Miocene vesicomyid species (Bivalvia) from Wakayama in southern Honshu, Japan

Abstract

A fossil association of potentially chemosymbiotic bivalves is reported from the lower Miocene Shikiya Formation in Kii Oshima Island, Wakayama Prefecture, Japan. The association is dominated by the elongate vesicomyid species Adulomya uchimuraensis (Kuroda, 1931); a second vesicomyid of lower abundance is here described as Archivesica sakoi new species. It represents the oldest species of the genus Archivesica and has a similar hinge structure as " Calyptogena " laubieri (Okutani and Mé tivier, 1986). Additional faunal elements include chemosymbiotic bivalves, namely the solemyid Acharax cf. yokosukensis Kanie and Kuramochi, 1995, and the lucinids Lucinoma? sp. and Poumea? sp., as well as the naticid gastropod Euspira meisensis (Makiyama, 1926). This association of fossil bivalves all with well-known chemosymbiotic living relatives suggests that these specimens lived at an ancient cold seep, and their mode of occurrence as mostly articulated shells parallel to the bedding plane indicates that were transported a short distance and then quickly buried.

Full text

Miocene vesicomyid species (Bivalvia) from Wakayama
in southern Honshu, Japan
Kazutaka Amano
Department of Geoscience
Joetsu University of Education
Joetsu 943-8512, JAPAN
amano@juen.ac.jp
Robert G. Jenkins
School of Natural System
College of Science and Engineering
Kanazawa University Kanazawa City
Ishikawa 920-1192, JAPAN
robertgj@staff.kanazawa-u.ac.jp
Masaaki Ohara
Wakayama Prefectural Museum
of Natural History, Kainan City
Wakayama 642-0001, JAPAN
ohara_m0002@pref.wakayama.lg.jp
Steffen Kiel
Georg-August University Go
¨ttingen
Geoscience Center, Geobiology Group
Goldschmidtstr. 3
37077 Go
¨ttingen, GERMANY
skiel@uni-goettingen.de
ABSTRACT
A fossil association of potentially chemosymbiotic bivalves is
reported from the lower Miocene Shikiya Formation in Kii
Oshima Island, Wakayama Prefecture, Japan. The association
is dominated by the elongate vesicomyid species Adulomya
uchimuraensis (Kuroda, 1931); a second vesicomyid of lower
abundance is here described as Archivesica sakoi new spe-
cies. It represents the oldest species of the genus Archivesica
and has a similar hinge structure as “Calyptogena” laubieri
(Okutani and Me
´tivier, 1986). Additional faunal elements
include chemosymbiotic bivalves, namely the solemyid
Acharax cf. yokosukensis Kanie and Kuramochi, 1995, and
the lucinids Lucinoma? sp. and Poumea? sp., as well as
the naticid gastropod Euspira meisensis (Makiyama, 1926).
This association of fossil bivalves all with well-known
chemosymbiotic living relatives suggests that these specimens
lived at an ancient cold seep, and their mode of occurrence
as mostly articulated shells parallel to the bedding plane
indicates that were transported a short distance and then
quickly buried.
Additional keywords: Miocene, Archivesica, Adulomya
INTRODUCTION
Vesicomyidae is one of the six extant bivalve families
with a chemosymbiotic mode of life, with have a fossil
history ranging back to the middle Eocene some 47
Million years ago (Kiel, 2010a). The vesicomyid fossil
record of the Northwestern Pacific region is increas-
ingly well studied and revealed the successive appear-
ance, dominance, and disappearance of several genera.
Hubertschenckia Takeda, 1953 was the only genus present
during the late Eocene and Oligocene and it disap-
peared afterward; the elongate Adulomya Kuroda,
1931 dominated during the early and middle Miocene;
andinthelateMiocenebothArchivesica Dall, 1908
and Calyptogena Dall, 1891 appeared and are the
dominant vesicomyid genera until the present day
(Amano and Jenkins, 2011; Amano and Kiel; 2007;
2010; 2011; 2012; Sasaki et al. 2005). Among those
two genera, Archivesica was, and still is, much more
species rich than Calyptogena.
When Amano and Jenkins (2011) investigated the fos-
sil record of extant vesicomyid species in Japan, they
questioned an identification by Katto and Masuda
(1978) of an early Miocene species from the lower
Miocene Shikiya Formation in southern Honshu as
Akebiconcha cf. kawamurai Kuroda, which is an older
name for the extant Archivesica kawamurai. Amano and
Jenkins (2011) suspected that the Shikiya species may
belong to Archivesica, but was not A. kawamurai. More-
over, Katto and Masuda (1978) identified an elongate
bivalve from the same locality as Cultellus izumoensis
Yokoyama, although the lot documentation was poor.
Based on observations on the original material of Katto
and Masuda (1978) and new material collected at their
locality, both species are here identified as members of
the Vesicomyidae: the ovate specimens previously iden-
tified as Akebiconcha cf. kawamurai is named as a new
species of the genus Archivesica, and the specimens
assigned to Cultellus izumoensis are identified as
Adulomya uchimuraensis Kuroda, 1931. Here we
describe these species and discuss their evolutionary sig-
nificance. We also describe the locality in more detail,
including its paleoecology and the mode of occurrence of
these fossils and additional taxa.
THE NAUTILUS 128(1):9–17, 2014 Page 9

MATERIALS AND METHODS
The specimens were recovered from a large block (ca.
744 m) and several small blocks in its vicinity along
the boulder beach near a cliff about 400 m north of
Oshima fishing port in Kii Oshima Island, Wakayama
Prefecture, Japan (Figure 1; Katto and Masuda, 1978,
Loc. no. 10). These blocks are derived from the cliff and
consist of sandy siltstone of the uppermost lower to low-
ermost middle Miocene Shikiya Formation of the
Kumano Group (Hisatomi, 1981). All shell material is
dissolved and rubber casts were made from the internal
and external molds to document the internal and exter-
nal morphology of the species, including the hinge struc-
ture. However, many small pyrite grains and the rough
surface of weathered shells prevent us from making rub-
ber casts without bubbles. All specimens reported here
are deposited at the Wakayama Prefectural Museum of
Natural History (WMNH).
MODE OF OCCURRENCE
The bivalves occur in shell beds consisting of mostly
articulated specimens lying parallel to the bedding plane,
although valves that are opened to various degrees were
also found (Figures 2–5). The matrix of these shell beds
does not differ from the surrounding sandy siltstone.
This mode of occurrence indicates that shells are not
preserved in life position but may instead have been
transported for a short distance and then quickly buried.
In addition to the two vesicomyid species described
here, two specimens of the giant solemyid Acharax cf.
yokosukensis Kanie and Kuramochi, 1995 (reaching
about 30 cm in length), one specimen each of the lucinid
bivalves Lucinoma? sp. and Poumea? sp., and of the
naticid gastropod Euspira meisensis (Makiyama, 1926)
were found (Figures 610).
SYSTEMATICS
Family Vesicomyidae Dall and Simpson, 1901
Subfamily Pliocardiinae Woodring, 1925
Genus Archivesica Dall, 1908
Type Species: Callocardia gigas Dall, 1896, from the
Gulf of California.
Archivesica sakoi new species
(Figures 11–17)
Akebiconcha cf. kawamurai Kuroda. Katto and Masuda,
1978: pl. 3, fig. 7.
Diagnosis: Medium-sized Archivesica with elongate
ovate shell, no lunular incision, shallow and wide pallial
sinus, and very shallow subumbonal pit; subumbonal
cardinal tooth consisting of very small anterior (3a) and
rather thick posterior (3b) teeth and thick cardinal tooth
Figure 1. Locality of the vesicomyid fossils (left-side base map is from the “Kushimoto”, scale 1:25,000 topographic map published
by the Geospatial Information Authority of Japan).
Page 10 THE NAUTILUS, Vol. 128, No. 1

(1) in right valve; subumbonal cardinal tooth consisting
of thin anterior ramus (2a) and very thick triangular
posterior tooth (2b) and posterior cardinal tooth (4b)
very thin in left valve.
Description: Shell medium-sized for genus (up to
80.4 mm in length), moderately inflated, elongate-ovate
(height/length ¼0.48–0.56), equivalve and inequilateral.
Antero-dorsal margin concave and continuing to rounded
anterior margin; postero-dorsal margin nearly straight,
posterior margin subtruncated; ventral margin broadly
arcuate. Beak prominent, prosogyrate and located at ante-
rior one-fifth to one-third of shell length (U%¼22–34).
Lunule absent. Surface ornamented with irregular com-
marginal lines. Right valve hinge wide for size, with three
cardinal teeth and very shallow subumbonal pit. Anterior
tooth (3a) in right valve very short and thin, connected
with posterior tooth (3b). Posterior tooth (3b) rather thick,
oblique posteriorly; middle cardinal tooth (1) moderately
thick, oblique anteriorly. Left valve hinge wide, with three
cardinal teeth. In left valve, anterior tooth (2a) thin, con-
nected with middle tooth (2b); middle tooth (2b) triangu-
lar and very thick; posterior cardinal tooth (4b) very thin.
Anterior adductor scar semicircular and deeply excavated;
anterior pedal adductor scar narrow, quadrate and deeply
excavated, distinct from adductor muscle scar; posterior
adductor scar indistinct, ovate, situated directly above a
weak ridge running from umbo to postero-ventral corner;
pallial sinus shallow and wide.
Type Material: Holotype: Right valve, length, 63.2 mm,
height, 31.3 mm, WMNH-Ge-5. Paratypes: Left valve,
length, 48.7 mmþ, height, 26.0 mm, WMNH-Ge-6;
left valve, length, 59.5 mm, height, 30.5 mmþ, WMNH-
Ge-7; left valve, length, 80.4 mm, height, 39.2 mm,
WMNH-Ge-8.
Type Locality: The coastal near cliff about 400 m
north to Oshima fishing port in Kii Oshima Island,
Kushimoto-cho, Wakayama Prefecture, Japan.
Figures 25. Field images of the outcrop. 24. Large block yielding vesicomyid fossils, white arrow indicates a gregarious
occurrence; enlargement of the southern side of the block (3), and close up of the gregarious occurrence (4). 5. Cluster of Adulomya
uchimuraensis Kuroda in a small block in the immediate vicinity of the large block.
K. Amano et al., 2014 Page 11

Material Examined: Twelve specimens from the
type locality.
Remarks: Archivesica sakoi new species was previously
reported as Akebiconcha cf. kawamurai Kuroda, 1943 by
Katto and Masuda (1978), but their illustration did not show
the hinge structure. The outline of Archivesica kawamurai
is indeed somewhat similar to that of A. sakoi except for
being higher and larger. However, the subumbonal cardinal
tooth of right valve enables us to easily separate both spe-
cies. The right valve of the present new species has a small
anterior tooth (3a) and posteriorly oblique posterior tooth
(3b), while that of A. kawamurai has a larger anterior tooth
and a posterior tooth that is slightly inclined anteriorly
(Kuroda, 1943; Sasaki et al., 2005).
Comparisons: The right valve hinge of Archivesica
sakoi is most similar to that of the Recent “Calyptogena”
laubieri Okutani and Me
´tivier, 1986 in having a small 3a
tooth and a shallow subumbonal pit (Figure 18). A recent
molecular phylogenetic analysis (Audzijonyte et al., 2012)
indicates that this species is phylogenetically close to the
type species of Archivesica,A. gigas (Dall, 1895). How-
ever, “C.” laubieri can be separated from the present new
species by having a distinct blunt ridge running from beak
to posterior end, a tapering posterior end, a very narrow
hinge plate, a less stout middle tooth of left valve, and
many distinct irregular growth lines on the outer surface
(Figures 19, 22). The extant Archivesica ochotica Scarlato,
1981 also resembles this new species in a having broadly
rounded ventral margin, a shallow pallial sinus, a rather
small 3a tooth, a shallow subumbonal pit in the right valve,
and a stout 2b tooth in the left valve (Figures 20, 21, 23).
However, A. ochotica has a larger (105.0 mm long) and
higher shell (H/L ¼0.58), an overhanging posterior tooth
above anterior and middle cardinal teeth of left valve, and
Figures 610. Mollusks associated with the vesicomyid bivalves described here. 6, 7. The lucinid bivalve Poumea? sp. (WMNH-
Ge-1120210284); 6, rubber cast of the inner surface of a left valve showing elongate anterior adductor scar (AAS); 7, internal mold. 8.
The naticid gastropod Euspira meisensis (Makiyama) (WMNH-Ge-1120210285), rubber cast. 9. The lucinid bivalve Lucinoma? sp.
(WMNH-Ge-1120210287), rubber cast of the external surface of a right valve. 10. The solemyid bivalve Acharax cf. yokosukensis
Kanie and Kuramochi (WMNH-Ge-1120210291), internal mold of a left valve.
Page 12 THE NAUTILUS, Vol. 128, No. 1

a more roundly curved postero-dorsal margin than
Archivesica sakoi new species.
Distribution: Lower Miocene Shikiya Formation at
the type locality.
Etymology: Named after Mr. Yukio Sako who col-
lected the type specimens and kindly offered them to
the authors for study.
Genus Adulomya Kuroda, 1931
Type Species: Adulomya uchimuraensis Kuroda,
1931, from the middle Miocene Bessho Formation, cen-
tral Honshu, Japan.
Adulomya uchimuraensis Kuroda, 1931
(Figures 24–29)
Adulomya uchimuraensis Kuroda, 1931: 27–28, pl. 13, figs. 111–
114; Tanaka, 1959: 117–118, pl. 1, fig. 1–10; Tanaka, 1960:
24–26, pl. 32, figs. 1–7; Amano and Kiel, 2011: 77–80, fig. 2.
Figures 1117. Archivesica sakoi new species. 11, 12, 15. Paratype (WMNH-Ge- 1120210286); 11, a rubber cast showing details
of the hinge of a right valve; SP¼subumbonal pit; 12, ventral view showing the anterior pedal retractor scar (APRS); 15, internal
mold of left valve. 13, 16. Paratype (WMNH-Ge-1120210288); 13, rubber cast showing details of the hinge area of a left valve;
16, view on the left valve of an internal mold, white arrow indicates the very shallow pallial sinus; PAS¼posterior adductor scar.
14. Holotype (WMNH-Ge-1120210283); view on the right valve of an internal mold; AAS¼anterior adductor scar. 17. Paratype
(WMNH-Ge-1120210289), rubber cast showing the outer surface of a left valve.
K. Amano et al., 2014 Page 13

Calyptogena (Adulomya)uchimuraensis Kuroda.—Kanno and
Tanaka in Kanno et al., 1998: 20–22, figs. 7–8.
Calyptogena (Adulomya)uchimuraensis kurodai Kanno and
Tanaka in Kanno et al., 1998: 22–25, figs. 9–10.
Akebiconcha chitanii (Kanehara).—Kanno and Ogawa, 1964:
pl.1, figs. 17–18.
Adulomya uchimuraensis Kuroda.—Hayashi and Miura,
1973: pl. 1, fig. 15.
Cultellus izumoensis Yokoyama.—Katto and Masuda, 1978:
pl. 3, figs. 8–9.
non Akebiconcha uchimuraensis Kuroda.—Matsumoto and
Hirata, 1972: 755–757, pl. 1, figs. 1–8, pl. 2, figs. 1–2.
Material Examined: Fifty-three specimens were
examined.
Remarks: Katto and Masuda (1978) illustrated two
elongate specimens as Cultellus izumoensis. Our exami-
nation of the hinge of their illustrated specimen (Katto
and Masuda, 1978: pl. 3, fig.9; Figures 15, 19 herein)
indicates that they belong to Adulomya uchimuraensis
for the following reasons: the left valve has an anterior
(2a) and a middle cardinal (2b) tooth and a low posterior
tooth (4b), and the right valve has only two teeth (cardi-
nals 1, 3b); the anterior retractor scar is very deep, and
some examined specimens show that the pallial line is
entire (Figure 16). Such characteristics are never seen
in any cultellid species. Cultellid species have only
two protruded cardinal teeth in each valve and pallial
sinus present.
Comparison: As discussed by Amano and Kiel (2011),
Adulomya uchimuraensis can be separated from the
early to middle Miocene Adulomya chitanii Kanehara,
1937 by its larger size and the lack of a pallial sinus.
Although the maximum size of Adulomya uchimuraensis
at Kii Oshima (94.1 mm long) is smaller than the maxi-
mum size of the specimens of A. uchimuraensis at its
type locality in the Bessho Formation (180 mm; Amano
and Kiel, 2011), it is still larger than A. chitanii,
which reaches only up to 70.4 mm in length (Amano
and Kiel, 2011).
Measurements: Left valve, length, 83.2 mm, height,
21.4 mm, WMNH-Ge-9; right valve, length, 89.7 mm,
height, 17.8 mm, WMNH-Ge-10; right valve, length,
94.1 mm, height, 23.9 mm, WMNH-Ge-11.
Distribution: Lower Miocene Shikiya Formation in
Wakayama Prefecture, lowest middle Miocene Takinoue
Formation in Hokkaido and middle Miocene Bessho
Formation in Nagano Prefecture.
Figures 18, 19, 22. “Calyptogena” laubieri Okutani and Me
´tivier, 1986, Holotype (NSMT Mo 64158); 18, hinge dentition
of a right valve; SP¼subumbonal pit; 19, hinge dentition of a left valve; 22, right side view of the holotype. Figures 20, 21, 23.
Archivesica ochotica Scarlato, 1981, Holotype (ZIN AN SSSR no. 9912); 20, hinge dentition of a right valve; 21, hinge dentition
of a left valve; 23, left side view of the holotype.
Page 14 THE NAUTILUS, Vol. 128, No. 1

DISCUSSION
Recent studies on presumed Paleogene members of
Archivesica from western North America (Amano and
Kiel, 2007; Kiel and Amano, 2010) indicate that these
species may not belong to Archivesica but instead to the
genus Pliocardia or a related new genus (Amano and
Kiel, 2012). Thus, the early Miocene Archivesica sakoi
new species described here from southern Japan repre-
sents the oldest species of the genus Archivesica. How-
ever, this does not change our earlier observation that
Adulomya was the dominant vesicomyid taxon during
the early and middle Miocene in Japan because
Adulomya uchimuraensis is far more common at the
Kii Oshima site than Archivesica sakoi.
The association reported here from Kii Oshima
consists almost exclusively of potentially chemosym-
biotic bivalves (e.g., Taylor and Glover, 2010): domi-
nant are the vesicomyids Adulomya uchimuraensis
and Archivesica sakoi with a minor number of lucinids
(Lucinoma?sp.andPoumea? sp.) and a solemyid
(Acharax cf. yokosukensis). Such an association domi-
nated by vesicomyid bivalves is typical for extant cold
seep communities (Paull et al. 1985; Levin 2005; Majima
et al. 2005; Campbell, 2006) and suggests that the
Kii Oshima fauna might have lived at an ancient cold
seep. Predatory gastropods such as the naticid Euspira
meisensis found along with the chemosymbiotic bivalves
are common in soft sediments in many marine environ-
ments, but they are also frequently found at ancient
cold-seep communities, especially in the North Pacific
realm (Amano et al. 2010; Kiel, 2010b). Due to the
transported nature of the association and the lack of
carbonate, the usual approach to identify an ancient
cold-seep deposit by stable carbon isotope analysis
(cf. Peckmann and Thiel, 2004) is not possible. This is
so far the only record of a vesicomyid-dominated
faunule from the Shikiya Formation and from southern
Honshu (cf., Majima et al. 2005).
Only about 4 km west of the Kii-Oshima locality, a
hydrocarbon-seep fauna has been recently been reported
by the present authors from the upper Eocene to lower
Oligocene(?) Tanamigawa Formation at Tanosaki
(Amano et al., 2013). That Paleogene community differs
from the early Miocene fauna from Kii Oshima
reported here by consisting of a different vesicomyid
genus (Hubertschenckia), the bathymodiolin mussel
Bathymodiolus, the thyasirid bivalve Conchocele and the
absence of lucinids. As already pointed out by Amano
et al. (2013), Paleogene seep communities in Japan
generally lack lucinid bivalves, while lucinids are generally
present at Miocene seep communities as in the the
Kii Oshima fauna reported here.
ACKNOWLEDGMENTS
We thank Yukio Sako (Kushimoto-cho), Yukito Kurihara
(Mie University) and Hiroshi Saito (National Science
Museum) for their help in examining some fossil or
recent specimens; Elena Krylova (P.P. Shirshov Institute
of Oceanology, Russian Academy of Sciences) for
discussions on vesicomyid systematics, for her helpful
review of the manuscript, and for allowing us to use her
images of A. ochotica. An anonymous reviewer is
thanked for suggestions that helped improve on the
manuscript. This study was partly supported by a Grant-
in-aid for Scientific Research from the Japan Society for
Promotion of Science (C, 23540456, 2011-2013) to KA,
Grant-in-Aid for JSPS Fellows to RGJ, and by the
Deutsche Forschungsgemeinschaft through grant
Ki802/6-1 to SK.
Figures 24–29. Adulomya uchimuraensis (Kuroda), 24, 26,
28. Right valve (WMNH-Ge-1120210293); 24,rubbercast
showing the hinge dentition and the anterior pedal retractor scar
(APRS) of a right valve; 26, rubber cast showing the anterior
(AAS) and posterior adductor scars (PAS) in a right valve, white
arrow indicates the posteriormost portion of the entire pallial
line; 28, rubber cast showing the outer surface of this shell. 25,
29. The specimen illustrated by Katto and Masuda (1978, pl. 3,
fig. 9)(WMNH-Ge-1120210292); 25, rubber cast showing
details of the hinge area; 29, left valve. 27. Rubber cast of the
outer surface of a right valve (WMNH-Ge-1120210290).
K. Amano et al., 2014 Page 15

LITERATURE CITED
Amano, K. and R.G. Jenkins. 2011. Fossil records of extant
vesicomyid species from Japan. Venus 69: 163176.
Amano, K and S. Kiel. 2007. Fossil vesicomyid bivalves from
the North Pacific region. The Veliger 49: 270293.
Amano, K and S. Kiel. 2010. Taxonomy and distribution of
fossil Archivesica (Bivalvia: Vesicomyidae) in Japan. The
Nautilus 124: 155165.
Amano, K and S. Kiel. 2011. Fossil Adulomya (Vesicomyidae,
Bivalvia) from Japan. The Veliger 51: 7690.
Amano, K and S. Kiel. 2012. Two Neogene vesicomyid species
(Bivalvia) from Japan and their biogeographic implica-
tions. The Nautilus 126: 7985.
Amano, K., R.G. Jenkins, M. Aikawa, and T. Nobuhara, 2010. A
Miocene chemosynthetic community from the Ogaya
Formation in Joetsu: evidence for depth-related eco-
logic control among fossil seep communities in the Japan
Sea back-arc basin. Palaeogeography, Palaeoclimatology,
Palaeoecology, 286: 164-170.
Amano, K., R. G. Jenkins, Y. Sako, M. Ohara and S. Kiel. 2013.
A Paleogene deep-sea methane-seep community from
Honshu, Japan. Palaeogeography, Palaeoclimatology,
Palaeoecology 387: 126133.
Audzijonyte, A., E.M. Krylova, H. Sahling, and R.C.
Vrijenhoek. 2012. Molecular taxonomy reveals broad
trans-oceanic distributions and high species diversity of
deep-sea clams (Bivalvia: Vesicomyidae: Pliocardiinae) in
chemosynthetic environments. Systematics and Biodiver-
sity 10: 403-415.
Campbell, K.A. 2006. Hydrocarbon seep and hydrothermal
vent paleoenvironments and paleontology: Past develop-
ments and future research directions. Palaeogeography,
Palaeoclimatology, Palaeoecology 232: 362-407.
Dall, W.H. 1891. Scientific results of explorations by the U.S.
Fish Commission Steamer Albatross. XX. On some new
or interesting West American shells obtained from
dredgings of the U.S. fish commission steamer Albatross
in 1888. Proceedings of the U.S. National Museum 14:
174191.
Dall, W.H. 1896. Diagnoses of new species of mollusks from
the west coast of America. Proceedings of the U.S.
National Museum of Natural History 18: 7–20.
Dall, W.H. 1908. Reports on the dredging operations off the
west coast of Central America ... The Mollusca and
Brachiopoda. Bulletin of the Museum of Comparative
Zoology at Harvard University 43: 205487.
Dall, W.H. and C.T. Simpson. 1901. The Mollusca of Porto
Rico. Bulletin of the United States, Fish and Fisheries
Commission 20: 351524.
Hayashi, T. and Y. Miura. 1973. The Cenozoic sediments in
the southern part of Okazaki City, central Japan. Bulle-
tin of the Aichi University of Education 22: 133–149
[in Japanese with English abstract].
Hisatomi, K. 1981. Geology and sedimentology of the Kumano
Group in the southeastern part of the Kumano basin, Kii
Peninsula. Journal of Geological Society of Japan 87:
157174 [in Japanese with English abstract].
Kanehara, K. 1937. Miocene shells from the Joban coal-field.
Bulletin of the Imperial Geological Survey of Japan
27: 112.
Kanie, Y. and T. Kuramochi. 1995. Acharax yokosukensis, n. sp.
(gigantic Bivalve) from the Miocene Hayama Formation
of the Miura Peninsula, south-central Japan. Science
Report of Yokosuka City Museum 43: 51–57.
Kanno, S. and H. Ogawa. 1964. Molluscan fauna from the
Momijiyama and Takinoue districts, Hokkaido, Japan. Sci-
ence Reports of the Tokyo Kyoiku Daigaku, Section C 8:
269–294.
Kanno, S., K. Tanaka, H. Koike, K. Narita, and T. Endo. 1998.
Adulomya uchimuraensis Kuroda (Bivalvia) from the Mio-
cene Bessho Formation in Shiga-mura, Nagano Prefec-
ture, Japan. Research Report of the Shinshushinmachi
Fossil Museum 1: 17–28.
Katto, J. and K. Masuda. 1978. Tertiary Mollusca from the
southern part of Kii Peninsula, Wakayama Prefecture,
Southwest Japan. Research Report of Kochi University,
Natural Science 27: 97–111.
Kiel, S. 2010a. The fossil record of vent and seep mollusks. In,
Kiel, S. (ed.). The vent and seep biota, Topics in
Geobiology 33: 255277.
Kiel, S. 2010b. On the potential generality of depth-related
ecologic structure in cold-seep communities: Cenozoic
and Mesozoic examples. Palaeogeography, Palaeocli-
matology, Palaeoecology 295: 245–257.
Kiel, S. and K. Amano. 2010. Oligocene and Miocene
vesicomyid bivalves from the Katalla district in southern
Alaska, USA. The Veliger 51: 7684.
Kuroda, T. 1931. Fossil Mollusca. In: Homma, F. (ed.). Geol-
ogy of the central part of Shinano, part 4. Kokin Shoin,
Tokyo: 190 [in Japanese].
Kuroda, T. 1943. Akebiconcha, a new pelecypod genus. Venus
13: 14–18. [in Japanese with English description].
Levin, L.A. 2005. Ecology of cold seep sediments: inter-
actions of fauna with flow, chemistry and microbes.
Oceanography and Marine Biology Annual Review
43: 1–46.
Makiyama, J. 1926. Tertiary fossils from north Kankyo-do,
Korea. Memoirs of College of Science, Kyoto Imperial
University, Series B 2: 143160.
Majima, R., T. Nobuhara, and T. Kitazaki, 2005. Review of fossil
chemosynthetic assemblages in Japan. Palaeogeography,
Palaeoclimatology, Palaeoecology 227: 86–123.
Matsumoto, E. and M. Hirata. 1972. Akebiconcha uchimuraensis
(Kuroda) from the Oligocene formations of the Shimanto
terrain. Bulletin of the National Science Museum, Tokyo
15: 753–760.
Okutani, T. and B. Me
´tivier. 1986. Description of three new
species of vesicomyid bivalves collected by the submers-
ible Nautile from abyssal depths off Honshu, Japan. Venus
45: 147–160.
Paull, C.K., A.J.T. Jull, L.J. Toolin, and T. Linick. 1985. Stable
isotope evidence for chemosynthesis in an abyssal seep
community. Nature 317: 709–711.
Peckmann, J. and V. Thiel, 2004. Carbon cycling at ancient
methane-seeps. Chemical Geology 205: 443–467.
Sasaki, T., T. Okutani, and K. Fujikura. 2005. Molluscs
from hydrothermal vents and cold seeps in Japan: A
review of taxa recorded in twenty recent years. Venus
64: 87–133.
Scarlato, O.A. 1981. Bivalve molluscs from the middle latitude
of the western part of the Pacific Ocean. Taxonomic
Monograph of Fauna USSR 126: 1–479 [in Russian].
Takeda, H. 1953. The Poronai Formation (Oligocene Tertiary)
of Hokkaido and South Sakhalin and its fossil fauna. Stud-
ies on Coal Geology, the Hokkaido Association of Coal
Mining Technologists 3: 1103.
Tanaka, K. 1959. Molluscan fossils from central Shinano,
Nagano Prefecture, Japan (Part 1)—fossils from Akanuda
Page 16 THE NAUTILUS, Vol. 128, No. 1

limestone. Journal of Shinshu University, Faculty of Edu-
cation 8: 115–133.
Tanaka, K. 1960. Illustrated Cenozoic fossils, 31. On
Adulomya from the Bessho Formation and Anadara
from the Ogawa Formation. Cenozoic Research 31:
783– 787.
Taylor, J.D. and E.A. Glover. 2010. Chemosynthetic bivalves.
In: Kiel, S. (ed.). The vent and seep biota. Topics in
Geobiology. Springer, Heidelberg, pp. 107–136.
Woodring W.P. 1925. Miocene mollusks from Bowden,
Jamaica. Part I: Pelecypods and Scaphopods. Carnegie
Institution of Washington, Publication 366: 1–222.
K. Amano et al., 2014 Page 17