Oceanogr., 53, 193–197.
Naganuma, T., C. J. Meisel, H. Wada, Y. Kato, A. Takeuchi, K.
Fujikura, J. Naka and K. Fujioka (1999): Sea-floor fissures,
biological communities and sediment fatty acids of the
Northern Okushiri Ridge, Japan Sea: Implications for possible methane seepage. The Island Arc, 8, 232–244.
Naganuma, T., M. Hattori, K. Akimoto, J. Hashimoto, H.
Momma and C. J. Meisel (2001): Apparent microfloral response to organic degradation on bathyal seafloor: an analysis based on sediment fatty acids. PSZN Mar. Ecol., 22, 267–
282.
Nagaoka, S., S. Kasuga and Y. Kato (1992): Geology of Mokuyo
Smt., Doyo Smt. and Suiyo Smt. In the Sitiyo Seamounts
on the Ogasawara arc. Proc. JAMSTEC Symp. Deep Sea
Res., 8, 237–248 (in Japanese with English abstract).
Naka, J., K. Fujikura and J. Hashimoto (1991): New geological
information of Calyptogena colony off Hatsushima, Sagami
Bay. JAMSTEC Deepsea Res., 7, 25–31 (in Japanese with
English abstract).
Newman, W. A. and T. Yamaguchi (1995): A new barnacle
(Cirripedia, Brachylepadomorpha) from the Lau Back-Arc
Basin, Tonga; first record of a living representative since
the Miocene. Bulletin du Muséum national d’Histoire
naturélle, Paris, 4e série, 17A, 221–243.
Ogawa, Y., K. Fujioka, K. Fujikura and Y. Iwabuchi (1996): En
echelon patterns of Calyptogena colonies in the Japan
Trench. Geology, 24, 807–810.
Ohta, S. and D. S. Kim (1992): Coupling behaviour of deepsea red crab and the nutritional ecology of vent mussel observed during the Shinkai 2000 Dive #542 surveying hydrothermal vent fields on the South Ensei Knoll, Okinawa
Trough, Japan. Proc. JAMSTEC Symp. Deep Sea Res., 8,
279–285 (in Japanese with English abstract).
Ohta, S. and D. Kim (2001): Submersible observations of the
hydrothermal vent communities on the Iheya Ridge, Mid
Okinawa Trough, Japan. J. Oceanogr., 57, 663–677.
Ohta, S. and L. Laubier (1987): Deep biological communities
in the subduction zone of Japan from bottom photographs
taken during “nautile” dives in the Kaiko project. Earth
Planet. Sci. Lett., 83, 329–342.
Ohta, S., S. Kojima, S. Yieh and J. Hashimoto (1995): Search
for the chemosynthetic ecosystem in the bathyal zone of
Enshunada—Report of the observations of the Dive 771 of
“Shinkai 2000”—. JAMSTEC J. Deep Sea Res., 11, 219–
225 (in Japanese with English abstract).
Okabayashi, Y., T. Tanaka and K. Nakamura (1986): Diving
survey of the knolls on the trench slope break off Kochi,
Southwest Japan. JAMSTEC Deepsea Res., 2, 173–192 (in
Japanese with English abstract).
Okutani, T. (1957): Two new species of bivalves from the deep
water in Sagami Bay collected by the R.V. Soyo-Maru. Bull.
Tokai. Reg. Fish. Res. Lab., 17, 27–30.
Okutani, T. (1990): Two new species of Provanna (Gastropoda:
Cerithiacea) from “snail pit” in the hydrothermal vent site
at the Mariana Back-Arc Basin. Venus (Jap. Jour. Malac.),
59, 83–101.
Okutani, T. and K. Egawa (1985): The first underwater observation on living habit and thanatocenoses of Calyptogena
soyoae in bathyal depth of Sagami Bay. Venus (Jap. Jour.
Malac.), 44, 285–289.
Okutani, T. and K. Fujikura (1992): A new turrid gastropod associated with metachromatic seabed near the Calyptogenacommunity in Sagami Bay. Venus (Jap. Jour. Malac.), 51,
1–7.
Okutani, T. and K. Fujikura (2002): Abyssal gastropods and
bivalves collected by SHINKAI 6500 on the slope of the
Japan Trench. Venus, 60, 211–224.
Okutani, T. and Y. Fujiwara (2000): Gastropod fauna of a thermal vent site on the North Knoll of Iheya Ridge, Okinawa
Trough. Venus (Jap. Jour. Malac.), 59, 123–128.
Okutani, T. and J. Hashimoto (1997): A new species of lucinid
bivalve (Heterodonta: Lucinidae) from Kanesu-no-Se Bank
near the mouth of Suruga Bay, with review of the recent
species of the chemosynthetic genus Lucinoma from Japan.
Venus (Jap. Jour. Malac.), 56, 271–280.
Okutani, T. and B. Métivier (1986): Descriptions of three species of vesicomyid bivalves collected by the submersible
Nautile from abyssal depths off Honshu, Japan. Venus (Jap.
Jour. Malac.), 45, 147–160.
Okutani, T. and S. Ohta (1988): A new gastropod mullusk associated with hydrothermal vents in the Mariana back-arc
basin, Western Pacific. Venus (Jap. Jour. Malac.), 47, 211–
218.
Okutani, T. and S. Ohta (1993): New buccinid and turrid gastropods from North Fiji and Lau Basins. Venus (Jap. Jour.
Malac.), 52, 217–221.
Okutani, T., J. Hashimoto and K. Fujikura (1992a): A new species of vesicomyid bivalve associated with hydrothermal
vents near Amami-Oshima Island, Japan. Venus (Jap. Jour.
Malac.), 51, 225–233.
Okutani, T., E. Tsuchida and K. Fujikura (1992b): Five bathyal
gastropods living within or near the Calyptogena community of the Hatsushima Islet, Sagami Bay. Venus (Jap. Jour.
Malac.), 51, 137–148.
Okutani, T., K. Fujikura and J. Hashimoto (1993a): Another
new species of Calyptogena (Bivalvia: Vesicomyidae) from
bathyal depth in Suruga Bay, Japan. Venus (Jap. Jour.
Malac.), 52, 121–126.
Okutani, T., K. Fujikura and T. Sasaki (1993b): New taxa and
new distribution records of deepsea gastropods collected
from or near the chemosynthetic communities in the Japanese waters. Bull. Nat. Sci. Mus. Ser. A, 19, 123–143.
Okutani, T., S. Kojima and J. Ashi (1996): An unusual
vesicomyid Bivalve, Calyptogena nankaiensis n. sp. from
bathyal depth off Honshu, Japan. Venus (Jap. Jour. Malac.),
55, 257–263.
Okutani, T., S. Kojima and J. Ashi (1997): Further discovery of
new taxon of vesicomyid clam from the Nankai Trough, off
Honshu, Japan. Venus (Jap. Jour. Malac.), 56, 185–188.
Okutani, T., K. Fujikura and S. Kojima (1999): Two new hadal
bivalves of the family Thyasiridae from the plate convergent area of the Japan Trench. Venus (Jap. Jour. Malac.),
58, 49–54.
Okutani, T., K. Fujikura and S. Kojima (2000a): New taxa and
review of vesicomyid bivalves collected from the Northwest Pacific by deep sea research systems of Japan Marine
Science & Technology Center. Venus (Jap. Jour. Malac.),
59, 83–101.
Okutani, T., T. Sasaki and S. Tsuchida (2000b): Two additional
new species to gastropod fauna of chemosynthetic site on
North Knoll of Iheya Ridge, Okinawa Trough. Venus (Jap.
Jour. Malac.), 59, 267–275.
Ono, T., K. Fujikura, J. Hashimoto, Y. Fujikura and S. Segawa
(1996): The hydrothermal vent community at the Kaikata
Seamount near Ogasawara (Bonin) Islands, South Japan.
JAMSTEC J. Deep Sea Res., 12, 221–230 (in Japanese with
English abstract).
Pautot, G., K. Nakamura, P. Huchon, J. Angelier, J. Bourgois,
K. Fujioka, T. Kanazawa, Y. Nakamura, Y. Ogawa, M.
Séguret and A. Takeuchi (1987): Deep-sea submersible survey in the Suruga, Sagami and Japan Trenches: preliminary
results of the 1985 Kaiko cruise, Leg 2. Earth Planet. Sci.
Lett., 83, 300–312.
Peek, A. S., R. G. Gustafson, R. A. Lutz and R. C. Vrijenhoek
(1997): Evolutionary relationships of deep-sea hydrothermal vent and cold-water seep clam (Bivalvia:
Vesicomyidae): results from the mitochondrial cytochrome
oxidase subunit I. Mar. Biol., 130, 151–161.
Peek, A. S., R. A. Feldman, R. A. Lutz and R. C. Vrijenhoek
(1998): Cospeciation of chemoautotrophic bacteria and deep
sea clams. Proc. Natl. Acad. Sci. USA, 95, 9962–9966.
Saino, T. and S. Ohta (1989): 13C/ 12C and 15N/ 14N ratios of
vesicomyid clams and a vestimentiferan tube worm in the
subduction zone east of Japan. Palaeogeogr. Palaeoclimatol.
Palaeoecol., 71, 169–178.
Scarlato, O. A. (1981): Bivalve mollusks from Middle Latitudes
of the Western Pacific. Academy of Sciences of USSR, Institute of Zoology, Leningrad, 479 pp. (in Russian).
Shinjo, R., H. Machiyama, Y. Maki, I. Motoyama, G. Toyama,
S. Hokakubo and S. Chung (2001): Outline of SHINKAI2000 dive surveys at the westernmost part of the Okinawa
Trough and the Kuroshima Knoll—preliminary report of the
NT00-06 Leg-1 Cruise—. JAMSTEC J. Deep Sea Res., 19,
109–121 (in Japanese with English abstract).
Shirayama, Y. (1992): Studies of meiofauna collected from the
Iheya Ridge during the dive 541 of the “SHINKAI 2000”.
Proc. JAMSTEC Symp. Deep Sea Res., 8, 287–290 (in Japanese with English abstract).
Shirayama, Y. and S. Ohta (1990): Meiofauna in a cold-seep
community off Hatsushima, Central Japan. J. Oceanogr. Soc.
Japan, 46, 118–124.
Sibuet, M. and K. Olu (1998): Biogeography, biodiversity and
fluid dependence of deep-sea cold-seep communities at active and passive margins. Deep-Sea Res. II, 45, 517–567.
Sibuet, M., S. K. Juniper and G. Pautot (1988): Cold-seep
benthic communities in the Japan subduction zones: Geological control of community development. J. Mar. Res.,
46, 333–348.
Southward, E. C. (1991): Three new species of Pogonophora,
including two vestimentiferans, from hydrothermal sites in
the Lau Back-arc Basin (Southwest Pacific Ocean). J. Nat.
Hist., 25, 859–881.
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
361
Southward, E. C. and S. V. Galkin (1997): A new vestimentiferan
(Pogonophora: Obturata) from hydrothermal vent fields in
the Manus Back-Arc Basin (Bismark Sea, Papua New
Guinea, Southwest Pacific Ocean). J. Nat. Hist., 31, 43–
55.
Stein, J. L., S. C. Cary, R. R. Hessler, S. Ohta, R. D. Vetter, J. J.
Childress and H. Felbeck (1988): Chemoautotrophic symbiosis in a hydrothermal vent gastropod. Biol. Bull., 174,
373–378.
Strizhov, V. P., A. P. Kuznetsov and N. V. Gurina (1990): Compounds of early diagenesis of sediments as energy source
in food web of bottom community of Conchocera sp.
(Bivalvia) around gasohydrate vents near the Paramushir
Island (the Sea of Okhotsk). Okeanologia, 30, 666–671 (in
Russian with English abstract).
Suzuki, T., T. Takagi and S. Ohta (1988): N-Terminal amino
acid sequence of the deep-sea tube worm haemoglobin remarkably resembles that of annelid haemoglobin. Biochem.
J., 255, 541–545.
Suzuki, T., T. Takagi and S. Ohta (1989a): Primary structure of
a dimeric hemoglobin from the deep-sea cold seep clam,
Calyptogena soyoae. Biochem. J., 260, 177–182.
Suzuki, T., T. Takagi and S. Ohta (1989b): Amino acid sequence
of the dimeric hemoglobin (Hb I) from the deep-sea coldseep
clam, Calyptogena soyoae and the phylogenetic relationship with other molluscan globlins. Biochem. Biophys. Acta,
999, 254–259.
Suzuki, T., T. Takagi, K. Okuda, T. Furukohri and S. Ohta
(1989c): The deep-sea tube worm haemoglobin: subunit
structure and phylogenetic relationship with annelid
hemoglobin. Zool. Sci., 6, 915–926.
Suzuki, T., H. Kawamichi, R. Ohtsuki, M. Iwai and K. Fujikura
(1998): The cDNA-derived amino acid sequence of
hemoglobin from Calyptogena kaikoi. JAMSTEC J. Deep
Sea Res., 14, 507–513 (in Japanese with English abstract).
Takai, K. and K. Horikoshi (1999): Genetic diversity of archaea
in deep-sea hydrothermal vent environments. Genetics, 152,
1285–1297.
Takasugi, H., T. Naganuma, K. Fujikura, T. Yamamoto, M.
Hattori, H. Yamamoto, M. Iwai and Y. Ogawa (1998): Analysis of methane-seep microbial communities based on sediment fatty acid analysis. JAMSTEC J. Deep Sea Res., 14,
609–619 (in Japanese with English abstract).
Takeda, M. and J. Hashimoto (1990): A new species of the genus Paralomis (Crustacea, Decapoda, Lithodidae) from the
Minami-Ensei Knoll in the Mid-Okinawa Trough. Bull. Nat.
Sci. Mus. Ser. A, 16, 79–89.
Takeda, M., J. Hashimoto and S. Ohta (2000): A new species of
the family Bythograeidae (Crustacea, Decapoda, Brachyura)
from the hydrothermal vents along volcanic front of the
Philippine Sea Plate. Bull. Nat. Sci. Mus. Ser. A, 26, 159–
172.
Takeuchi, A. (2000): Bottom response to a tsunami earthquake:
Submersible observations in the epicenter area of the 1993
earthquake off southwestern Hokkaido, Sea of Japan. J.
Geophys. Res., 103, 24109–24125.
Takeuchi, A., E. Tokuyama, S. Miyashita, J. Naka, H. Momma,
K. Ohtsuka, W. Soh, J. Ishii, T. Sagayama and S. Kuramoto
(1992): Discovery of open gashes with bacteria mat and liv-
362
S. Kojima
ing communities at the frontal thrust of the tectonic belt
along eastern margin of Japan Sea. Proc. JAMSTEC Symp.
Deep Sea Res., 8, 29–39 (in Japanese with English abstract).
Takeuchi, A., Y. Okamura, Y. Kato, K. Ikehara, J. Zhang, K.
Satake, T. Nagao, M. Hirano and M. Watanabe (2000): Large
earthquakes and bottom disturbances in the Okushiri ridge
along the eastern margin of Japan Sea. JAMSTEC J. Deep
Sea Res. II, 16, 29–46 (in Japanese with English abstract).
Tanaka, T. and J. Hashimoto (1991): Distribution of recent
Calyptogena bivalves in Sagami Bay. Chikyu Monthly, 13,
42–46 (in Japanese).
Tanaka, T., H. Hotta, H. Sakai, J. Ishibashi, T. Oomori, E. Izawa
and N. Oda (1990): Occurrence and distribution of the hydrothermal deposits in the Izena Hole, Central Okinawa
Trough. JAMSTEC Deepsea Res., 6, 11–26 (in Japanese with
English abstract).
Tanishima, M., J. Hashimoto, K. Fujikura, Y. Fujiwara and J.
C. Hunt (1996): A trial to elucidate locomotory activity
patterns in a deep-sea crab (Family: Bythograeidae).
JAMSTEC J. Deep Sea Res., 12, 231–238 (in Japanese with
English abstract).
Tsuchida, S. and K. Fujikura (2000): Heterochely, relative
growth, and gonopod morphology in the bythograeid crab,
Austinograea williamsi (Decapoda, Brachyura). J. Crust.
Res., 20, 407–414.
Tsuchida, S., K. Fujikura, Y. Fujiwara, J. C. Hunt, D. J. Lindsay
and J. Hashimoto (1998): Molting of bythograeid crabs
under laboratory conditions. JAMSTEC J. Deep Sea Res.,
14, 515–520 (in Japanese with English abstract).
Tsuchida, S., K. Watanabe, J. Ishibashi, H. Miyake, H. Watabe,
T. Yamaguchi, T. Kitajima, A. Nakano, M. Matsumura and
H. Watanabe (2000): Preliminary report of a biological,
geological, and geochemical survey on hydrothermalism at
the Hatoma and Minna Knolls. JAMSTEC J. Deep Sea Res.,
17, 35–42 (in Japanese with English abstract).
Tsuchida, S., H. Kumagai, J. Ishibashi, H. Watanabe and F.
Kouzuma (2001): Preliminary report of hydrothermalism
and volcanism at the Kaikata Seamount. JAMSTEC J. Deep
Sea Res., 18, 209–215 (in Japanese with English abstract).
Tsukahara, J. (1994): Transfer of endosymbiotic bacteria in the
ovary of the giant clam, Calyptogena soyoae. Zool. Sci.,
11, Suppl., 75 (abstract).
Tsunogai, U., J. Ishibashi, H. Wakita, T. Gamo, T. Masuzawa,
T. Nakatsuka, Y. Nojiri and T. Nakamura (1996): Fresh water
seepage and pore water recycling on the seafloor: Sagami
Trough subduction zone, Japan. Earth Planet. Sci. Lett., 138,
157–168.
Tsunogai, U., J. Ishibashi, H. Wakita and T. Gamo (1998):
Methane-rich plumes in the Suruga Trough (Japan) and their
carbon isotope characterization. Earth Planet. Sci. Lett.,
160, 97–105.
Tunnicliffe, V. and C. M. R. Fowler (1996): Influence of seafloor spreading on the global hydrothermal vent fauna. Nature, 379, 531–533.
Tunnicliffe, V., A. G. McArthur and D. McHugh (1998): A biogeographical perspective of the deep-sea hydrothermal vent
fauna. Adv. Mar. Biol., 34, 353–442.
Van Dover, C. (2000): The Ecology of Deep-Sea Hydrothermal
Vents. Princeton University Press, Princeton, 424 pp.
Von Cosel, R., B. Métivier and J. Hashimoto (1994): Three new
species of Bathymodiolus (Bivalvia: Mytilidae) from hydrothermal vents in the Lau Basin and the North Fiji Basin,
western Pacific, and the Snake Pit area, Mid-Atlantic Ridge.
The Veliger, 37, 374–392.
Vrijenhoek, R. C., R. A. Feldman, R. A. Lutz, C. Craddock and
J. Hashimoto (1997): Genetic characterization of
Lepetodrilus limpets from hydrothermal vents in the
Mariana Trough. JAMSTEC J. Deep Sea Res., Special Volume “Deep sea research in subduction zones, spreading
centers and backarc basins”, 111–116.
Watabe, H. and H. Miyake (2000): Decapod fauna of the hydrothermally active and adjacent fields on the Hatoma Knoll,
southern Japan. JAMSTEC J. Deep Sea Res., 17, 29–34 (in
Japanese with English abstract).
Yamaguchi, T. and W. A. Newman (1990): A new and primitive
barnacle (Cirripedia: Balanomorpha) from the North Fiji
Basin abyssal hydrothermal field, and its evolutionary implications. Pacific Sci., 44, 135–155.
Yamaguchi, T. and W. A. Newman (1997): The hydrothermal
vent barnacle Eochinelasmus (Cirripedia: Balanomorpha)
from the North Fiji, Lau and Manus Basins, South-West
Pacific. Zoosystema, 19, 623–649.
Yamamoto, T., T. Kobayashi, K. Nakasone and S. Nakao (1999):
Chemosynthetic community at North Knoll, Iheya Ridge,
Okinawa Trough. JAMSTEC J. Deep Sea Res. I, 15, 19–24.
Yamanaka, T., C. Mizota, T. Murae and J. Hashimoto (1999): A
currently forming petroleum associated with hydrothermal
mineralization in a submarine caldera, Kagoshima Bay, Japan. Geochem. J., 33, 355–367.
Yamanaka, T., C. Mizota, Y. Maki, K. Fujikura and H. Chiba
(2000): Sulfur isotope composition of soft tissues of deepsea mussels, Bathymodiolus spp., in Japanese waters.
Benthos Res., 55, 63–68.
Yoshida, N. and M. Ishida (1989): A gamma-ray spectral and
bacteria population surveys on the giant clam colonies,
Okinoyama Bank, Sagami Bay. JAMSTEC Deepsea Res.,
5, 15–21 (in Japanese with English abstract).
Yoshida, N. and H. Tsukahara (1997): The upwelling fluid on
the biological community on the subduction zone Sagami
Bay. JAMSTEC J. Deep Sea Res., Special Volume “Deep
sea research in subduction zones, spreading centers and
backarc basins”, 229.
Zhang, J. and T. Ishii (2000): Rare earth and trace element biogeochemistry of white clam in off Hatsushima cold seepage. JAMSTEC J. Deep Sea Res. II, 16, 155–161.
Deep-Sea Chemoautosynthesis-Based Communities in the Northwestern Pacific
363