THE VELIGER# CMS, Inc., 2006

The Veliger 49(2):84–90 (July 2, 2007)

A New Gigantic Nucinella (Bivalvia: Solemyoida) from the Cretaceous
Cold-seep Deposit in Hokkaido, Northern JapanKAZUTAKA AMANO
Department of Geoscience, Joetsu University of Education, Joetsu 943-8512, Japan(e-mail: amano@juen.ac.jp)

ROBERT G. JENKINS
Department of Earth and Planetary Science, University of Tokyo, Tokyo 113-0033, JapanAND

YOSHINORI HIKIDA
Nakagawa Museum of Natural History, Nakagawa 098-2626, Japan
Abstract. Nucinella gigantea, sp. nov is described from the Campanian cold-seep deposits in Hokkaido, northern
Japan. This species is characterized by its gigantic size within the genus and by its radiating interior striation. Its
morphological features, associated fauna, lithofacies and carbon isotope suggest that N. gigantea lived in the bathyal
cold-seep site, and depended on chemosymbiosis. In addition, this new species is the oldest known Nucinella invaderof deep cold-seep environments.

INTRODUCTION
Species of the genus Nucinella Wood, 1851 are rare
bivalves ranging from shallow waters to lower bathyal
depths (La Perna, 2005, Table 1; Okutani & Iwasaki,2003). Based on the soft-part anatomy and shellstructure, the genus was transferred from Arcoida of
Pteriomorphia to Solemyoida of Protobranchia (Allen& Sanders, 1969; Waller, 1978).
In Japan, seven species of Recent and fossil
nucinellids have been described: Nucinella kanekoi
Matsukuma, Okutani & Tsuchi, 1982, N. viridis
Matsukuma, Okutani & Tsuchi, 1982, N. surugana
Matsukuma, Okutani & Tsuchi, 1982, Huxleya sulcata
A. Adams, 1860, H. decussata A. Adams, 1862, H.
ochiaiensis (Chinzei, 1959) and H. cavernicola Hayami& Kase, 1993. H. sulcata and the extinct H. ochiaiensis
have been recorded from the Plio-Pleistocene formations in the Northeast Honshu and Kanto district of
central Honshu (Baba, 1990) whereas N. kanekoi is also
known from the Pliocene Takanabe Formation of the
Miyazaki Group in Miyazaki Prefecture, southern
Kyushu (Matsukuma et al., 1982).
A new gigantic species of Nucinella was collectedfrom the Cretaceous cold-seep site in Nakagawa Town
in northern Hokkaido. This is the second record of
Nucinella from cold seep sites, after the Recent species
N. viridis (Okutani & Iwasaki, 2003; Sasaki et al.,2005). Therefore, this is the oldest record of Nucinella
in Japan and the first record from a fossil cold-seep site.

MATERIALS AND METHODS
Fossil specimens of the present new species werecollected from the Campanian Omagari Formation ofUpper Yezo Group (Takahashi et al., 2003) at about
400 m downstream of Abeshinai River from thejunction with Tanno-sawa Creek (Figure 1). The namecoined for the locality is Yasukawa site (Jenkins, 2006).
Four carbonate bodies, each larger than 50 cm indiameter, here named A to D, were identified at the
Yasukawa site. We examined the largest carbonate
body and surrounding sandy siltstone (Carbonate body
A, 1 m 3 2 m) which yields some small bivalves,
including the new species of Nucinella and gastropods(Jenkins, 2006). The new species of Nucinella was alsocollected from mudstone overlying the cemented
sandstone, and mudstone located laterally 5 m away
but on nearly the same horizon as Carbonate body A(the name coined for the locality is Yasukawa
Peripheral Site; YPS). All fossil specimens are housed
at the University Museum, the University of Tokyo(UMUT).
According to Jenkins et al. (2007), the carbonate
body can be divided into upper and lower parts (40 cm
and 60 cm thick, respectively). The lower part iscomposed mainly of fractured micritic limestone. The

K. Amano et al., 2006
Table 1
Associated species with a new species of Nucinella.
Species
Serradonta sp.
Ataphridae gen. et sp. indet.
Abyssochrysidae gen. et sp. indet.
Acharax cretacea Kanie and Nishida
Leionucula formosa (Nagao)
Acila (Truncacila) hokkaidoensis (Nagao)
Nuculana (Ezonuculana) mactraeformis Nagao
Propeamussium yubarense (Yabe and Nagao)
Myrtea ezoensis (Nagao)
M.? sp.
Miltha sp.
Thyasira sp.
Fissidentalium cf. otatumei (Nagao)

Page 85
fact, as well as the variable sulphide sulphur isotopic
compositions, high carbonate contents, and in situ
fractures in the carbonates, strongly suggest that thislocality was a methane-seep site (Jenkins, 2006).
On the measurement of shells, Matsukuma et al.
(1982) proposed to measure ‘‘shell length’’ along theantero-posterior axis passing through the supposed
locations of mouth and anus. However, most authors
measured shell length as parallel to hinge axis. We use
this traditional method to measure the shell length andheight, as claimed by La Perna (2005) for comparing
the new species with other ones. La Perna (2005)
measured height as the beak-ventral distance andlength as the antero-posterior one at right angle toheight.

SYSTEMATICS
upper part is mainly composed of cemented sandstone(i.e., grainstone) with cements of radiaxial bladedcalcite and sparry calcite. Isotopic composition of early
cemented phases, i.e., micrite and radiaxial bladedcalcite, ranges from 243.5 to 37.7% for d13C (vs. PDB)
and from 22.7 to 20.4% for d18O (vs. PDB). These
oxygen isotope values closely resemble the values from
Campanian shells of benthic animals examined fromthe outer shelf to upper slope settings in the northwestern Pacific (Moriya et al., 2003). The negative d13C
values of the early precipitated carbonates indicate that
their carbon was derived from biogenic and/or
thermogenic methane (Peckmann & Thiel, 2004). This

Family Manzanellidae Chronic, 1952
Remarks: This family includes Manzanella Girty, 1909
(Early Permian), Nucinella Wood, 1851 (Early Jurassic
to Recent) and Huxleya A. Adams, 1860 (Pliocene to
Recent). Assignment of these genera to a single family(Manzaniellidae) or two families (Manzaniellidae andNucinellidae Vokes, 1956) is controversial. Habe (1977)
and Pojeta (1988) separated Nucinellidae including
some living species from the extinct Manzanellidae.
Vokes (1956) considered Manzanella is included in thesame family to Nucinella and Huxleya when hesummarized supraspecific group of Nucinellidae andall species of Nucinella. Then, he regarded Nucinellidae
as a junior synonym of Manzanellidae (Vokes, 1980).
In his cladistic work on bivalves, Waller (1990, 1998)
shows high-rank segregation of Nucinellidae andManzanellidae from other solemyoids and consequently combined two families. Thus, we follow theconclusions of Vokes (1980) and Waller (1990, 1998).

Genus Nucinella Wood, 1851
Nucinella gigantea Amano, Jenkins and Hikida,sp. nov.
(Figures 2–7)
Diagnosis: A gigantic Nucinella having nuculid shape
and with many radiating interior striae and narrow flatarea below dentition.
Holotype: Length—18.4 mm, height- 18.8 mm, width10.4 mm, both valves, UMUT MM29245; Carbonate
body A.

Figure 1.

Locality of a new species of Nucinella.

Paratypes: Length—8.9 mm, height—9.8 mm, width—
4.9 mm, both valves, UMUT MM29246; length—
7.9 mm, height—7.9 mm, width—4.2 mm, both valves,
UMUT MM29247; length—6.5 mm, height—6.9 mm,

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The Veliger, Vol. 49, No. 2

Figures 2–7. Nucinella gigantea Amano, Jenkins & Hikida. Figure 2, Paratype, length 5 7.9 mm, UMUT MM29247. Figure 3,
Paratype, length 5 13.2 mm, UMUT MM29250. Figure 4, Paratype, length 5 12.1 mm, UMUT MM29249. Figure 5, Paratype,
length 5 6.5 mm, UMUT MM29248. Figures 6a–b, Paratype, length 5 8.9 mm, UMUT MM29246. Figures 7a–c, Holotype,
length 5 18.4 mm, UMUT MM29245; Figure 7b, rubber cast of Holotype.

width—3.1 mm, both valves, UMUT MM29248;
length—12.1 mm, height—5.4 mm (deformed), rightvalve, UMUT MM29249; YPS: Length—13.2 mm,height—11.2 mm, left valve, UMUT MM29250; Carbonate body A.
Type locality: 400 m downstream of Abeshinai River
from the junction with Tanno-sawa Creek, Nakagawa
Town, Hokkaido (44u409270N, 142u19450E); Omagari
Formation.
Stratigraphic and geographic distribution: Late Cretaceous (Campanian), Omagari Formation; known only
from the type locality.
Description: Shell gigantic in size, thin-walled, moderately inflated, nuculiform, equivalve, inequilateral.
Umbo projecting above dorsal margin, opisthogyrate,posteriorly situated at about one-third of shell length.
Antero-dorsal margin long, broadly arcuated, grading
into rounded anterior margin; antero-ventral margin
well rounded; postero-ventral margin nearly straight;
postero-dorsal margin short, steeply sloping, making
right angle with postero-ventral margin. Surface
smooth, except for very weak growth lines. Hinge
plate not so wide, consisting of subumbonal taxodont
teeth and anterior lateral tooth. In right valve; twocentral teeth large, but thin; more than two anterior
teeth small; two posterior teeth small, chevron shaped,
parallel to hinge, just behind central teeth; anterior

lateral tooth long and elevated. In left valve; one
central tooth strong; more than five posterior teeth
small; three anterior teeth parallel to hinge; one
anterior lateral tooth strong and long. Ligament pit
wide and deep, immediately situated at posterior ofbeak, occupying postero-dorsal margin. Monomyarian
condition advanced, anterior adductor muscle scarlarge and ovate. Inner surface with very fine radialstriations. Pallial line entire.
Comparison: Most species of fossil and Recent Nucinella have very small adult shells (length 5 about 1 to
3 mm). Exceptionally, three Recent species attaining
and even exceeding 1 cm in shell length are known
from the deep sea: Nucinella maxima (Thiele & Jackel,1931) from Zanzibar Channel, eastern Africa (463 m
deep), N. viridis Matsukuma, Okutani & Tsuchi, 1982
from off Nojimazaki, central Honshu (2630–3581 m
deep) and N. boucheti La Perna, 2005 from Bondoc
Peninsula, Philippines (1580–1610 m deep). N. gigantea
is a member of such a large shelled species group.
Among them, N. boucheti closely resembles N. gigantea
in its large shell size (length 5 25 mm), nuculiform
outline and fine radiating interior striation. However,N. boucheti differs from N. gigantea in having a wide
triangular flat area below dentition and more numerous teeth (13 in the holotype). N. maxima is anotherspecies similar to N. gigantea, with two vertical central
teeth, similar number of teeth (6 or 7) and interior

K. Amano et al., 2006
striation. Smaller size (L 5 12.5 mm) and the obtusely
rounded posterior corner of N. maxima enable us toseparate this species from N. gigantea. N. viridis differsfrom N. gigantea by having a smaller shell (L ,10.4 mm), fewer teeth (5) and obtusely rounded
posterior corner.
Two other species of Nucinella have been recorded
from Cretaceous deposits: N. glabrata Stoliczka, 1871
from the Ootatoor Group in South India and N. sohli
Pojeta, 1988 from the Blufftown Formation in Georgia,
USA (Vokes, 1956; Pojeta, 1988). However, their small
size (length 5 3.1 mm in N. glabrata and length 5
2 mm in N. sohli) easily distinguishes them from N.
gigantea. Although the inner surface of N. glabrata is
unknown in detail, N. sohli distinctly lacks the radiatinginterior striation.
Associated species and paleoenvironment: At the type
locality, this new species was collected from theCarbonate body A, and its surrounding sandy siltstone
and mudstone, and YPS together with many taxodont
bivalves, lucinids and thyasirids (Table 1). Amongthem, Acharax cretacea Kanie & Nishida, 2000, Myrtea
ezoensis (Nagao, 1938), Miltha sp. and Thyasira sp.
might be chemosymbiotic bivalves. In contrast, Leionucula formosa (Nagao, 1930), Acila (Truncacila)
hokkaidoensis Nagao, 1932, Nuculana (Ezonuculana)
mactraeformis Nagao, 1932 might be deposit feeders.
Propeamussium yubarense (Yabe & Nagao, 1928) isthought to be a carnivore (Hickman, 1984). The highratio (0.5) of protobranch bivalves to total bivalve
species shows a habitat deeper than the lower sublittoral zone (Amano et al., 1987; Amano & Nonaka,
2001; Amano, 2003). On the other hand, the Recent
records of small associated gastropod Serradonta
Okutani, Tsuchida & Fujikura, 1992 is restricted tobathyal depth in the Pacific (Warén & Bouchet, 2001;
Sasaki et al., 2005). Judging from these associatedfauna and lithology, this new species lived at aroundthe bathyal cold-seep site.
Etymology: This species named after its size.

DISCUSSION
La Perna (2005) suggested that the large body size of N.
boucheti, together with deep sea colonization, may be
an expression of the evolutionary diversification of
Nucinella. In fact, other than N. maxima living in theupper bathyal zone, large species of Nucinella now livein the middle to lower bathyal depth while most small
species live in the upper sublittoral to upper bathyalzones (La Perna, 2005). However, it is difficult to
explain how low nutrient conditions of the deep sea
could sustain such large species.
Taylor & Glover (2000) estimated chemosymbiosis
of fossil Lucinidae, partly based on the study ofPage 87
symbiosis-associated morphological features. Reid
(1990) listed the genera possessing sulphide-oxidizing
bacteria including Solemya Lamarck, 1818, Acharax
Dall, 1908, Calyptogena Dall, 1891, Bathymodiolus
Kenk & Wilson, 1985, Lucinoma Dall, 1901, Parvilucina Dall, 1901, Anodontia Link, 1807, Codakia
Scopoli, 1777, Linga De Gregorio, 1884, Myrtea
Turton, 1822, Thyasira Lamarck, 1818 and so on.
Although the exact reason is unknown, most species ofthese genera have radiating interior striation. The
large-sized N. boucheti, N. maxima and N. gigantea
have such striation while small-sized N. serrei Lamy,
1912 and N. adamsi (Dall, 1889) lack this feature (LyleCampbell, personal communication). Moreover, Reid
(1990) inferred that the large species N. maxima may
harbor sulphide-oxidizing bacteria based on its gutlesscondition pointed out by Kuznetsov & Schileyko(1984). In contrast, Allen & Sanders (1969) describeda normal gut in the small species, N. serrei. In addition,
as above noted, another large species, N. viridis, hasbeen found at the cold seep sites (Okutani & Iwasaki,
2003; Sasaki et al., 2005). Furthermore, all solemyid
species live in symbiosis with sulfur-oxidizing bacteria(Fisher, 1990). The solemyid bivalves first entered coldseep conditions in the Carboniferous (Peckmann et al.,2001) and are widely distributed in post-Carboniferous
hot-vent and cold-seep environments (Campbell, 2006).
Molecular phylogeny of the endosymbiotic bacteria
suggest that the solemyid bivalves harbored thebacteria in ancient times (Imhoff et al., 2003). Basedon these criteria and on the ecology of related species,
we believe that the large size of Nucinella may be
attributed to chemosymbiosis. Conversely, small size offossil and living species of Nucinella may indicate
deposit feeding, as in other protobranchs and shallow
water dwellers. Of course, to settle this hypothesis, it isnecessary to make a direct examination of thesymbiosis on the Recent large Nucinella. N. gigantea
appears to be the earliest Nucinella to invade the cold
seeps, presumably because of its chemosymbiosis.
When he discussed paleobiogeographic history ofEuropean nucinellids, La Perna (2004) pointed out thatthe fossil species, other than the Pleistocene N.
seguenzae (Dall, 1898), lived in shelf environment andthen later migrated to deeper water. The oldest recordof Nucinella is from the early Jurassic of Europe(Vokes, 1956). Based on this record, La Perna (2004)
also postulated that this genus originated in the western
Tethys region. Our new data show that the migrationfrom the western Tethys to the eastern part occurred in
pre-Cretaceous time and the invasion to the deep sea
dates back to the Campanian.
In the northeastern Pacific region, a small species, N.
oregona (Vokes, 1956) has been recorded from the
Nestucca Formation in Oregon whose age is late
middle Eocene (Prothero, 2001). From the middle

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The Veliger, Vol. 49, No. 2

Eocene Cook Mountain Formation, N. alleni Vokes,
1966 was found in the eastern coast of America. By ourdata, it is possible to infer two migration routes of thenortheastern Pacific Nucinella. As the Central American Seaway was present at that time (e.g. Coates &Obando, 1996), it was possible for Nucinella to migratefrom the Atlantic to the Pacific side as did the venerid
genus Securella Parker, 1949 (Vermeij, 2001). Anotherpossibility is the eastward trans-Pacific migration asshown in some cold-water mollusks (Amano, 2005).
The current through the Central American Seaway in
the Eocene (Iturralde-Vinent, 2003) supports theformer alternative. For settling this problem, much
more fossil data of Nucinella are needed especially fromthe North Pacific region.
Acknowledgments. We are very grateful to Prof. G.J. Vermeij(University of California at Davis) and Prof. K. Tanabe(University of Tokyo) for their critical reading of thismanuscript. We thank Dr. R. La Perna (University of Bari),
Dr. Lyle Campbell (University of South Carolina at Spartanburg), Dr. A. Kaim (Institute of Paleobiology, Polish
Academy of Sciences and Department of Earth and Planetary
Science, University of Tokyo) and Dr. T. Sasaki (University
Museum of University of Tokyo) for their information on the
Recent and fossil species. Thanks are extended to Dr. A.
Takahashi (Waseda University) for his information on thegeology and fossils in the Nakagawa area. We also thank Mr.
M. E. Jenkins for his linguistic advice. This study was
financially supported by scientific research grants from the
Fukada Geologocal Institute, the Japan Science Society, andthe 21st Century COE Program of the University of Tokyo(G3, Leader T. Yamagata), and the Japanese Society for the
Promotion of Science (no. 18403013).

LITERATURE CITEDADAMS, A. 1860. On some new genera and species of
Mollusca from Japan. Annals and Magazine of Natural
History (Series 3) 5:299–303.
ADAMS, A. 1862. On some new species of Mollusca fromJapan. Annals and Magazine of Natural History (Series 3)
9:298–299.
ALLEN, J. A. & H. L. SANDERS. 1969. Nucinella serrei Lamy
(Bivalvia: Protobranchia), a monomyarian solemyid andpossible living actinodont. Malacologia 7:381–396.
AMANO, K. 2003. Predatory gastropod drill holes in Upper
Miocene cold-seep bivalves, Hokkaido, Japan. Veliger 46:
90–96.
AMANO, K. 2005. Migration and adaptation of late Cenozoic
cold-water molluscs in the North Pacific. Pp. 127–150 in
A. M. T. Elewa (ed.), Migration of organisms Climate?
Geography?Ecology. Springer-Verlag Berlin Heidelberg.
AMANO, K., S. KANNO, A. ICHIKAWA & Y. YANAGISAWA.
1987. Molluscan fauna from the Tanihama Formation in
the western part of Joetsu City. Studies on the molluscan
fossils from the western part of Joetsu district, Niigata
Prefecture (Part 2). Bulletin of Joetsu University of
Education 6:157–170. [in Japanese with English abstract].
AMANO, K. & T. NONAKA. 2001. A method for estimating
paleobathymetric depth from molluscan community
structure. A case study in the Miocene Mizunami Group

in central Japan. Fossils (Palaeontological Society ofJapan) 70:23–34. [in Japanese with English abstract].
BABA, K. 1990. Molluscan fossil assemblages of the Kazusa
Group, south Kanto, central Japan. Keio Gijuku
Yochisha: Tokyo. 445 pp. [in Japanese].
CAMPBELL, K. A. 2006. Hydrocarbon seep and hydrothermalvent paleoenvironments and paleontology: past developments and future research directions. Palaeogeography
Palaeoclimatology Palaeoecology 232:362–407.
CHINZEI, K. 1959. Molluscan fauna of the Pliocene SannoheGroup of Northeast Honshu, Japan. 1. The faunule of the
Kubo Formation. Journal of the Faculty of Science, theUniversity of Tokyo, sec.2 12:103–132.
CHRONIC, H. 1952. Molluscan fauna from the Permian
Kaibab Formation, Walnut Canyon, Arizona. GeologicalSociety of America Bulletin 63:95–166.
COATES, A. G. & J. A. OBNDO. 1996. The geologic evolutionof the Central American Isthmus. Pp. 21–56 in J. B. C.
Jackson, A. F. Budd & A. G. Coates (eds.), Evolution andenvironment in tropical America. University of Chicago:
Chicago.
DALL, W. H. 1889. A preliminary catalogue of the shellbearing marine mollusks and brachiopods of the southeastern coast of the United States, with illustrations of
may of the species. Proceedings of the United States
National Museum 37:1–221.
DALL, W. H. 1891. On some new or interesting West
American shells obtained from dredgings of the U.S. fish
commission steamer ‘‘Albatross’’ in 1888. Proceedings of
the United States National Museum 14:173–191.
DALL, W. H. 1898. Contributions to the Tertiary fauna of
Florida, with special reference to the silex-beds of Tampa,
and the Pliocene beds of the Caloosahatchie River,including many cases a complete revision of the genericgroups treated of and their American Tertiary species.
Part IV. 1. Prionodesmacea: Nucula to Julia. 2. Teleodesmacea: Teredo to Ervilia. Transactions of the Wagner
Free Institute of Science, Philadelphia 3:571–947.
DALL, W. H. 1901. A synopsis of the Lucinacea and of theAmerican species. Proceedings of the United States
National Museum 23:779–833.
DALL, W. H. 1908. A revision of the Solenomyacidae.
Nautilus 22:1–2.
FISHER, C. R. 1990. Chemoautotrophic and methanotrophic
symbioses in marine invertebrates. Reviews in Aquatic
Sciences 2:399–436.
GIRTY, G. H. 1909. Paleontology of Manzano Group, in W. T.
Lee & G. H. Girty (eds.), The Manzano Group of the Rio
Grande valley, New Mexico. U.S. Geological Survey,
Bulletin 239:1–141.
GREGORIO, A. de. 1884. Studi su talune conchiglie Mediteranee viventi et fossili, con una rivista del gen. Vulsella.
Bulletino della Società Malacologica Italiana 10:36–288.
HABE, T. 1977. Systematics of Mollusca in Japan. Bivalvia
and Scaphopoda. Hokuryukan: Tokyo. 372 pp. [inJapanese].
HAYAMI, I. & T. KASE. 1993. Submarine cave Bivalvia fromthe Ryukyu Islands. University of Tokyo Press: Tokyo.
133 pp.
HICKMAN, C. S. 1984. Composition, structure, ecology, andevolution of six Cenozoic deep-water mollusk communities. Journal of Paleontology 58:1215–1234.
IMHOFF, J. F., H. SAHLING, J. SULING & T. KATH. 2003. 16S
rDNA-based phylogeny of sulphur-oxidising bacterial

K. Amano et al., 2006
endosymbionts in marine bivalves from cold-seep habitats. Marine Ecology Progress Series 249:39–51.
ITURRALDE-VINENT, M. A. 2003. A brief account of theevolution of the Caribbean Seaway: Jurassic to present.
Pp. 386–396 in D. R. Prothero, L. C. Ivany & E. A.
Nesbitt (eds.), From Greenhouse to Icehouse. The marine
Eocene-Oligocene transition. Columbia University Press:
New York.
JENKINS, R. G. 2006. Late Cretaceous marine chemosynthetic
paleoecosystem in northern Hokkaido, Japan, reconstructed from paleontological, stable isotopic and organic
geochemical analyses. Unpublished Ph.D. Thesis, Department of Earth and Planetary Science, The Universityof Tokyo. 78 pp.
JENKINS, R. G., A. KAIM, Y. HIKIDA & K. TANABE. insubmit. Methane flux dependent lateral faunal changes ina Late Cretaceous chemosymbiotic assemblage from the
Nakagawa area of Hokkaido, Japan: Geobiology.
KANIE, Y. & T. NISHIDA. 2000. New species of chemosynthetic bivalves, Vesicomya and Acharax, from theCretaceous deposits of northwestern Hokkaido. Science
Report of Yokosuka City Museum 47:79–84.
KENK, V. C. & B. R. WILSON. 1985. A new mussel (Bivalvia,
Mytilidae) from hydrothermal vents in the Galapagos
Rift zone. Malacologia 26:253–271.
KUZNETSOV, A. P. & A. A. SCHILEYKO. 1984. On the gutless
Protobrachia (Bivalvia). Biologicheskie Nauki 2:39–49. [inRussian with English abstract].
LAMARCK, J. B. P. 1818. Histoire naturelle des Animaux sans
Vertèbres. Vol. 5. 612 pp.
LAMY, E. 1912. Sur le genre Pleurodon ou Nucinella S. Wood,avec description d’une espèce nouvelle. Bulletin deMuséum d’Histoiere Naturelle 18:429–433.
LA PERNA, R. 2004. Nucinella alibrandi (Conti, 1864) and N.
seguenzae (Dall, 1898), the last European nucinellids(Bivalvia, Protobranchia). Rivista Italiana di Paleontologia e Stratigrafia 110:571–577.
LA PERNA, R. 2005. A gigantic deep-sea Nucinellidae from thetropical West Pacific (Bivalvia: Protobranchia). Zootaxa
881:1–10.
LINK, H. F. 1807. Mollusken. In Beschreibung der NaturalienSammlung der Universität zu Rostock, section 2:82–100,
section 3:101–160.
MATSUKUMA, A., T. OKUTANI & R. TSUCHI. 1982. Three new
species of the Nucinellidae (Bivalvia: Protobranchia) from
Pacific coast of Japan. Venus 40:177–186.
MORIYA, K., H. NISHI, H. KAWAHATA, K. TANABE & Y.
TAKATANAGI. 2003. Demersal habitat of Late Cretaceousammonoids: evidence from oxygen isotopes for theCampanian (Late Cretaceous) northwestern Pacific thermal structure. Geology 31:167–170.
NAGAO, T. 1930. On some Cretaceous fossils from the Islandsof Amakusa, Kyushu, Japan. Journal of Faculty of
Science, Hokkaido Imperial University (Series IV) 1:1–25.
NAGAO, T. 1932. Some Cretaceous Mollusca from Japanese
Saghalin and Hokkaido (Lamellibranchia and Gastropoda). Journal of Faculty of Science, Hokkaido Imperial
University (Series IV) 2:23–50.
NAGAO, T. 1938. Some molluscan fossils from the Cretaceous
deposits of Hokkaido and Japanese Saghalien. Journal of
Faculty of Science, Hokkaido Imperial University (SeriesIV) 4:117–142.
OKUTANI, T., E. TSUCHIDA & K. FUJIKURA. 1992. Fivebathyal gastropods living within or near the Calyptogena-

Page 89
community of Hatsushima Islet, Sagami Bay. Venus 51:
137–148.
OKUTANI, T. & N. IWASAKI. 2003. Noteworthy abyssal
mollusks (excluding vesicomyid bivalves) collected fromthe Nankai Trough off Shikoku by the ROV Kaiko of theJapan Marine Science & Technology Center. Venus 62:1–
10.
PARKER, P. 1949. Fossil and recent species of the pelecypod
genera Chione and Securella from the Pacific coast.
Journal of Paleontology 23:577–593.
PECKMANN, J., E. GISHLER, W. OSCHMANN & J. REITNER.
2001. An Early Carboniferous seep community andhydrocarbon-derived carbonates from the Harz Mountains, Germany. Geology 29:271–274.
PECKMANN, J. & V. THIEL. 2004. Carbon cycling at ancientmethane-seeps. Chemical Geology 205:443–467.
PROTHERO, D. R. 2001. Chronostratigraphic calibration of
the Pacific coast Cenozoic: a summary. Pp. 377–394 in
D. R. Prothero (ed.), Magnetic stratigraphy of the Pacificcoast Cenozoic. Pacific Section of SEPM, Book 91,
Tempo Offset Printing: Santa Fe Springs.
POJETA, J., JR. 1988. The origin and Paleozoic diversificationof solemyid pelecypods. New Mexico Bureau of Minesand Mineral Resources Memoir 44:201–222.
REID, R. G. B. 1990. Evolutionary implications of sulphideoxidizing symbioses in bivalves. Pp. 127–140 in B. Morton(ed.), The Bivalvia. Hong Kong University Press: Hong
Kong.
SASAKI, T., T. OKUTANI & K. FUJIKURA. 2005. Molluscs
from hydrothermal vents and cold seeps in Japan: a review
of taxa recorded in twenty recent years (1984–2004).
Venus 64:87–133.
SCOPOLI, G. A. 1777. Introductio ad historiam naturalem
sistens genera lapidum, plantarum et animalium. Prague.
506 pp.
STOLICZKA, F. 1871. Cretaceous fauna of South India. III.
The Pelecypoda, with a review of all known genera of thisclass, fossil and recent. Memoirs of Geological Survey ofIndia, ser. 6 3:223–409.
TAKAHASHI, A., H. HIRANO & T. SATO. 2003. Stratigraphy
and fossil assemblage of the Upper Cretaceous in theTeshionakagawa area, Hokkaido. The Journal of theGeological Society of Japan 109:77–95. [in Japanese withEnglish abstract].
TAYLOR, J. D. & E. A. GLOVER. 2000. Functional anatomy,
chemosymbiosis and evolution of the Lucinidae. Pp. 207–
225 in E. M. Harper, J. D. Taylor & J. A. Crame (eds.),
The evolutionary biology of the Bivalvia. The GeologicalSociety: London.
THIELE, J. & S. JACKEL. 1931. Muscheln der deutschen
Tiefsee-Expedition. Wissenschaftliche Ergebnisse derdeutschen Tiefsee-Expedition auf dem Dampfer ‘‘Valdivia’’ 1898–1899 21:159–268.
TURTON, W. 1822. Conchylia insularum Brittanicarium. The
shells of the British islands, systematically arranged.
Rodwell & Martin: London. 279 pp.
VERMEIJ, G. J. 2001. Community assembly in the sea.
Geologic history of the living shore biota. Pp. 39–60 in
M. D. Bertness, S. D. Gaines & M. E. Hay (eds.), Marine
community ecology. Sinauer Associates, Inc.: Sunderland.
VOKES, H. E. 1956. Notes on the Nucinellidae (Pelecypoda)with description of a new species from the Eocene ofOregon. Journal of Paleontology 30:652–671.
VOKES, H. E. 1966. A new species of the bivalve genus

Page 90
Nucinella from the Eocene of Louisiana. Tulane Studiesof Geology and Paleontology 5:38–40.
VOKES, H. E. 1980. Genera of the Bivalvia: a systematic andbibliographic catalogue (revised and updated). Paleontological Research Institution: Ithaca, New York. 307 pp.
WALLER, T. R. 1978. Morphology, morphoclines and a newclassification of the Pteriomorphia (Mollusca: Bivalvia).
Philosophical Transactions of the Royal Society of
London (B) 284:345–365.
WALLER, T. R. 1990. The evolution of ligament systems in the
Bivalvia. Pp. 49–71 in B. Morton (ed.), The Bivalvia.
Hong Kong University Press: Hong Kong.
WALLER, T. R. 1998. Origin of the molluscan class Bivalvia and a phylogeny of major groups. Pp. 1–45 in

The Veliger, Vol. 49, No. 2
P. A. Johnston & S. J. Gould (eds.), Bivalves: an
eon of evolution. University of Calgary Press: Calgary.
WARÉN, A. & P. BOUCHET. 2001. Gastropoda and Monoplacophora from hydrothermal vents and seeps; new taxa
and records. Veliger 44:116–231.
WOOD, S. V. 1851. A monograph of the Crag Mollusca withdescriptions of shells from the Upper Tertiaries of the
British Isles. Paleontograqphical Society (London), Bivalves 2:1–150.
YABE, H. & T. NAGAO. 1928. Cretaceous fossils from
Hokkaido: Annelida, Gastropoda and Lamellibranchia.
Science Report of Faculty of Science, Tohoku Imperial
University, Ser. 2 (Geology) 9:77–96.