Korean J. Malacol. 35(2): 149-238 2019
https://doi.org/10.9710/kjm.2019.35.2.149
Marine bivalve mollusks of Jeju Island (Korea).
Part 1
Konstantin A. Lutaenko1, Ronald G. Noseworthy2 and Kwang-Sik Choi2
1
A.V. Zhirmunsky National Scientific Center of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences,
Vladivostok 690041, Russia
2
School of Marine Biomedical Sciences, Jeju National University, Jeju-si 690-756,
Republic of Korea
ABSTRACT
The bivalve molluscan fauna of Jeju Island (Jeju-do), the largest island of Korea, is reviewed based on original and
literature data. An annotated and illustrated catalogue of species with indication of local distribution and taxonomic
comments is prepared, supplemented also with data on general distribution, habitats, and primary synonyms. This
first part of the paper includes 105 species belonging to 21 families (Nuculidae through Limidae). The history of
Jeju molluscan research, a bibliography on all aspects of mollusk biology, and the general ecology of Jeju bivalves
in various habitats are also included. Five species are reported as new for Jeju Island, two species as new for
Korea, and 11 species have been reported in literature but were not included in the first bivalve catalogue of Jeju
by Noseworthy et al. (2007); in total 16 species are added to the fauna.
Key words: Bivalvia, fauna, taxonomy, ecology, Korea, Jeju-do
Introduction
lifestyle and customs of its people, tourism is the
region’s most important industry. The island is also
There are about 3000 islands in Korea, located
mostly off the west and south coasts; only a handful of
famous for its subtropical fruits, such as tangerines
(Kwon Y.W. and Lee J.D., 2010).
them lie off the East/Japan Sea coast. Jeju Island
The blending of warm and cool currents around the
(Jeju-do; also previously as Cheju-do) is Korea’s largest
island has resulted in an abundance of marine life.
island with an area of about 1800 km², and it belongs
Jeju Island has a rich mollusk fauna, with over 1000
to the Jeju Special Self-Governing Province (Fig. 1).
species presently recorded. This fauna is a combination
The island is located about 140 kilometers south of the
of warm temperate (Central Japan to the Yellow Sea)
port of Mokpo, in continental Korea, and about 80 km
and tropical and subtropical species (Southern Japan
south of the Korean Peninsula. Due to its volcanic
to Indo-West Pacific) (Noseworthy et al., 2007). The
origin, the island has a topography of numerous hills,
great majority of Jeju mollusks are gastropods,
gorges, and waterfalls. Because of its subtropical
followed by bivalves and the three smaller classes,
climate and the influence of the warm Tsushima
Cephalopoda,
Current, a branch of the Kuroshio, and the unique
respectively.
Polyplacophora,
and
Scaphopoda,
Jeju Island is regarded as a world heritage area by
Received: April 17, 2019; Revised: June 11, 2019; Accepted:
June 26, 2019
Corresponding author: Konstantin A. Lutaenko
e-mail: lutaenko@mail.ru
1225-3480/24734
This is an Open Access Article distributed under the terms
of the Creative Commons Attribution Non-Commercial
License with permits unrestricted non-commercial use,
distribution, and reproducibility in any medium, provided
the original work is properly cited.
UNESCO, which had designated it as a World
Biosphere Reserve, a World Natural Heritage Site, and
a Global Geopark. Biodiversity is one of the most
important components of nature, and all efforts should
be made to protect and manage Jeju’s marine
biodiversity as a Korean and world natural heritage.
We are well aware that biodiversity in marine and
coastal zones worldwide has experienced considerable
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Fig. 1. A map of Jeju and Korea.
pressure, and some marine species have become
extinct, either globally, regionally, or locally, due to
human activity. The main causes of biodiversity loss
are habitat change, climate change, introduction of
invasive exotic species, overexploitation of biological
resources, and pollution. A very important factor
influencing the natural environment of Jeju Island is
tourist pressure: it is estimated that 13 million people
visited the island in 2016. Jeju Island is seeing a
sharp increase in Chinese travelers; nearly half of
Chinese tourists to South Korea visit Jeju.
Understanding
and
conservation
of
biodiversity
requires preparing full inventories, and this is the
Fig. 2. A map of Jeju Island with survey stations: 1.
Dong-gwi, 2. Iho, 3. Jeju-shi, 4. Hwabuk, 5. Samyang, 6.
Hamdeok, 7. Bukchon-ri, 8. Gimnyeong, 9. Wolpyeong,
10. Haengwong, 11. Handong, 12. Sehwa, 13. Hado, 14.
Udo (Sanhosa), 15. Udo (Hagosudong), 16. Udo
(Geomeollae), 17. 5 km east of Seongsan, 18.
Jongdal-ri, 19. Ojo-ri, 20. Seongsan, 21. Seopjikoji, 22.
Shinyang, 23. Pyoseon, 24. Namwon, 25. Wimi, 26.
Bomok, 27. Supseom, 28. 2 km south of Seogwipo, 29.
Munseom, 30. Seogwipo, 31. Seogundo, 32. Bumseom,
33. Bophwan, 34. Gangjeong, 35. Daepo, 36. Jungmun,
37. Yerae, 38. Daepyeong, 39. Hwasoon, 40.
Yeongmeori, 41. Hyungjaeseom, 42. Sagye, 43.
Songaksan, 44. Gapado, 45. Marado, 46. Hamo, 47.
Yongnak, 48. Gosan, 49. Jagwido, 50. Sinchang, 51.
Keumneung, 52. Biyangdo, 53. Hyeopjae, 54. Suwon,
55. Gwideok, 56. Gwakji.
purpose of this paper. Taxonomic inventories are a top
years of research and fieldwork on the mollusks of
priority for biodiversity science. Mollusks are the
Jeju Island. The bivalve fauna was surveyed in order
second largest group in terms of the number of species
to ascertain its distribution and habitat, and update
in marine environments, and they play an important
its taxonomy. Detailed locality lists were prepared
role in communities and ecosystems, including those
which included the number of specimens, living or
unique
to
dead, maturity of specimens, and other relevant
summarize all available data on Jeju bivalve mollusks,
comments. Bivalves were collected from a total of 56
and we believe that this study will be of use not only
general survey stations encompassing the entire coast
for
environment-friendly
of the island (Fig. 2). An attempt was made to sample
education. Since the publication of the above catalogue
as many localities as possible with a wide variety of
(Noseworthy et al., 2007), new species have been
habitats. Some stations were visited on several
added to the fauna, and there have been many
occasions, and many of them contained several specific
changes
This
collecting localities. Therefore the surveys included 87
illustrated catalogue covers the present Jeju bivalve
marine localities. The numbers on the map correspond
fauna and utilizes the current taxonomy.
with those in the list of stations. Specimens collected
to
Jeju
scientists
in
Island.
but
also
nomenclature
We
have
for
and
attempted
systematics.
are kept either at the Shellfish Aquaculture and
Research Laboratory at Jeju National University, Jeju
Materials, Surveys, and Study Methods
City, Republic of Korea (JNU), in the R.G. Noseworthy
This catalogue is the result of more than twenty
collection (RGN collection), a major part of which was
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Korean J. Malacol. 35(2): 149-238 2019
Marine
(Xu and Zhang, 2008), “Mollusks of the Yellow Sea and
Biodiversity Institute, Seocheon, Republic of Korea
Bohai Sea” (Zhang et al., 2016), “Compendium of
(MABIK) and those collected by K.A. Lutaenko, partly
Bivalves” (Huber, 2010, 2015), and “Catalogue of the
at the Zoological Museum, Science and Educational
Living Bivalvia of the Continental Coast of the Sea of
Museum, Far Eastern Federal University, Vladivostok,
Japan (East Sea)” (Lutaenko and Noseworthy, 2012).
Russia
Many
recently
transferred
(ZMFU).
to
Additional
the
National
locality
records
were
research
papers,
taxonomic
revisions
and
obtained mainly from “Mollusks in Korea” (Min D.-K.
reviews, regional guides and, if necessary, those
et al., 2004) and a number of research papers; these
containing original descriptions were consulted, and
records are all listed in the catalogue. Additionally, we
many
list other records, mostly from ecological works
nomenclature,
containing bivalve records.
distribution. The taxonomy of bivalves still lacks
Collecting of bivalves was done mainly from the
intertidal
and
upper
subtidal
zones,
and
beach
driftlines. Numerous taphonomic, actuopaleontological,
species
have
comments
type
material,
with
regard
taxonomy
to
and
agreement, and the taxonomic remarks here will be
helpful for future researchers and students for further
identifications.
All
and faunal studies suggest that dead shells should be
shells
illustrated
in
this
paper
were
included in biodiversity studies, as they generally
photographed in the laboratory of the Zoological
reflect the distribution of live mollusks (e.g., Kidwell
Museum, ZMFU, and all photographs are original.
and Bosence, 1991; Lutaenko, 1994a; b; Lutaenko et
Field photographs were taken by the staff members of
al., 2002; Kidwell, 2008). Live-taken material, after
Shellfish Aquaculture and Research Laboratory, JNU,
identification, was stored in 70% ethanol. Samples of
unless otherwise stated in the captions.
We
beach drift, rock brushings, and detritus obtained from
follow
mainly
the
higher
classification
of
depths of 0.5 to two meters were also sorted, and the
Bivalvia by Carter et al. (2011). Habitat data are
micromolluscs extracted and identified with the help of
mostly obtained from Higo et al. (1999) and Huber
a Zeiss dissecting microscope and a handheld 50x
(2010; 2015, including supplementary CD files). Our
magnifier.
original data on Jeju habitats are included and other
In 2012, we identified a large number of mostly
works were also consulted.
live-taken mollusk specimens obtained by SCUBA on
behalf of ECO21, an environmental survey company;
Format
all species, including bivalves, identified from many
locations around the island, were listed. Also, a survey
The entries in this catalogue follow a specific format.
was conducted by the Shellfish Aquaculture and
For each species, major synonyms (names proposed as
Research Laboratory, JNU, from 2012 to 2014, on the
new for science) are given, followed by the local and
distribution of the encrusting coral Alveopora japonica
general distribution ranges, habitat, and comments.
Eguchi, 1968, which was rapidly extending its range
Specific Jeju survey stations are listed. Other records,
around the island. Coral specimens were obtained by
either additional or from the same localities, are
SCUBA diving from several localities, at depth of 5-15
provided in the “other records” local distribution
m.
section.
Mollusks,
mainly
bivalves,
were
sorted
and
identified, then preserved for further study.
Zonal-geographical groupings are included to show
Identification of specimens from the field surveys, as
more clearly the biogeographical affinities of each
well as from those above, was facilitated by several
species. For each species we give zonal-geographical
important recent books, mainly “Marine Mollusks in
characteristics. For information about this method and
Japan” (Okutani, 2000), “Mollusks in Korea” (Min
a clarification of terms see: Scarlato (1981) and
D.-K. et al., 2004), “Seashells of China” (Qi, 2004), “An
Kussakin (1990). We recognize six major zonal-
Illustrated Bivalvia Mollusca Fauna of China Seas”
geographical groups: 1. tropical-subtropical (distributed
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
southward to the Philippines, Vietnam, and Indonesia),
Hallasan. While drainage systems created from wide
2. subtropical (distributed southward to Taiwan and
lava plateaus formed on the gentle east-west slope of
the northern part of the South China Sea) 3.
Hallasan are less evolved, most watercourses were
subtropical-lowboreal (limited both to subtropical seas
developed on the north-south slopes and run either
and the Sea of Japan/East Sea, southeastern Sakhalin,
southbound or northbound (Woo K.-S. et al., 2013).
and the southern Kuril Islands, 4. lowboreal (limited
With the exception of the time during typhoons and
to the Sea of Japan/East Sea from Peter the Great
summer monsoons, most of the flowing water in Jeju’s
Bay,
to
rivers and streams is short-lived, and there is little
southwestern Sakhalin, Aniva and Terpenya Bays, and
standing water in the form of ponds because of the
northern
Korea,
and
northern
Honshu
the southern Kurile Islands), 5. widely distributed
porous nature of the extensive basaltic areas. Water
boreal (limited to the Sea of Japan/East Sea and
from upper streams runs into the
Hokkaido to Bering Strait, along the Asian coast, and
through permeable layers developed on the edges of
along the northern American coast southward to
lava flows or along columnar joints in the streambeds
California),
underground
to
(Woo K.-S. et al., 2013). However, although dry
temperate latitudes, both in the Atlantic and Pacific
riverbeds are often seen, flowing rivers are more
Oceans, but also partly to subtropical and arctic
common along the coast because the water usually
zones), and 6. boreal-arctic (limited to both the
rises in the lower reaches of the channels; the
temperate zone of the Pacific Ocean and the Arctic,
Seogwipo region in the south usually has waterfalls.
and partly to the temperate Atlantic). We checked the
Rivers in Jeju Island are usually small, less that 26
geographical distributions of
km (16 miles) long, and their estuaries are not usually
and
circumboreal
limited
mainly
many species using
heavily polluted or directed into channels. Our field
numerous literature sources not cited herein.
on
surveys have revealed that the rather small estuaries
taxonomy, distinguishing characterizes, location of
exert little influence on the salinity of nearby
types, and other relevant information. We use only
seawater,
subclasses, orders, superfamilies, families, subfamilies,
Crassostrea gigas (Thunberg, 1793) commonly occur in
genera, and subgenera in the taxonomic arrangement.
those areas.
Comments
include,
if
necessary,
remarks
although
estuarine
species
such
as
Although there are several small estuaries, mainly
on the south coast, there is only one lagoon on Jeju
Brief Regional Setting
Island, at Ojo-ri, on the east coast near Seongsan
Jeju Island, 33°10′-33°50′N, 126°10′-127°0′E, is
Ilchulbong
(Sunrise
Peak),
(33°27′27.18″N,
an
extinct
secondary
a volcanic island, elliptical in shape, created mainly
volcano
from eruptions of Hallasan, a shield volcano with a
lagoon is almost flat and shallow, and protected from
height of 1950 m, from about 1.2 million years ago
wave action by the volcano which forms a cape. This
until historic times; the eruptions also caused more
lagoon has mainly a coarse sandy mud substrate with
than 360 small parasitic volcanoes, called oreum
patches of small boulders (Lee H.-J. et al., 2014;
(Kwon S., 2006). The east and northeast coasts face
Silina, 2014).
126°56′00.39″E).
This
the Korea/Tsushima Strait; the south coast faces the
Sandy beaches of several types are found on all
East China Sea; and the west and northwest coasts
coasts of Jeju Island (Figs. 3, 4). The beach sands are
face the Yellow Sea. The coastline is mainly rocky
mainly of three types: silicate sand, carbonate sand,
with several sandy beaches and a few sand tidal flats.
and
Mud flats are virtually nonexistent (Choi K-S. et al.,
carbonate materials (Kim T.-H. et al., 2008). Silicate
2000).
sand is formed from volcanic rocks by weathering
mixed
sands
containing
both
silicate
and
Jeju’s drainage system is made up of streams that
caused by strong wind and waves, and feldspar and
radiate outwards from the central high-point of
quartz are the main elements; this type is found at
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Korean J. Malacol. 35(2): 149-238 2019
Fig. 3. Coastline and beaches of Jeju Island: A. Seongsan (2007); B. Beach thanatocoenoses at Gwakji
(2011); C. Jungmun (2011); D. Yeongmeori (2013); E. Jongdal-ri (2011); F. Hyopjae (2011). All
photographs by K.A. Lutaenko.
Samyang, Seongsan, Hamo (Sangmo), and Gummulae
sands. Mixed sands are found at Jongdal-ri and
at
Jungmun areas, as well as other beaches.
Udo,
a
small
island
adjacent
to
Seongsan.
Carbonate sand is formed from crushed mollusk shells
Most Jeju beaches are carbonate sand and the
and skeletons of coralline algae caused by wave action
remainder are an equal number (6) of mixed sand and
and is found in Hamdok, Gymnyeong, and Hyupjae, as
silicate sand. Most beaches (9) are found on the north
well as other beaches. Calcite and analcime are the
and east coasts, with an equal number (5) found on
principal minerals of the carbonate sand. Mixed sands,
the south and west coasts (Table 1). Two of the most
the third type, contain both silicate and carbonate
speciose
- 153 -
beaches,
Yeongmeori
and
Hwasoon,
are
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Fig. 4. Coastline and beaches of Jeju Island: A. Jungmun (2007); B. Yerae (2012); C. Pyeoson (2007); D.
North coast (2007); E. Seongsan (2007); F. Tongbatarl (Ojo-ri) Lagoon (2007). All photographs by
K.A. Lutaenko.
located on the south coast, and a third, Seongsan, on
development.
the east coast (Figs. 3, 4). Another rich area is Yerae,
Apart from the sandy beaches found on all coasts of
a coastline with deep tidepools and patches of sand, on
the island, most of the coastline is composed of
the south coast. The highest number of mollusk
volcanic rock of various types (Figs. 3, 4). Steep cliffs
species, 176, of which 73 are bivalves, has been
and headlands are found at various locations, and
reported from Hwasoon; however this area has now
cliffs of columnar basalt occur on the south coast. Also,
been significantly modified by port and tourism
low-lying coasts, such as the north coast, may be
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Korean J. Malacol. 35(2): 149-238 2019
Tabel 1. Beach sediments in studied localities of Jeju Island
Coast
Beach
Sand type
North
Gwakji
Carbonate
Gwakji (Handam)
Carbonate
East
Udo
South
West
Iho
Mixed
Samyang
Silicate
Hamdeok
Carbonate
Gimnyeong
Carbonate
Woljeong
Carbonate
Haengwon
Carbonate
Handong
Carbonate
Sehwa
Carbonate
Hado
Carbonate
Jongdal
Carbonate
Songsan
Silicate
Shinyang
Mixed
Pyoseon
Carbonate
Hagosudong
Carbonate
Sanhosa
Coralline algae (carbonate)
Geomeollae
Silicate
Beomeok
Cobble and coarse sand (silicate)
Jungmun
Mixed
Hwasoon
Mixed
Yeongmeori
Silicate
Sagyei
Mixed
Hamo (Sangmo)
Silicate
Hamo
Mixed
Geumneung
Carbonate
Hyeopjae
Carbonate
Gwideok
Carbonate
composed of jagged basalt rocks interspersed with
Beardsley, 2002). The Changjiang (Yangtze) River, the
tidepools, and projecting rocky ledges. The coastline
longest river in Asia, drains into the East China Sea;
may also consist of wide lava flows, such as at Yerae,
this drainage water, having moved southward from the
on the south coast, and boulder shores, also with
Yangtze River mouth, joins the Kuroshio Current, and
tidepools, such as those at Seogundo, on the south
is carried by the Kuroshio Branch Current to Jeju
coast, and Suwon, on the northwest coast. Offshore,
Island and the Korea/Tsushima Strait. In winter, the
however, the sea bottom is largely composed of muddy
Tsushima Current has a single source, the Kuroshio
sand, with areas of sandy mud near the north coast
Branch Current,
(Je J.-G. et al., 1991).
Kuroshio Current water and Yangtze River drainage
The Tsushima Current flows past Jeju Island
which transports a mixture of
water northward. In summer, the Tsushima Current,
through the Korea/Tsushima Strait. The northward
besides
current flowing west of Kyushu, southwestern Japan,
influenced by the Taiwan Warm Current, flowing from
called the Kuroshio Branch Current, is considered the
the south (Ishikawa and Beardsley, 2002).
origin of the Tsushima Warm Current (Ishikawa and
- 155 -
the
Kuroshio
Branch
Current,
is
also
Although cooler water currents, particularly from
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
the eastern area of the Yellow Sea flowing past the
western
Korean
coast,
exert
some
influence,
been approximately 6 mm, which is about three times
the global average (Kim Y. et al., 2012). Furthermore,
particularly to the north of Jeju Island, the warm
Yamano
Tsushima Current predominates, giving the island a
temperature
subtropical climate with mild winters and hot, humid
including Jeju Island, has risen by 1.2° C in the past
summers, and an average yearly temperature of 16°C
100 years.
et
al.
in
[2011]
the
demonstrated
southern
Yellow
that
Sea
sea
area,
(World Climates, 2015). The prevailing winds from the
Pacific Ocean are southeasterly in summer, and also
Previous Research on Bivalve Mollusks
help to raise the temperature of the water. The water
temperature is highest in August (26.1°C) and the
The Korean fauna received little attention from
lowest in February or March (13.5°C) (World Sea
malacologists in the nineteenth and the first half of
Temperature, 2015). Monthly mean temperature and
the twentieth century. In the middle of the nineteenth
salinity of sea water around Jeju Island is influenced
century some mollusk collections from Korean waters
by precipitation and weather conditions as well as by
were made during the visit of the British Navy vessel
oceanographic conditions of the open sea such as the
H.M.S. Samarang. Arthur Adams, a malacologist and
Tsushima Warm Current and sea water in coastal
assistant surgeon on the Samarang, supervised these
areas (Ko J.-C. et al., 2003). Salinity is usually over 3
collections, and they were described in the “Zoology of
4‰ with a tiny variation between December and June,
the Voyage of H.M.S. Samarang” (Adams and Reeve,
but large fluctuations occur during summer, and
1848-1850). Five bivalve species described here also
salinity may decrease to 29-30‰ or even to 23.94‰
occur in Jeju Island, and several others described
(Sinsan, September) or 25.65‰ (Jungmun, July) in
elsewhere are also listed. Adams, in his introduction to
shallow-water stations. The place of the lowest water
the mollusk section of the above publication, stated:
temperature was the north-eastern coast of Jeju: in
“In Quelpart [Jeju Island], for instance, where the
February, the minimum temperature goes down as low
perpendicularity of the sides of this deep-seated
as 11.0°C at Hado and in March, 12.1°C at Sehwa.
submarine mountain will not allow of the formation of
The maximum water temperature can increase in
coral, few shells are found” (l.c., p. ix). He also
August up to 27.7°C at Hwasoon and 26.1°C at
described a lucinid species (Myrtea delicatula A.
Jungmun, and in September up to 26.6° in Jungmun
Adams, 1862) from Korea Strait (Adams, 1862), and
(Ko J.-C. et al., 2003).
later described several bivalve species with type
Jeju Island has moderately wide tidal ranges, with
localities as Quelpart, e.g., Cuspidaria nobilis (A.
an average of about three meters; the south coast
Adams, 1864) (as Neaera), Cuspidariidae (“Quelpart,
experiences slightly wider ranges than the north coast.
52 fathoms”) and Callocardia guttata A. Adams, 1864,
Off the south coast the tidal current sets in an east
Veneridae (“off Quelpart, 48 fathoms”) (Adams, 1864a;
and west direction, roughly parallel to the coast, at a
b). Thus, the history of mollusk collecting in Jeju has
velocity of 0.5 to 0.75 knots. Tidal currents off the
continued for more than 160 years.
north coast of the island set parallel to the coast.
Jeffreys (1879) described a collection of bivalves,
Climate change caused by global warming is a
gastropods, and brachiopods, without illustrations,
strong possibility for the addition of new bivalve
from Korea Strait, between Jeju Island and the
species to Jeju Island’s fauna. There has been a 1.6°C
southern Korean mainland. However, his list was
increase in average winter temperatures since the
based on the erroneous idea that a number of species
1930s. As a result, both the depth of snow at higher
from Japan or Korea were the same as those in
elevations and the total number of days of snowfall
Europe: “certain species ... are identical or varieties of
have been moving downwards. Since 1960, the average
European species” (l.c., p. 418). Although it would be
annual rate of sea-level rise around Jeju Island has
most interesting to ascertain the Korean species to
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Korean J. Malacol. 35(2): 149-238 2019
which Jeffreys was referring, it would be extremely
Munseom and Jigwido. These authors compiled a list
difficult, if not impossible, to do so without access to
of all species found around Jeju Island, including those
the actual specimens.
from previous works, and among them were 123
In the first half of the twentieth century Japanese
bivalve species.
malacologists published several lists of the Korean
In a series of papers, J.J. Lee and co-workers
fauna (see for details: Lutaenko and Noseworthy
published important data on the species composition
(2012), but they dealt mostly with continental Korea.
and distribution of intertidal benthic invertebrates in
The earliest Korean papers documenting some bivalve
many parts of Jeju Island. A study of 25 intertidal
mollusks from Jeju Island were by B.D. Lee (1956;
stations around the island in 1987-1989 listed 13
1958), a pioneer of Korean malacology. He mentioned
species of bivalves (Lee J.J. and Jwa Y.W., 1988; Lee
“specimens from Isl. Quelpart” for several mollusk
J.J. et al., 1989). In the northern part of Jeju Island,
species (Lee B.D., 1958). H.S. Kim and B.J. Rho (1971)
at Hamdeok, Oedo, and Guiduck, 13 species of bivalves
published the first check-list of Jeju Island benthic
were found (Lee J.J., 1990). At 8 stations around
animals based on literature records and field studies
Seogwipo, on the southern coast (August 1990-July
in 1970-1971. They enumerated only 10 bivalve
1991), a study on the distribution and community
species,
mostly
from
Seogwipo
but
also
from
structure of benthic mollusks was conducted, with 8
bivalve species collected; among them, two bivalves
Seongsanpo, Moseulpo, and Jungmun.
J.-G. Je et al. (1991) sampled benthic mollusks in
were dominant species in the middle and lower
the soft substrate of the Southern Sea of Korea (Korea
intertidal zones (Lee J.J. and Hyun J.M., 1991). A
Strait and East China Sea) at 83 stations in July and
similar study in the intertidal zone of Gapa and Mara
August, 1990, including about ten station around Jeju
islets (March-June 1991) revealed 11 bivalve species
Island. At least, three species of dominant bivalves are
(Lee J.J., 1991). From October 1991 to October 1992,
shown on maps as living around the island. A total of
the
33 bivalve species were found in this area; however,
community on rocky shores were studied in the
the list is provided for the entire Southern Sea, and
eastern part of the island, at Shinyang, Ohcho (Ojo-ri),
Jeju coastal waters cannot be separated. Photographs
Jongdal, and Hado, and also around Udo Island, with
of many species collected in this and other areas of
12 species reported (Lee J.J. and Hyun J.M., 1992).
Korea are given by J.-G. Je (1993).
Lee J.J. and Kim J.C. (1993) studied the distribution
seasonal
changes
in
the
intertidal
benthic
Kim H.S. et al. (1992) published a report on bivalve
and seasonal changes of the benthic macroinvertebrate
mollusks of Jeju enumerating 22 species (9 identified
community in four intertidal rocky shore areas of
to genus level only) and illustrating Arca avellana
Biyangdo from January 1992 to January 1993; they
Lamarck, 1819, Septifer keenae Nomura, 1936, Limaria
collected 10 bivalve species. In the upwelling area on
basilanica (Adams et Reeve, 1850), Crassostrea gigas
the western coast, around Jagwi-do, 10 bivalves were
(Thunberg, 1793) and Ruditapes variegatus (G.B.
identified, and one mytilid species was found among
Sowerby II, 1852) (= Ruditapes aspera (Quoy et
the dominant fauna (Lee J.J. et al., 1995). A study on
Gaimard, 1835)).
the distribution and seasonal changes of the benthic
J.-G. Je et al. (2002) conducted a baseline study for
invertebrate community, performed on intertidal rocky
conserving the coastal and marine biological diversity
shore of Chuja-do in 1996-1997, revealed 15 species of
of Jeju Island. Coastal habitats, and zoobenthic species
bivalves, and among them S. keenae was a dominant
composition and distribution, were investigated for
species in the middle and sub-middle intertidal zones
three years (1999-2000) in 18 intertidal areas along
(Lee J.-J. and Hyun J.-M., 1997). A study of species
the coast of the island. They sampled four subtidal
diversity
rocky shores and three subtidal sandy substrate areas
(September 1999-August 2000) revealed 10 bivalve
near Seogwipo, including the islets of Beomseom,
species (Lee J.J. et al., 2001). J.J. Lee and J.M. Hyun
- 157 -
in
the
Hwasoon
rocky
intertidal
area
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
(2002) conducted a study on species diversity, vertical
intertidal and subtidal bottom communities are given
distribution, and community structure of intertidal
in the book “Current and Future Living Resources”
invertebrates
near
(2015) published by Korea Institute of Ocean Science
Songacksan, on the southern coast, from June 2000 to
and Technology (KIOST). Another book, a field guide
May 2001; in this area, they recorded 12 bivalve
to the Jeju seashore animals and plants, features 21
species.
species of bivalves with color illustrations (Yang H.-S.
at
nine
sampling
stations
The biodiversity of benthic invertebrates in the
and Choi K.S., 2011).
intertidal and subtidal regions of Gapado, Beomseom,
There are a number of regional ecofaunal molluscan
and Munseom islets was surveyed twice, in May and
studies in Jeju. J.-G. Je et al. (1994a) studied
September 2013, to study the state of biodiversity in
composition and distribution patterns on rocky shores
Seogwipo; as a result, a total of 29 species of bivalves
in the Munseom area, in June-July 1993; a total of 77
were found (Cho I.-Y. et al., 2014). A report on the
bivalve species were identified, and the authors
distribution of common species of benthic animals in
discussed the zoogeography of this fauna. A special
the Munseom area was published by Je J.-G. et al.
study was devoted to the molluscan fauna of the
(1994b).
Seongsan area in eastern Jeju; 17 species were
K.-S. Choi (1996) and K.-S. Choi et al. (2000)
collected and illustrated (Noseworthy et al., 2002).
surveyed 25 of the island’s edible bivalves to ascertain
Surveys of Seogundo, a small island adjacent to the
their
species
southern coast of Jeju and connected by a boulder
belonged to the Veneridae, Mytilidae, Pectinidae, and
beach at low tide, were conducted from 2001-2009 to
Ostreidae. The majority of these species occur in the
enumerate the mollusks, and examine their diversity,
subtidal area, and harvesting wild populations for
relative abundance, and ecological relationships; in
commercial purposes would require special techniques
total, 16 bivalves were found (Noseworthy and Choi,
and equipment. However, because of the relatively
2010). Suzuki (2010) collected 26 species of bivalves
small populations of edible species and the exposed,
from beach drift at 11 localities in Jeju Island. The
mainly rocky, shoreline, no commercial bivalve mollusk
molluscan fauna of the Tongbatarl (Ojo-ri) Lagoon
commercial
potential.
Most
of
the
aquaculture may be possible in Jeju Island.
near Seongsan was studied by H.-J. Lee et al. (2014)
H. Qi and B.L. Choe (2000) published a report on
in July, 2012; a total of 9 bivalve species were found,
five unrecorded ostreoidean species in Korea with four
the Manila clam (Ruditapes philippinarum (Adams et
new species records from Jeju Island. K.-S. Choi (2008)
Reeve, 1850)) population having low density, and the
surveyed the Gryphaeidae and Ostreidae of Korea,
lucinid Pillucina pisidium (Dunker, 1860) high density.
showing their distribution, value in aquaculture, and
Noseworthy et al. (2016a) studied an assemblage of
type of aquaculture employed for some species.
mollusks
Since the 1970s, Korean malacologists published
several
identification
guidebooks
with
associated
with
the
high
latitude
scleractinian coral Alveopora japonica (Eguchi, 1968)
color
in Jeju, and identified 20 species of bivalve mollusks.
photographs of mollusks (Yoo, 1976; Kwon O.-K. et al.,
Noseworthy and K.-S. Choi (2018) collected mollusk
1993; 2001; Min D.-K. et al., 2004), also containing
species associated with the coralline algae Corallina
records of bivalves from Jeju Island. The most
confusa Yendo, 1902 from a tide pool at Hamo beach
important is the above-mentioned book “Mollusks in
on the south-west coast of Jeju Island in May 2012,
Korea” (Min D.-K. et al., 2004). In two reviews of the
and a total of 24 species of mollusks were identified,
Korean Arcoida and Mytiloida (Lee J.-S., 2013) and
including 4 bivalves.
Solemyoida and Nuculida (Lee J.-S., 2014), many
Special reports deal with possible invasive bivalves
species records for Jeju are enumerated. Images of 7
have appeared in Jeju recently. Oh et al. (2017)
common bivalve species living around Busan and Jeju,
reported the first finding of the tropical blacklip pearl
along with data on distribution and schemes of the
oyster Pinctada margaritifera (L., 1758) (Pteriidae) in
- 158 -
Korean J. Malacol. 35(2): 149-238 2019
Jeju
(Beomseom)
and
suggested
that
intensive
Limpanont et al. (2011) reported on the annual
monitoring surveys are necessary for confirming the
reproductive
northward expansion of this species in Korean waters
reproduction, and larval morphology of the tellinid
cycle,
environmental
influences
on
in the near future. A malleid, Malleus regula (Forsskål
Macoma irus (Hanley, 1845). Kang D.-H. et al. (2007)
in Niebuhr, 1775) was recorded for the first time from
reported
Korea, being the second species of this family found in
gametogenesis in Coecella chinensis (Deshayes, 1855).
this country; coastal warming and possible changes in
The relatively smaller eggs observed were believed to
for
the
first
time
on
the
annual
the northward-flowing Tsushima Current may account
be a reproductive strategy of C. chinensis, an adaptive
for the addition of new mollusk species to the island’s
trade-off between reproduction and growth in the
fauna (Noseworthy et al., 2016b).
food-poor upper intertidal environment. Limpanont et
The first detailed check-list of all mollusks of Jeju
al. (2010) also issued a first report on the annual
Island was published by Noseworthy et al. (2007)
reproductive cycle of Burchard's cockle, Acrosterigma
based on a four-year survey and literature records.
burchardi (Dunker, 1877).
Forty-eight survey stations were visited, with a total of
The commercially important venerid R. philippinarum
82 specific localities. The list contained 225 marine
was the object of several Jeju Island studies. Ngo and
bivalves, with the Veneridae, Mytilidae, and Arcidae
K.-S. Choi (2004) investigated seasonal changes in the
having the largest number of species, and also data on
infection intensity of the parasites Perkinsus and
local and world-wide distributions. This check-list is a
Cercaria in R. philippinarum. This study clearly
basis for the current paper.
showed that parasitism impacts clam reproduction, at
The above studies have contributed much to the
least during part of the annual reproductive cycle.
understanding of the species composition of the
Also, Uddin et al. (2013) investigated the gonad
molluscan fauna of Jeju Island specifically and also
development and reproductive effort of this species.
the Korean Peninsula. Since then, studies of this
Size and age structures of R. philippinarum from
fauna have continued.
coastal subrecent shell accumulations derived from
There are a number of studies on the biology,
populations inhabiting the east coast of Jeju Island
ecology, age and growth, reproduction, and diseases of
were studied by Silina (2009; 2011). Silina (2014) also
bivalve
studied
mollusks
inhabiting
Jeju
coastal
waters:
the
growth
and
some
morphological
Pinctada fucata (Gould, 1850) (= Pinctada imbricata
parameters of Ruditapes bruguieri (Hanley, 1845) (= R.
Röding,
aspera) as well as its habitat preferences in Jeju
1798)
(Cho
C.-H.
et
al.,
1988),
Ostrea
circumpicta Pilsbry, 1904 (= Striostrea circumpicta)
(Kang
D.-H.
2004),
Island.
japonicum
On the coast of Jeju, marine Holocene deposits with
(Gmelin, 1791) (= Ylistrum japonicum) (Son P.-W. et
molluscan shells are distributed, with the youngest
al., 1996). N.L. Lim (2005) reported for the first time
stratigraphic units being the Sinyangri and Hamori
on the microscopic features of larvae and gonads of S.
formations; however, the molluscan shells are mostly
circumpicta. H.-K. Hong et al. (2013) investigated the
bivalve fragments which cannot be identified to species
types of hemocytes and immune activity of the oysters
level; their ages are mid-Holocene (3400-5000 years
Saccostrea
et
kegaki
al.,
Torigoe
Amusium
and
Inaba,
1981,
S.
BP) (Cheong C.-S. et al., 2006). We believe that some
circumpicta, and Hyotissa hyotis (L., 1758). Lee H.-J.
of the subfossil shells collected by us from beaches
et al. (2013) made first observation on the early
(e.g., Anadara granosa (L., 1758) may belong to a
embryonic and larval development of S. kegaki. Their
re-deposited Holocene fauna washed ashore. Another
results provide the basic information concerning the
fossil molluscan fauna in Jeju is more ancient and
hemocyte populations of those three subtropical oysters
belongs to the Seogwipo Formation (Late Pliocene-
for
Early Pleistocene) (Yoon S., 1988; Kang S.-S., 1995).
further
investigation
associated
with
various
environmental disease stresses. In the Heterodonta,
- 159 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Fig. 5. Jeju bivalves in natural habitats: A, C, D. Clumps of Septifer virgatus in the intertidal zone; B.
Lithophaga lischkei: specimens and boreholes; E, F. Clumps of Septifer keenae in the intertidal zone;
G, H. Clumps of Brachidontes mutabilis in the intertidal zone.
- 160 -
Korean J. Malacol. 35(2): 149-238 2019
Ecology of Jeju Bivalves
sand with a slight admixture of silt. The silt content in
the bottom sediments may constitute 1-2%. It indicates
Most of the Jeju coastline is of volcanic origin,
mainly basalt and lava flows. A wide variety of
that the species is, most likely, sensitive to mud
resuspension in the water (Silina, 2014).
bivalves inhabit this area, and the dominant bivalve
At Seongsan, the sandy beach is behind the slabs,
species vary from one area to another. Mytilids are
and along their seaward edge are other sandy areas.
rarely found in some areas, such as Seogundo, but A.
Three species of Veneridae and single species of
avellana and Cardita leana Dunker, 1860 inhabit
Mactridae and Psammobiidae are moderately common
crevices in the undersides of rocks in shallow water, as
in those areas. Besides sandy beaches, bivalves may
well as other species, such as Striarca symmetrica
also inhabit other areas of soft substrate, such as
(Reeve, 1844), Irus irus (L., 1758), and Anomia
sandy areas at the bottom of large tidepools. Samples
cytaeum Gray, 1850. Clumps of Septifer virgatus
of coarse sand from the bottoms of tidepools at
(Wiegman, 1837) and S. keenae (Fig. 5) occur byssally
Seogundo yielded eight bivalves: Nucula paulula A.
attached between rocks; C. leana and Brachidontes
Adams, 1856, Musculus viridulus (H. Adams, 1871), B.
mutabilis (Gould, 1861) (Fig. 5) are frequently found
mutabilis, A. avellana, Barbatia trapezina (Lamarck,
among the clumps.
1819), Barbatia stearnsi (Pilsbry, 1895), R. aspera, and
At Seongsan Ilchulbong, the shoreline is composed of
Irus irus (L., 1758). M. viridulus, which prefers rocky
flat slabs of basalt which are exposed at low tide. The
substrates (Okutani, 2000; as “Musculus nanus”) and
rock slabs are several meters in width from a coarse
marine
sandy beach to the water’s edge, and contain several
(Noseworthy and Choi, 2010). The pholadid Barnea
tide pools. A. avellana and S. keenae are found exposed
manilensis (Philippi, 1847) inhabits hard sandy mud at
on the rocks, and in tide pools and crevices. At the
Hamo Beach.
flora,
was
also
found
in
this
habitat
eastern end of the Hwasoon area, Barbatia bistrigata
A few sand tidal flats occur on the east coast of Jeju
(Dunker, 1866) is one of the dominant species in the
Island, mainly at Jongdal-ri, Ojo-ri, and Hado-ri. A
intertidal zone of the rocky coastline (Lee J.J. et al.,
macrobenthic fauna survey at Jongdal-ri revealed that
2001); while in the Seogwipo area S. keenae is the
one of the dominant species at this locality was
dominant bivalve in the lower intertidal zone (Lee J.J.
Cycladicama
and Hyun J.M., 1991).
cf.
lunaris
(Yokoyama,
1927).
Other
common bivalves were Solen strictus Gould, 1861, R.
Bivalves colonize all hard substrates, including
philippinarum, Gomphina multifaria (Kong, Matsukuma
artificial ones such as breakwaters and wharves which
et Lutaenko, 2012), and Nuttallia obscurata (Reeve,
present a smoother surface for encrusting organisms.
1857) (Fig. 7). Other bivalves occurring in this area
A survey of intertidal invertebrates inhabiting natural
were mainly infaunal species, one each of eleven
and artificial substrata on the northeast coast of the
families, and two species of Carditidae (Ecological
island revealed that the ostreids C. gigas and S.
Studies of Jongdal-ri…, 2005).
kegaki (Fig. 6), and the mytilid S. virgatus, are
Marine flora, seaweed and coralline algae, constitute
dominant species on natural substrata; however, only
a distinct mollusk habitat. Although gastropod species
S. virgatus is the dominant species on artificial
make up the largest number of mollusks, bivalves,
substrata (Cha J.-H. et al., 2013).
usually smaller sessile filter feeders, are often found
Although rocky coastlines predominate in Jeju
attached to various species of seaweed and coralline
Island, several beaches of coarse and fine sand, and a
algae, and some may be dominant. This habitat
few muddy-sand areas, are distributed along all coasts
provides both some protection from predators and a
of the island. Bivalves occur mainly in the subtidal
source of food. Organic detritus can collect in and
zone,
For
around the algae and provide a ready food supply for
example, R. aspera prefers coarse and medium-grained
the attached bivalves. For example, smaller species of
although some
inhabit
the intertidal.
- 161 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Fig. 6. Jeju bivalves in natural habitats: A, B. Mytilus coruscus. C. Musculus pusio in seaweed Chondrus
crispus; D. Specimens of Saccostrea kegaki in the intertidal zone; E. Coecella chinensis; F. Pinctada
margaritifera; G. Macoma irus; H. Irus mitis.
- 162 -
Korean J. Malacol. 35(2): 149-238 2019
Fig. 7. Jeju bivalves in natural habitats: A-C. Ruditapes philippinarum; D, F. Solen strictus; E. Cyclina
sinensis.
Mytilidae and Arcidae may be found in species of red
dominant mollusk species there (Fig. 6).
and brown algae, e.g., Sargassum sp., and also
Coralline algae appear to contain the largest number
coralline algae. Species of Musculus, as well as some
of mollusk species. The number of bivalve specimens of
small arcids, are often found byssally attached to the
small species or juveniles and subadults of the larger
algae.
at
species may vary among different localities. For
Seogundo on the south coast (Noseworthy and Choi,
example, a study of mollusks associated with coralline
2010) revealed that, although only three bivalve
algae at Hamo Beach, on the southwestern coast of the
A
study
of
mollusk-algae
associations
species occurred live in four different algae species, the
island revealed that some species, e.g., M. viridulus,
mytilid M. pusio was not only the most common live
that were very common in the Seogundo survey were
bivalve obtained in this survey but was also one of the
quite scarce at Hamo. Most bivalves obtained so far
- 163 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
have been filter feeders; however, N. paulula, a deposit
coral communities. J.-C. Ko et al. (2008), in their study
feeder, has also been obtained from the coralline algae
of the macrobenthos of Munseom, an islet off the
Alatocladia cf. modesta Johansen, 1969 (Noseworthy
southern coast of the island, listed the dominant soft
and Choi, 2010). It is possible that organic detritus
coral species but provided no specific information on
may be trapped in the blades of the algae and become
bivalve associations. I.-Y. Cho et al. (2014) studied the
avalaibale for deposit-feeding bivalves.
biodiversity of benthic invertebrates in the Seogwipo
At Yeongmeori coast, approximately 2 km west
area, which is rich in soft coral species, especially the
southwest of Hwasoon, south coast of Jeju Island, a
subtidal zones around Munseom and the neighboring
large sample of coralline algae and brown seaweed
islet
washed up at the water’s edge by heavy wave action,
occurring in this area were listed, along with 14
at very low tide, yielded a wide variety of bivalves,
bivalve families associated with the coral. Of those, the
mainly seven species of Mytilidae: four small species,
Mytilidae, Arcidae, and Chamidae were the most
M. viridulus, Musculus pusio (A. Adams, 1862), B.
speciose with five species each.
juvenile
specimens
of
three
larger
Beomseom.
Six
protected
coral
species
There are only a few hard coral species in Jeju
mutabilis, and Xenostrobus atratus (Lischke, 1871),
and
of
species,
Island. A. japonica, the dominant species, forms a
Modiolus
coral carpet, a relatively thin layer of coral framework,
nipponicus (Oyama, 1950), and S. keenae. Also, six
in relatively shallow water, 5-15 m, along many areas
other
Lasaeidae,
of the Jeju coastline. Many mollusks have developed
Ungulinidae, and Nuculidae, all containing small
thriving communities mainly at the base of the coral
Modiolus
kurilensis
families
were
Bernard,
found,
1983,
including
species. Although other bivalve species were found in
(Noseworthy et al., 2016a). At present, 61 species,
the sample, they may have inhabited other habitats
about half of them bivalves, have been obtained from
and been washed up with the seaweed.
A. japonica colonies, although bivalves have the
Seagrass is uncommon in Jeju Island, being found
largest
number
of
specimens.
The
Mytilidae,
only in a few localities on the northeast coast. At
Chamidae, and Arcidae are the most speciose of the
present,
Bivalvia,
Zostera
marina
L.,
1753,
the
most
with
five,
four,
and
three
species,
wide-ranging marine flowering plant in the Northern
respectively, and Lithopaga lischkei (Huber, 2010), B.
Hemisphere, is the only seagrass species reported from
stearnsi, Chama cerinohodon Hamada et Matsukuma,
the island (Park J.-I. et al., 2012). Many bivalves
2005, and C. leana are the dominant species. Almost
inabit Z. marina, expecially the Mytilidae. At Ojo-ri, a
all bivalves inhabiting the coral are suspension
large lagoon-like area, about 1 km west of Seongsanpo,
feeders,
on the northeast coast, large dried clumps of seagrass
commensals. Bivalves can be classified according to
may be found along the low tide line on the eastern
their habitats in coral: borers, encrusters, and crevice
side of the lagoon. Four mytilid species inhabit the
dwellers, or nestlers (Noseworthy et al., 2016a). Most
although
some
are
detritus
feeders
or
seagrass; three are small species, B. mutabilis, X.
of the bivalve species inhabiting A. japonica colonies
atratus, and M. viridulus, and juvenile specimens of
are
M. kurilensis are also present. Three species of
Carditidae, which attach themselves by a byssus to the
Pillucina (Lucinidae) and juvenile specimens of the
substrate or nestle in crevices. Other bivalves are
crevice
dwellers,
such
as
the
Arcidae
and
venerids Paphia cf. vernicosa (Gould, 1861) and R.
either
philippinarum, as well as a species of Irus, are also
Spondylidae, which cover dead coral or other hard
found there. Among other species, the Ungulinidae and
substrates,
Lasaeidae also occur, as well as N. paulula, the most
penetrate corals or other calcareous substrata. A
common small species in the seagrass.
zonal-geographical examination of the distribution
Although there are several studies of soft corals in
Jeju Island, there is little information on bivalves-soft
encrusters,
and
such
borers,
as
e.g.,
the
Chamidae
Lithophaga,
and
which
ranges of the mollusks associated with A. japonica
demonstrates
- 164 -
that
the
majority
of
species
have
Korean J. Malacol. 35(2): 149-238 2019
tropical-subtropical (warm-water) affinities, and most
1837), Pecten sinensis G.B. Sowerby II, 1842, Pecten
of them are bivalves. Bivalve larvae can remain viable
puncticulatus Dunker, 1877, Isognomon ephippium (L.,
in the water column longer that gastropod larvae,
1758), Laevichlamys lemniscata (Reeve, 1853), Chlamys
in
squamata (Gmelin, 1791), Chlamys irregularis (G.B.
northward-flowing currents (Brusca and Brusca, 2003),
Sowerby II, 1842) (preoccupied name) (= Laevichlamys
enabling
them
to
travel
longer
distances
and this may account for the preponderance of
cuneata (Reeve, 1853)), Chlamys larvata (Reeve, 1853)
tropical-subtropical bivalve species occurring in the
(a synonym of Laevichlamys squamosa (Gmelin, 1791),
coral carpet in Jeju Island. In general, the sizes of
Isomonia umbonata (Gould, 1861). These bivalves are
bivalve
A.
valid species living either in other parts of Korea or in
japonica are smaller than usual, suggesting a size
other parts of the Indo-Pacific but not found in Jeju.
adjustment to a more restricted habitat.
The rest of the species names regarded in current
specimens
living
in
association
with
taxonomic literature as synonyms and enumerated by
Noseworthy et al. (2007), and also mentioned in some
Annotated Catalogue of Marine Bivalve
Mollusk Species of Jeju-do
other, mainly ecological, papers are listed in this
catalogue under different binomina but they do live in
In first part of this catalogue, 105 species and 21
Jeju; these names are discussed in the comments
families are included (orders Nuculida, Nuculanida,
sections
Solemyida, Arcida, Mytilida, Ostreida and Pectinida).
mentioned in the catalogue below are odd, doubtful, or
Among
trapezina
unreliable and, thus, are not included here because
them,
five
species
(Barbatia
of
relevant
species.
Some
records
not
(Lamarck, 1819), Anadara kagoshimensis (Tokunaga,
there are no further literature confirmations of their
1906), Anadara sp., Anadara granosa (L., 1758),
existence, no voucher material is available to be
Isognomon nucleus (Lamarck, 1819)) are recorded for
consulted, or they are fossil species not known from
the first time in Jeju Island, and one species (I.
recent seas. In the Pectinidae, Patinopecten tokyoensis
nucleus) is a new record for Korea. Another 11 species
(Tokunaga, 1906) and Chlamys cosibensis (Yokoyama,
(Nucula tokyoensis Yokoyama, 1920, Acila insignis
1911) (Je J.-G. et al., 1994a; 2002) are Japanese
(Gould, 1861), Megayoldia japonica (A. Adams et
Neogene-Early Pleistocene species (Masuda, 1962); in
Reeve, 1850), Modiolus comptus G.B. Sowerby III,
the Limidae, Limaria basilanica (A. Adams et Reeve,
1915, Lithophaga cf. malaccana (Reeve, 1857), Atrina
1850) and Limaria hakodatensis (Tokunaga, 1906) (Je
japonica (Reeve, 1858), Saccostrea scyphophilla (Peron
J.-G. et al., 1994a; 2002) may be found in Jeju when
et Lesueur, 1807), Parahyotissa numisma (Lamarck,
additional surveys are undertaken; and in the Arcidae,
1819),
Pinctada
Barbatia lacerata (Bruguière, 1789) (Suzuki, 2010) is a
margaritifera (L., 1758), Malleus regula (Forsskål in
tropical species not known in Korea (Min D.-K. et al.,
Niebuhr, 1775)) were not listed in the check-list of
2004; Lee J.-S., 2013), and distributed in Japan only in
Jeju bivalves by Noseworthy et al. (2007), but are
the Amami Islands and southwards (Higo et al., 1999).
Anomia
cytaeum
Gray,
1850,
known from literature either published before or after
2007, and, thus, they are added here to the Jeju
fauna. Some species listed by Noseworthy et al. (2007)
Subclass PROTOBRANCHIA Pelseneer, 1889
are excluded from the Jeju fauna because their true
Order NUCULIDA Dall, 1889
identity was checked and re-assessed based on voucher
Superfamily Nuculoidea Gray, 1824
material: Arca boucardi Jousseaume, 1894, Anadara
Family Nuculidae Gray, 1824
inaequivalvis (Bruguière, 1789)”, Glycymeris imperialis
Genus Nucula Lamarck, 1799
Kuroda, 1934, Glycymeris munda (G.B. Sowerby III,
Subgenus Nucula Lamarck, 1799
1903), Musculus nanus (Dunker, 1857), Gregariella
1. Nucula (Nucula) paulula A. Adams, 1856
semigranata (Reeve, 1858), Septifer excisus (Wiegmann,
- 165 -
Pl. 1, Figs. B, E
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Seopjikoji,
General distribution: Japan (Izu Islands, Miyake
Munseom, Seogundo, Hwasoon, Hamo, Tongbatarl;
Island, Boso Peninsula and southwards, Kii Peninsula,
other records: Je J.-G. et al. (2002) (Hado); Noseworthy
southern East/Japan Sea), Taiwan, Okinawa, East
Local
distribution:
Udo
(Sanhosa),
and Choi K.-S. (2010) (Seogundo); Lee H.-J. et al.
China Sea, Korea (Higo et al., 1999; Min D.-K. et al.,
(2014) (Tongbatarl (Ojo-ri) Lagoon); Lee J.-S. (2014)
2004; Huber, 2010); subtropical.
(Seopjikoji, Hwasoon).
Habitat: On sandy mud, at depth of 50-1350 m (Higo
General distribution: Korea and Japan to Taiwan, East
et al., 1999; Huber, 2010).
China Sea (Yangtze River delta) (Higo et al., 1999;
Comments: Je et al. (1991) included this species among
Min D.-K. et al., 2004; Qi, 2004); subtropical.
the seven most abundant soft bottom species in the
Habitat: Under rocks in tidepools, on sandy mud or
Southern Sea of Korea, including the area around Jeju
fine sandy substrate, from the intertidal zone to depth
Island; an illustration was given by Je (1993). N.
of 50 m (Huber, 2010). It inhabits coarse sand in the
tokyoensis appears to be a somewhat variable species
bottoms of tidepools around Seogundo, off southern
in shape and it might be possible that Nucula
Jeju Island (Noseworthy and Choi, 2010).
gemmulata Habe, 1953 is based on a more trigonal
Comments: Robba et al. (2007, p. 5, fig. 4a, b) recorded
juvenile (Huber, 2010). Lectotype and paralectotype of
and illustrated this species from the Gulf of Thailand,
N. tokyoensis are figured by Oyama (1973, pl. 20, figs.
and stated that the Thai specimens conform to the
28, 29); the holotype of N. gemmulata is figured by
Japanese shells in all respects but they have a
Higo et al. (2001, p. 145, fig. B4). This species was not
somewhat more triangular outline. However, they also
listed in the previous checklist of bivalves of Jeju
mentioned (l.c., p. 5) that the outer surface of the
(Noseworthy et al., 2007).
specimens possesses uneven growth lines crossed by
faint radial striations, whereas Japanese and Korean
Genus Acila H. et A. Adams, 1858
specimens have a smooth and polished surface, e.g.,
Subgenus Acila H. et A. Adams, 1858
Okutani (2000, pl. 415, fig. 1; Min D.-K. et al., 2004, p.
3. Acila (Acila) mirabilis (A. Adams et Reeve, 1850)
366, fig. 1160; Lutaenko, 2014, pl. 1, fig. A), and their
Synonyms: Nucula sculpta Pease, 1860; Acila divaricata
postero-dorsal margin is more curved. A closer species
submirabilis Schenck, 1936 (non Makiyama, 1926); Acila
but of larger size, up to 7 mm, may be Nucula
schencki Habe, 1958; Acila (Acila) archibenthalis Okutani,
(Nucula) striolata A. Adams, 1856, originally described
1964.
from the “China Seas” (Adams, 1856, p. 52) and
having a crenulate margin, rather smooth surface and
Local distribution: Jeju (Lee J.-S., 2014; as Acila
the umbones rather acute. This was not recognized by
divaricata divaricata).
subsequent authors in China or Japan, and the
General distribution: Yellow Sea, East/Japan Sea,
identity of this species and its distribution are
Korea, Japan (Kyushu, Honshu) (Min D.-K. et al.,
unknown at present (Huber, 2010); there are four
2004; Huber, 2010; Zhang et al., 2014; Lutaenko and
syntypes of this species in the Natural History
Noseworthy, 2019); subtropical-lowboreal.
Museum, London (l.c.). A possible syntype of N.
Habitat: Subtidal-bathyal, at depth of 15-800 or 1470
paulula is figured by Higo et al. (2001).
m, mud, sandy mud and fine sand (Huber, 2010).
Subgenus Lamellinucula Schenck, 1944
2000) and Russian (Scarlato, 1981) workers believed
2. Nucula (Lamellinucula) tokyoensis Yokoyama, 1920
that Nucula mirabilis A. Adams et Reeve, 1850 is a
Synonyms: ?Nucula (Lamellinucula) gemmulata Habe, 1953.
synonym of Acila (Acila) divaricata (Hinds, 1843).
Comments: Japanese (e.g., Higo et al., 1999; Okutani,
Meanwhile, Chinese malacologists (Xu, 1984; Qi, 2004)
Local distribution: northern coast of Jeju Island (Je
separated A. divaricata and A. mirabilis, and showed
J.-G. et al., 1991).
that A. divaricata ranges only from the East China to
- 166 -
Korean J. Malacol. 35(2): 149-238 2019
the South China seas, whereas A. mirabilis is a
member
of
the
Yellow
Sea
Cold
Water
Mass
community. Later, this opinion was supported by
Order NUCULANIDA Carter, Campbell et Campbell, 2000
Superfamily Nuculanoidea H. et A. Adams, 1858
Family Nuculanidae H. et A. Adams, 1858
Huber (2010): A. divaricata, originally described from
Genus Nuculana Link, 1807
China, is smaller, generally less than 20 mm, with
Subgenus Nuculana Link, 1807
weaker ribbing, less rostrate posteriorly, with a very
5. Nuculana (Nuculana) scalata Prashad, 1932
fine, regular ribbing on the inner ventral margin. It is
Synonyms: ?Nuculana subscalata Okutani, 1962.
distributed in Philippines and Chinese waters, and not
found in Japan or in Russia. Recently, Zhang et al.
Local distribution: Seongsan (Min D.-K. et al., 2004).
(2014) clearly confirmed this geographical pattern and
General distribution: Korea and south-east Japan to
morphological separation by a genetic-molecular study:
Taiwan, South China Sea and Indonesia (Higo et al.,
according to them, A. mirabilis is commonly found in
1999;
the Yellow Sea, the northern East/Japan Sea, and the
tropical-subtropical.
Russian Far Eastern seas (Okutani and Saito, 2017;
Habitat: Sandy mud, at depth of 100-1640 m (Huber,
Lutaenko and Noseworthy, 2019). The distribution of
2010).
Min
D.-K.
et
al.,
2004:
Huber,
20100);
A. divaricata and A. mirabilis in the Yellow Sea and
Comments: Huber (2010) suggested that Nuculana
East China Sea is separated by the Changjiang
subscalata Okutani, 1962 might be a synonym of this
(Yangtze) River estuary. This species was not listed in
species. At least, specimens dredged off northern
the previous checklist of bivalves of Jeju (Noseworthy
Borneo at approximately 100 m deep are intermediate
et al., 2007).
in sculpture and shape between N. subscalata and N.
scalata (l.c.). Part of the records of N. scalata in Japan
Subgenus Truncacila Grant et Gale, 1931
are,
4. Acila (Truncacila) insignis (Gould, 1861)
Tsuchida et Okutani, 1985 distributed around Boso
Synonyms: Nucula (Acila) japonica Dall, 1898 (non A. Adams
and Kii peninsulas (Honshu) and southwards, from
et Reeve, 1850).
southern Kyushu to Tanegashima and Yakushima, at
in
fact,
Nuculana
(Nuculana)
tanseimaruae
depth of 700-1031 m (Higo et al., 1999). The holotype
Local distribution: Munseom (Je J.-G. et al., 1994a);
of N. subscalata was figured by Higo et al. (2001, p.
Jeju (Je J.-G. et al., 2002; Lee J.-S., 2014).
147, fig. B60).
General distribution: Korea (North Korea, Gangwon-do,
Gyeongsangbuk-do, Jeju), Japan (Hokkaido, East/Japan
6. Nuculana (Nuculana) leonina (Dall, 1896)
Sea,
western
Synonyms: Leda amiata Dall, 1916; Leda liogona Dall, 1916;
(Russia)
Nuculana sagamiensis Okutani, 1962.
Boso
Kyushu),
Peninsula
northwestern
and
northwards,
East/Japan
Sea
(Scarlato, 1981; Higo et al., 1999; Min D.-K. et al.,
2004; Lutaenko and Noseworthy, 2012; Lee J.-S.,
Local distribution: Seongsan (Min D.-K. et al., 2004).
2014); subtropical-lowboreal.
General distribution: Kamchatka to Korea and Japan;
Habitat: Fine sand and mud, muddy sand, from
southern Bering Sea to California (Scarlato, 1981; Higo
intertidal zone to depth of 200 m (Scarlato, 1981; Higo
et al., 1999, as Nuculana sagamiensis); widely distributed
et al., 1999).
boreal.
Comments: The holotype is figured by Higo et al.
Habitat: Brown mud, fine sandy mud and green ooze,
(2001, p. 145, fig. B12). This species was not listed in
at depth of 200-2560 m (Huber, 2010).
the previous checklist of bivalves of Jeju (Noseworthy
Comments:
et al., 2007).
sagamiensis as a distinct species, and morphologically
Huber
(2010)
did
not
recognize
N.
no traits were found separating Japanese specimens
from Washington State specimens; adult size and
- 167 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
depth of N. sagamiensis correspond well to N. leonina.
Yellow seas (Higo et al., 1999; Min D.-K. et al., 2004;
The holotype of N. sagamiensis is figured by Higo et
Xu and Zhang, 2008); subtropical.
al. (2001, p. 147, fig. B63).
Habitat: Sandy mud, from the intertidal zone to depth
of 50 m (Huber, 2010).
7. Nuculana (Saccella) gordonis (Yokoyama, 1920)
Comments:
Huber
(2010)
states
that,
although
Japanese authors and additionally Kamenev (2009)),
Local distribution: Seongsan (Min D.-K. et al., 2004;
due to ligament position, consistently refer this species
Lee J.-S., 2014).
to Acharax Dall, 1908, it does not belong to this genus.
General distribution: Korea and Japan (south-eastern
It is not close in broad anterior shape, neither does it
East/Japan Sea, to Niigata Prefecture; north-eastern
show the typical extended marginal fringes, nor is it
Honshu and southwards) to Yellow and East China
large (l.c.). For this species, he proposes a new
seas (Higo et al., 1999; Qi, 2004); subtropical.
subgenus, Pseudacharax Huber, 2010 (type species:
Habitat: Fine sand and mud, at depth of 15-300 m
Solenomya japonica Dunker, 1882). The lectotype of S.
(Huber, 2010).
yamakawai is fugured by Oyama (1973, pl. 27, fig. 1).
Comments: Lectotype and paralectotype are figured by
Oyama (1973, pl. 21, figs. 1, 2).
Subclass AUTOBRANCHIA Grobben, 1894
Order ARCIDA Gray, 1854
Family Yoldiidae Dall, 1908
Superfamily Arcoidea Lamarck, 1809
Subfamily Yoldiellinae Allen et Hannah, 1986
Family Arcidae Lamarck, 1809
Genus Megayoldia Verrill & Bush, 1897
Subfamily Arcinae Lamarck, 1809
Genus Arca L., 1758
8. Megayoldia japonica (A. Adams et Reeve, 1850)
Subgenus Arca L., 1758
Local distribution: northern coast of Jeju Island (Je
10. Arca (Arca) avellana Lamarck, 1819
Pl. 1, Fig. G, H
J.-G., 1993).
General distribution: Korea and Japan, East China
Synonyms: Arca retusa Lamarck, 1819; Byssoarca maculata
Sea, Yellow Sea (Higo et al., 1999; Huber, 2010);
Sowerby I, 1833; Arca ocellata Reeve, 1844; Arca cunealis
subtropical.
Reeve, 1844; Arca arabica Philippi, 1847; Arca kraussii
Habitat: Subtidal, at depth of 10-300 m; fine sandy
Krauss, 1848; Arca acuminata subnormalis Pilsbry, 1895;
and muddy bottoms (Huber, 2010).
Arca bicarinata Sowerby III, 1901; Navicula aladdin Iredale,
1939; Navicula terebra Iredale, 1939.
Order SOLEMYIDA Dall, 1889
Local distribution: Samyang, Hamdeok, Haengwon,
Superfamily Solemyoidea Gray, 1840
Udo
Family Solemyidae Gray, 1840
Daejong, Shinyang, Pyoseon, Wimi, Bomok, Munseom,
Genus Solemya Lamarck, 1818
Gangjeong, Yerae, Daepyeong, Jungmun, Hwasoon,
Subgenus Pseudacharax Huber, 2010
Yeongmeori, Sagyeri, Songaksan, Hamo, Keumneung,
9. Solemya (Pseudacharax) japonica (Dunker, 1882)
Biyangdo,
Synonyms: Solemya yamakawai Yokoyama, 1927.
Gwakji; other records: Lee J.J. and Jwa Y.W. (1988; as
Local distribution: Hwasoon (Min D.-K. et al., 2004).
boucardi); Lee J.J. (1990; as Arca boucardi) (northern
General distribution: Korea and Japan (southern
coast); Lee J.J. (1991; as Arca boucardi) (Gapado,
Hokkaido
Sea;
Marado); Lee J.J. and Hyun J.M. (1991; as Arca
south-eastern East/Japan Sea) to East China and
boucardi) (Seogwipo); Kim H.S. et al. (1992); Lee J.J.
(Sanhosa),
Weoljeong,
Hyeopjae,
Suwon,
Seongsan,
Gwideok,
Tongbatarl,
Soessogak,
Arca “bronchardi”); Lee J.J. et al. (1989; as Arca
and
southwards;
Seto
Inland
- 168 -
Korean J. Malacol. 35(2): 149-238 2019
and Hyun J.M. (1992) (eastern coast; Udo); Je J.-G. et
former one. The two latter taxa do not have anything
al. (1994a) (Munseom); Lee J.J. et al. (1995; as Arca
in common in morphology (A. dayi, perhaps, a distinct
boucardi) (Jagwi-do); Lee J.-J. and Hyun J.-M. (1997;
species), whereas a specimen figured by Huber (2010,
as Arca boucardi) (Chujado); Je J.-G. et al. (2002;
p. 129; as “A. ocellata”) from northern Borneo does
including Arca arabica and Arca boucardi) (Bumseom,
represent A. koumaci. This is a rather peculiar species,
Munseom, Supseom, Jigwido); Lee J.J. and Hyun J.M.
clearly differentiated from all known species of the
(2002; as Arca boucardi) (Songacksan); Noseworthy et
genus by its small size (to 12-13 mm), presence of
al. (2002; as Arca boucardi) (Seongsan); Noseworthy
spikes (up to 5-7) on posterior ridge, lack of ventral
and
boucardi)
sulcus, and yellowish color. The Borneo specimen was
(Seogundo); Suzuki (2010; as Arca arabica and Arca
found at 70 m deep, among coral rubble at the wall
boucardi) (Hwasun, Ojo-ri, Hyopjae); Ko J.-C. et al.
bottom, a rather characteristic habitat for A. koumaci.
Choi
K.-S.
(2010;
including
Arca
(2011; as Arca boucardi) (Gwakji, Sincheon, Woljeong,
Huber (2015) claims that type material of A. ocellata
Jungmun, Hacheon, Haengwon); Cho I.-Y. et al. (2014)
corresponds to his specimen from Borneo; however,
(Beomseom, Munseom); Je J.-G. et al. (2002); Lee J.-S.
Reeve’s (1844) original description and illustrations
(2013) (Seongsan; Sagyeri); Cho I.-Y. et al. (2014)
(Reeve, 1843-1844, pl. 15, sp. 102) do not show
(Beomseom,
Munseom); Noseworthy et al.
(2016;
characteristic features of A. koumaci, e.g., posterior
including Arca cf. boucardi) (Keumneung).
ridge spikes. Therefore, Huber’s (2015) view is not
General distribution: Korea and Japan (Tsugaru
accepted.
Peninsula and southwards) to Indo-West Pacific;
Arca boucardi Jousseaume, 1894, previously recorded
Australia; South Africa (Lamy, 1907; Higo et al., 1999;
for Jeju (Kim H.-S. et al., 1992; Je et al., 2002; Lee
Huber, 2010; Lutaenko and Noseworthy, 2012; Lee
J.J. and Hyun J.M., 2002; Noseworthy et al., 2002;
J.-S., 2013); tropical-subtropical.
2007;
Habitat: Sheltered bays and coral reefs; byssally
misidentification; only one species of the genus, A.
Lee
J.-S.,
2013;
and
others),
is
a
attached to rocks, shells, corals, or coarse sand, also in
avellana, is present. “Arca arabica” listed by Je et al.
crevices, from the intertidal zone to depth of 100 m
(2002) and Suzuki (2010) is a synonym of A. avellana.
(Huber, 2010).
Comments: Huber (2010) showed that Röding (1798)
Genus Barbatia J. Gray, 1842
erected Arca patriarchalis based on Chemnitz’s figure;
Subgenus Abarbatia Dall, Bartsch et Rehder, 1938
it was mentioned after 1899 by Lamy (1907) as a valid
11. Barbatia (Abarbatia) trapezina (Lamarck, 1819)
Pl. 5, Figs. A-E
name. Thus, unless an ICZN-petition is made to
invalidate Röding’s name, Lamarck’s well-known A.
Synonyms: Arca lima Reeve, 1844.
avellana has to be regarded a junior synonym (Huber,
2010). We still use the name A. avellana until a
Local
possible decision by the ICZN may be taken although
Seongsan, Bomok, Seogwipo, Munseom, Seogundo,
some Korean workers have started to use the name A.
Jungmun,
patriarchalis (Lee Y. et al., 2016). Syntypes of Arca
Gwakji; other records: ?Je J.-G. et al. (2002; as
avellana Lamarck, 1819 and Arca retusa Lamarck,
Barbatia lima) (Munseom); Yang H.-S. and Choi K.-S.
distribution:
Yerae,
Udo
(Sanhosa),
Hwasoon,
Daepyeong,
Yongmeori,
Sagyeri,
1819 are figured by Lutaenko and Maestrati (2007).
(2011; as Barbatia virescens).
Huber (2015) regarded Arca ocellata Reeve, 1844 (type
General distribution: Korea, Japan, China, Thailand,
locality: Singapore; Reeve (1844)) as a valid species
Philippines, Indonesia and the Indo-Pacific (Okutani,
and synonymized two other species, Arca acuminata
2000;
dayi Oliver et Chesney, 1994 (type locality: Masirah,
Volvenko, 2013); tropical-subtropical.
Oman) and Arca koumaci Lutaenko et Maestrati, 2007
Habitat: Byssally attached to rocks and coral blocks,
(type locality: New Caledonia, Koumac) with the
from intertidal zone to upper subtidal zone. Records of
- 169 -
García
and
Oliver,
2008;
Lutaenko
and
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
B. trapezina are often mixed with other species of the
in the literature (e.g., Huber (2010)) may belong to
complex.
this species, and the geographical distribution of B.
Comments: García and Oliver (2008) revised the
decussata
so-called “foliata/decussata complex”, in which they
synonymized this species with B. trapezina. Syntypes
included Barbatia foliata (Forsskål in Niebuhr, 1775),
of Byssoarca decussata G.B. Sowerby I, 1833 are
B. trapezina, Barbatia decussata (G.B. Sowerby I,
figured by Higo et al. (2001, p. 150, fig. B191 in color)
1833), and Barbatia grayana Dunker, 1867. We
and García and Oliver (2008, p. 13, fig. 3).
should
be
re-assessed.
Huber
(2010)
distinguish B. trapezina from B. decussata based on
morphological criteria and type designations of the
Subgenus Savignyarca Jousseaume, 1891
above authors: B. trapezina has spicate and erect
13. Barbatia (Savignyarca) virescens (Reeve, 1844)
Pl. 6, Figs. A-F; Pl. 7, Figs. A, B
periostracal bristles, rather than blunt and adpressed,
a more rectangular shell as compared to the more
Synonyms: Arca obliquata Reeve, 1844 (non Wood, 1828);
anteriorly reduced shell of B. decussata, only four or
Arca obtusa Reeve, 1844 (non de Konninck, 1842); Arca
five posterior riblets remaining deeply cut throughout
obtusoides Nyst, 1848; ?Arca sinensis Philippi, 1851; Arca
life and widely spaced, whereas other riblets are
decurvata Lischke, 1869.
narrow and weekly nodulose. According to Huber
(2010; 2015), Arca lima Reeve, 1844 is a synonym of
Local distribution: Hamdeok, Handong, Hado, Udo
Barbatia foliata (Forsskål in Niebuhr, 1775) whereas
(Hagosudong), Udo (Sanhosa), Jongdal-ri, Seongsan,
García and Oliver (2008) believe that the former is a
Shinyang, Pyoseon, Daepyeong, Weoljeong, Bomok,
synonym of B. trapezina. The lectotype of Arca
Munseom,
trapezina Lamarck, 1819 and a syntype of Arca lima
Keumneung, Soessogak; other records: Je J.-G. et al.
Reeve, 1844 are figured by García and Oliver (2008, p.
(1994a) (Munseom); Cho I.-Y. et al. (2014) (Munseom);
11, fig. 2). A new record for Jeju Island.
Je J.-G. et al. (2002); Cho I.-Y. et al. (2014)
Yerae,
Hwasoon,
Yongmeori,
Sagyeri,
(Munseom); Lee J.-S. (2013) (Seongsan, Sagyeri); Lee
12. Barbatia (Abarbatia) decussata (G.B. Sowerby I, 1833)
Pl. 4, Figs. C, D
Synonyms:
?Barbatia
J.U. et al. (2017).
General distribution: Korea, Japan (southern Hokkaido
oblonga
Dunker,
1868;
?Barbatia
and southwards), China, South China Sea, Vietnam,
Thailand, Philippines (Lamy, 1907; Higo et al., 1999;
petersii Dunker, 1871.
Min D.-K. et al., 2004; García and Oliver, 2008; Xu
Local distribution: Seongsan, Yeongmeori, Hwasoon,
and Zhang, 2008; Huber, 2010; Lee J.-S., 2013);
Yerae; other records: ?Je et al. (2002; as Barbatia
tropical-subtropical.
lima); Noseworthy and Choi K.-S. (2010) (Seogundo);
Habitat: Byssally attached to rocks, under stones, and
Cho I.-Y. et al. (2014) (Munseom); Cho I.-Y. et al.
in crevices, from the intertidal zone to depth of 20 m
(2014)
(Huber, 2010).
(Munseom);
Lee
J.-S.
(2013)
(Seongsan,
Hwasoon, Sagyeri).
Comments: This species has its velvety, dark greenish
General distribution: Japan (Boso Peninsula and
periostracum eroded in the central shell area, with
southwards; Oga Peninsula and southwards in the
periostracum remaining around the edge of the shell.
East/Japan Sea), Korea, China, Taiwan, Thailand,
It is variable in shape, usually white, occasionally
Philippines, Indonesia and the Indo-Pacific (Higo et al.,
greenish from algae, often deformed. Arca decurvata
1999; Min D.-K. et al., 2004; García and Oliver, 2008;
Lischke, 1869 is a replacement name (nom. nov. pro
Lee J.-S., 2013); tropical-subtropical.
Arca obliquata Reeve, 1844 non Arca obliquata W.
Habitat: Byssally attached to rocks, intertidal-upper
Wood, 1828) but the earlier replacement name for
subtidal.
Reeve’s species is Arca obtusoides Nyst, 1848 (Nyst,
Comments: Some of the synonyms of B. foliata listed
1848, p. 50). Voucher specimens of Arca decurvata of
- 170 -
Korean J. Malacol. 35(2): 149-238 2019
Lischke are figured by Lutaenko (2015, pl. 3, figs. E-F,
1939; Hawaiarca miikensis Noda, 1966.
and taxonomic discussion therein); syntypes of Arca
obtusa Reeve, 1844 and Arca virescens Reeve, 1844 are
Local
figured by Higo et al. (2001, p. 150, figs. 192, 192s).
Munseom,
distribution:
Seopjikoji,
Hwasoon,
Seongsan,
Yongmeori,
Pyoseon,
Sagyeri;
other
records: Lee J.J. et al. (2001; as “Nipponica” bistrigata)
Subgenus Ustularca Iredale, 1939
(Hwasoon); Cho I.-Y. et al. (2014) (Munseom); Lee J.-S.
14. Barbatia (Ustularca) stearnsii (Pilsbry, 1895)
(2013) (Hwasoon, Seopjikoji, Sagyeri).
General distribution: Widely distributed Indo-Pacific
Pl. 7, Figs. C-H
species:
Korea
and
Japan
(Boso
Peninsula
and
Local distribution: Iho, Jeju-si, Hamdeok, Bukchon-ri,
southwards), Yellow Sea, East China Sea, South China
Gimnyeong, Wolpyeong, Haengwon, Handong, Sehwa,
Sea,
Daejeong,
(Sanhosa),
Mozambique (Higo et al., 1999; Min D.-K. et al., 2004;
Jongdal-ri, Seongsan, Tongbatarl, Seopjikoji, Shinyang,
Qi, 2004; García and Oliver, 2008; Huber, 2010; Lee
Pyoseon,
Udo
Bomok,
(Hagosudong),
Supseom,
Udo
Seogwipo,
Munseom,
Thailand,
India,
Pakistan,
eastern
Africa,
J.-S., 2013; Zhang et al., 2016); tropical-subtropical.
Seogundo, Gangjeong, Jungmun, Yerae, Daepyeong,
Habitat: Byssally attached to rocks, in shell debris and
Hwasoon,
muddy areas, at depth of 3-55 m (Huber, 2010).
Yongmeori,
Sagyeri,
Songaksan,
Hamo,
Keumneung, Biyangdo, Hyeopjae, Gwideok, Gwakji;
Comments: This species was previously placed in the
other records: Lee J.J. (1990) (northern coast); Je J.-G.
genus Nipponarca Habe, 1951 of which it the type
et al. (1994a) (Munseom); Lee J.J. et al. (2001)
species. Lamprell and Healy (1998) and Xu and Zhang
(Hwasoon); Je J.-G. et al. (2002) (Munseom, Bomseom,
(2008) Synonymszed Barbatia signata (Dunker, 1868)
Jigwido);
with B. bistrigata. Stevenson (1972), Huber (2010), and
Noseworthy
et
al.
(2002)
(Seongsan);
Noseworthy and Choi K.-S. (2010) (Seogundo); Ko J.-C.
Lamprell and Healy (1998) synonymized “Nipponarca”
et al. (2011) (Daepo, Aewol, Woljeong); Lee J.-S. (2013)
adamsiana (Dunker, 1866) with B. bistrigata; García
(Seongsan, Sagyeri); Cho I.-Y. et al. (2014) (Munseom);
and Oliver (2008) stated that it is likely to be correct,
Noseworthy et al. (2016) (Keumneung); Lee J.U. et al.
but other authors do not share this opinion (Scott,
(2017).
1994; Lutaenko and Volvenko, 2013). A syntype of A.
General distribution: Korea, Japan (Boso Peninsula
obtusa var. duplicostata is figured by Coan et al.
and southwards, East/Japan Sea), and Taiwan to
(2015, fig. 5); the holotype of H. miikensis is figured by
South China Sea, Philippines (Higo et al., 1999; Min
Noda (1966, pl. 8, figs. 9, 14).
D.-K.
et
al.,
2004; Qi, 2004;
Lee J.-S.,
2013);
tropical-subtropical.
Genus Acar Gray, 1857
Habitat: Byssally attached to rocks, in coarse or
16. Acar plicata (Dillwyn, 1817)
Pl. 1, Fig. F
muddy sand, from the intertidal zone to depth of 83 m
(Huber, 2010).
Synonyms: Byssoarca divaricata Sowerby I, 1833; Arca
Comments: A common shell in beach drift. A syntype
irioides Menke, 1843; Arca donaciformis Reeve, 1844; Arca
of B. stearnsii is figured by Higo et al. (2001, p. 150,
(Byssoarca) dubia Baird in Brenchley, 1873; Acar hawaiensis
fig. B196).
Dall, Bartsch et Rehder, 1938; Acar laysana Dall, Bartsch et
Rehder, 1938; Acar dubia digma Iredale, 1939; Acar dubia
kerma Iredale, 1939; Barbatia (Acar) numaensis Noda, 1966.
Subgenus Mesocibota Iredale, 1939
15. Barbatia (Mesocibota) bistrigata (Dunker, 1866)
Pl. 4, Fig. G
Local
distribution:
Handong,
Hado,
Jongdal-ri,
Synonyms: ?Barbatia paulucciana Tapparone-Canefri, 1876;
Seopjikoji, Shinyang, Pyoseon, Munseom, Hwasoon,
Arca (Barbatia) fischeri Lamy, 1907; Arca obtusa var.
Jungmun,
duplicostata Grabau et King, 1928; ?Mesocibota luana Iredale,
Keunmeung, Hyopjae, Gwakji; other records: Je J.-G. et
- 171 -
Daepyeong,
Yeongmeori,
Sagyei,
Hamo,
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
al. (2002); Je J.-G. et al. (1994a) (Munseom); Je J.-G. et
Local distribution: Seongsan (Min D.-K. et al., 2004).
al. (2002; as Arca) Ko J.-C. et al. (2011) (Hacheon); Lee
General distribution: Northern Japan and southwards,
J.-S. (2013) (Hwasoon, Seongsan, Sagyeri).
East/Japan Sea, southern Korea to East China Sea
General distribution: Korea and Japan (Tsugaru
(Higo et al., 1999; Min D.-K. et al., 2004); subtropical.
Peninsula and Niigata Prefecture and southwards),
Habitat: Fine sand and mud, from depth of 20-500 m
East and South China seas, Philippines to Indo-Pacific
(Huber, 2010).
including South Africa, Red Sea, India, Seyshelles,
Comments: This species was sometimes synonymized
northern Australia, Hawaii, the Mediterranean Sea
with the Indo-Pacific Bathyarca anaclima (Melvill et
(Lessepsian migrant) (Higo et al., 1999; Qi, 2004;
Standen, 1907) (e.g., Higo et al., 1999; Min D.-K. et al.,
Zenetos et al., 2005; Galil, 2007; Xu and Zhang, 2008;
2004), which is somewhat close to the present species;
Huber, 2010; Lee J.-S., 2013); tropical-subtropical.
however, it appears that these two species are
Habitat: Bysally attached to rocks and gravel, from
different. B. kyurokusimana has a sculpture of fine
intertidal zone to depth of 20 m (Huber, 2010).
radial riblets and growth striae giving a cancellate
Comments: This is a very variable, widely-distributed
appearance and its shell reaches 6 mm (Okutani,
Indo-Pacific species. The closest species is Acar
2000), as opposed to B. anaclima which has fewer
congenita (Smith, 1885) from the Philippines (type
radial riblets and is smaller (to 3 mm long) (Huber,
locality: Mindanao (Smith, 1885)) and Indonesia to
2010). While this species was listed and illustrated by
Japan (Huber, 2010), with coarser sculpture and a
Lee J.-S. and Min (2002) and Min D.-K. et al. (2004; in
more compressed shell (see (Okutani, 2000)). Although
both cases as B. anaclima), it is missing in the review
A.
congenita
was
Synonymszed
with
A.
plicata
of the Korean Pteriomorphia (Lee J.-S., 2013).
(Stevenson, 1972), they seem to be separate species.
However, A. plicata may comprise a complex of species
Genus Samacar Iredale, 1936
(Oliver and Holmes, 2004). The holotype of B.
19. Samacar (Samacar) strabo (Hedley, 1915)
numaensis is figured by Noda (1966, pl. 5, figs. 11, 12).
Synonyms: Pseudogrammatodon pacificus Nomura et Zinbo,
1934; Arca acupicta Viader, 1951; Arca approximata Viader,
1951.
Genus Hawaiarca Dall, Bartsch et Rehder, 1938
17. Hawaiarca uwaensis (Yokoyama, 1928)
Local distribution: Seongsan (Min D.-K. et al., 2004).
Local distribution: Hwasoon (Min D.-K. et al., 2004;
General
distribution:
Korea,
Japan
(Honshu
to
Lee J.-S., 2013); Woljeong (Ko J.-C. et al., 2011).
Kyushu), East China Sea to Indo-West Pacific; eastern
General distribution: Korea and Japan (Boso and Noto
Australia; South Africa (Kilburn, 1983; Lamprell and
peninsulas and southwards) to Vietnam and South
Healy,
China Sea, Taiwan, Okinawa, Hawaii (Higo et al.,
tropical-subtropical.
1999;
Habitat: Byssally attached to rocks and sponges, at
Xu
and
Zhang,
2008;
Huber,
2010);
1998;
Higo
et
al.,
1999;
Qi,
2004);
tropical-subtropical.
depths from 20-500 m (Kilburn, 1983; Higo et al.,
Habitat: Byssally attached to rocks, shells, and sunken
1999).
wood, from depth of 10-500 m (Huber, 2010).
Comments: For a detailed discussion and re-description
of this species, see Kamenev (2007). This author also
suggests (l.c.) that along with S. strabo, another,
Subfamily Bathyarcinae Scarlato et Starobogatov, 1979
undescribed, species of this genus occurs off the coasts
Genus Bathyarca Kobelt, 1891
of Japan. S. strabo is missing in the review of the
Subgenus Bathyarca Kobelt, 1891
Korean Pteriomorphia (Lee J.-S., 2013).
18. Bathyarca (Bathyarca) kyurokusimana (Nomura et
Hatai, 1940)
- 172 -
Korean J. Malacol. 35(2): 149-238 2019
Subfamily Anadarinae Reinhart, 1935
Kalishevich, 1976 (Kalishevich, 1976; Talmi Lagoon,
Genus Anadara Gray, 1847
East/Japan Sea, near the Russian-Korean border)
Subgenus Scapharca Gray, 1847
(Lutaenko and Noseworthy, 2019). A. talmiensis was
20. Anadara (Scapharca) talmiensis Kalishevich, 1976
illustrated in the Korean literature as “Scapharca
satowi Dunker, 1882” (Min et al., 2004, fig. 1242). The
Pl. 2, Fig. 3
holotype is figured by Kalishevich (1976, pl. 23, fig. 1).
Local distribution: Iho, Udo (Geomeollae), Jongdal-ri,
Seongsan, Shinyang, Jungmun, Hwasoon, Yongmeori,
21. Anadara (Scapharca) sp.
Pl. 2, Figs. A-C
Wimi; other records: Suzuki (2010; as Scapharca
inaequivalvis) (Sehwa, Hwasoon); Lee J.-S. (2013; as
Scapharca inaequivalvis) (Hwasoon).
Local distribution: Seongsan, Jungmun, Hwasoon,
General distribution: Korea, Japan, and northern
Yeongmeori.
China (Higo et al., 1999; Xu and Zhang, 2008; Huber,
Comments: Shells of specimens of this unidentified
2010); subtropical.
species are rather small (up to 37 mm), nearly
Habitat: Muddy sand and mud, in upper subtidal zone.
subquadrate in shape with well-rounded anterior
Comments: Collected specimens from Jeju Island
margin,
belong to the species complex “Anadara (Scapharca)
thin-shelled, with low beaks, 32-34 radial ribs, and
inaequivalvis (Bruguière, 1789)”. Taxonomy of this
only slightly granulated anteriorradial ribs. A very
anadarine species complex, including a number of
distinctive feature is the nearly straight posterior
fossil species, is very complicated. This group includes,
margin, not typical of known species in north-east
in addition to A. inaequivalvis, at least, Anadara
Asia. In shell shape, specimens are similar to Anadara
covered
with
light-brown
periostracum,
disparilis (Reeve, 1844), Anadara rufescens (Reeve,
(Scapharca) globosa (Reeve, 1844) (see photographs of
1844), Anadara hispida (Philippi, 1849), Anadara
syntypes in: Lutaenko and Volvenko (2013, p. 92, fig.
penangana (Jousseaume, 1893). Arca rhomboidalis
5)) but the latter one grows much larger and has a
Schumacher 1817 is a synonym of A. inaequivalvis, see
swollen shell with dark-brown periostracum, whereas
discussion in Lutaenko and Volvenko (2013). Lutaenko
our specimens are rather flattened and small. A
(2006) showed that only material from southern India
similar specimen is figured from the Pleistocene
represents true A. inaequivalvis; its type locality is the
deposits of Kikaijima Island in Kagoshima Prefecture,
Coromandel coast and Tranquebar (Bruguière, 1789, p.
Kyushu, as Scapharca sp. (Yamashita et al., 1998, pl.
107), and it is clearly different from north-east Asian “A.
1, fig. 8a, b).
inaequivalvis aucct.” Following Huber (2010), we believe
that Japanese-Korean-Chinese “A. inaequivalvis”, as
well
as
subfossil
Holocene
species
from
22. Anadara (Scapharca) kagoshimensis (Tokunaga, 1906)
Pl. 3, Figs. A, B
the
north-western East/Japan Sea (Lutaenko, 1988; 1993a),
Synonyms: Arca amygdalum Philippi, 1845 (non Link, 1807);
is a distinct species and it differs from Anadara
Arca subcrenata Lischke, 1869 (non Michelotti, 1861); Arca
(Scapharca) kagoshimensis (Tokunaga, 1906) although
nodosocrenata Lischke, 1869; ?Arca (Scapharca) peitaihoensis
they were often confused in literature. Huber (l.c.)
Grabau et King, 1928; Scapharca sativa Bernard, Cai et
wrongly used for the above-mentioned species the name
Morton, 1993.
Anadara sativa (Bernard, Cai et Morton, 1993) which
has been proposed as a nom. nov. pro Arca subcrenata
Local distribution: Seongsan, Hwasoon, Yeongmeori,
Lischke, 1869 non Michelotti, 1861, a junior synonym
Daejeong, Handong, Wimi.
of A. kagoshimensis. We think that the correct name
General distribution: Korea and Japan (north-western
for the north-east Asian “A. inaequivalvis aucct.” is the
Honshu and southwards; East/Japan Sea, to northern
little-known, subfossil Holocene Anadara talmiensis
Niigata Prefecture); Yellow Sea, East China Sea (Higo
- 173 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
et al., 1999; Min D.-K. et al., 2004; Lee J.-S., 2013;
commercially in mainland Korea. Yokogawa (1997) and
Zhang et al., 2016); subtropical.
Feng et al. (2011) found that Chinese representatives
Habitat: In Japan, intertidal to 20 m, sandy mud
of this species are different from Japanese ones at the
(Higo et al., 1999); in China, to 56 m, very common
level of species or subspecies based on morphological
(Qi, 2004).
and genetic analyses. Cho E.-S. et al. (2007) detected
Comments: The name Anadara subcrenata (or Scapharca
considerable genetic differences between the Korean,
subcrenata) was widely accepted for this common
Chinese, and Russian populations of A. broughtonii;
north-east Asian anadarine species until the beginning
Tanaka
of the 1990s when Bernard et al. (1993) discovered
differences between populations from Japan/Korea and
and
Aranishi
(2016)
found
significant
that this name is preocuppied by earlier Arca
China using the mitochondrial DNA cytochrome c
subcrenata
kagoshimensis
oxidase subunit I gene, indicating that the divergence
Tokunaga, 1906 is the first available name for this
had reached subspecies level. Huber (2010) unreasonably
species and it was recognized as a synonym of A.
synonymized Anadara (Scapharca) kafanovi Lutaenko,
subcrenata for the first time by Habe (1951, 1965) and
1993 known from the Pacific coast of Japan, southern
Noda (1966). Bernard et al. (1993) proposed the
China and Vietnam (Lutaenko, 1993b; 2008; Xu and
unnecessary
Michelotti,
1861.
Arca
sativa
Zhang, 2008) with A. broughtonii, from which the
Bernard, Cai et Morton, 1993. Korean workers still use
former species is clearly distinguished by shell shape,
incorrect names A. subcrenata (Lee J.-S., 2013) or A.
smaller size, and fewer radial ribs, 35-38 vs. 41-45 on
sativa (Lee Y. et al., 2016). Type material of A.
average in A. broughtonii. Syntypes of A. tenuis are figured
replacement
name
Scapharca
by
online at: http://umdb.um.u-tokyo.ac.jp/DKoseibu/en/Collection/
Lutaenko (2015); syntypes of A. peitaihoensis are
detail.php?umutNo=13788 and http://umdb.um.u-tokyo.ac.jp/
figured by Coan et al. (2015, fig. 4). A new record for
DKoseibu/en/Collection/detail.php?umutNo=13789 (The
Jeju Island.
University
subcrenata
and
its
taxonomy
are
discussed
Museum,
the
University
of
Tokyo
Collection).
23. Anadara (Scapharca) broughtonii (Schrenck, 1867)
Subgenus Tegillarca Iredale, 1939
Pl. 4, Figs. A, B
Synonyms: Arca inflata Reeve, 1844 (non Brocchi, 1814); Arca
24. Anadara (Tegillarca) granosa (L., 1758)
reeveana Nyst, 1848 (non d’Orbigny, 1846); Arca tenuis
Tokunaga, 1906 (non Montagu, 1808).
Pl. 3, Figs. C, D
Synonyms: Arca nodulosa Lightfoot, 1786 (non O.F. Müller,
1776); Arca aculeata Bruguière, 1789; Arca corbicula Gmelin,
Local
distribution:
Ojo-ri,
Munseom,
Bumseom,
1791; Arca corbula Dillwyn, 1817; Arca cuneata Reeve, 1844;
Supseom, Wimi, Soessogak; other records: Kim H.S.
Arca zanzibarensis Nyst, 1847; ?Anomalocardia pulchella
and Rho B.J. (1971) (Seogwipo); Je J.-G. et al. (1994a)
Dunker, 1868; Anadara thackwayi Iredale, 1927; Anadara
(Munseom); Je J.-G. et al. (2002).
bisenensis Schenk et Reinhart, 1938; Tegillarca granosa
General distribution: Korea, north-western East/Japan
bessalis Iredale, 1939; Anadara (Tegillarca) obessa Kotaka,
Sea (Russia: Peter the Great Bay), Japan (southern
1953; Anadara (Tegillarca) granosa kamakuraensis Noda, 1966.
Hokkaido and southwards, East Sea/Sea of Japan;
southern Kyushu), China (Yellow and East China seas)
Local
(Higo et al., 1999; Qi, 2004; Lutaenko and Noseworthy,
Shinyang, Bomok, Hamo, Daejong, Yeongmeori, Wimi,
distribution:
Udo
(Geomeollae),
Seongsan,
2012; Lee J.-S., 2013; Zhang et al., 2016); subtropical.
Soessogak.
Habitat: In bays in sandy mud, sand, and mud, at
General distribution: Widely distributed Indo-Pacific
depths of 2-60 m (Scarlato, 1981; Higo et al., 1999; Qi,
species: Korea, Japan, and China (including Yellow
2004).
Sea) to Indo-Pacific and eastern coast of Africa
Comments: Uncommon in Jeju Island; harvested
(Kilburn, 1983; Higo et al., 1999; Qi, 2004; Lutaenko,
- 174 -
Korean J. Malacol. 35(2): 149-238 2019
2006;
Huber,
2010;
Lee
J.-S.,
2013);
Genus Didimacar Iredale, 1939
26. Didimacar tenebrica (Reeve, 1844)
tropical-subtropical.
Pl. 1, Fig. A
Habitat: Bays and sheltered shores, estuarine-marine
Iredale,
environment, in mangrove areas and brackish water,
Synonyms:
from the intertidal zone to depth of 10 m (Higo et al.,
koshibensis Hatai et Nishiyama, 1952.
Didimacar
repenta
1939;
Barbatia
1999; Huber, 2010).
Comments: Most specimens from Jeju Island appear
subfossil
and
may
originate
from
the
Local distribution: Udo (Hagosudong), Udo (Sanhosa),
Holocene
Yeongmeori, Seongsan, Hamo, Biyangdo; other records:
deposits. Lee J.-S. (2013) listed this species only for
Lee J.J. and Kim J.C. (1993; as Striarca) (Biyangdo);
Gyeongsangnam-do, Jeollanam-do and Gyeonggi-do in
Lee J.-J. and Hyun J.-M. (1997; as Striarca) (Chujado);
mainland Korea. Lectotype and paralectotype of A.
Lee J.-S. (2013).
granosa are figured by Schenk and Reinhart (1938, pl.
General distribution: Widely distributed Indo-Pacific
4, fig. 1); the holotype and a paratype of A. granosa
species: Korea, Japan (Boso Peninsula and southwards;
kamakuraensis are figured by Noda (1966, pl. 14, figs.
East/Japan Sea) and China (including Yellow Sea) to
1-3). A new record for Jeju Island.
tropical West Pacific, Pakistan, and Persian Gulf (Higo
et al., 1999; Huber, 2010; Zhang et al., 2016; Lee J.-S.,
2013); tropical-subtropical.
Subfamily Noetiinae R. Stewart, 1930
Habitat: Attached to the underside of large rocks, on
Genus Striarca Conrad, 1862
or under stones or dead corals from the intertidal zone
25. Striarca symmetrica (Reeve, 1844)
to depth of 31 m (Huber, 2010). A syntype is figured
Pl. 1, Figs. C, D
Synonyms:
Gabinarca
by Higo et al. (2001, p. 151, fig. B251).
protrita
Iredale,
1939;
Striarca
(Galactella) oyamai Habe, 1953.
Family Parallelodontidae Dall, 1898
Local distribution: Jeju-shi, Hamdeok, Gimnyeong,
Subfamily Parallelodontinae Dall, 1898
Wolpyeong, Sehwa, Haengwon, Handong, Sehwa, Udo
Genus Porterius Clark, 1925
(Hagosudong), Udo (Sanhosa), Jongdal-ri, Seongsan,
27. Porterius dalli (E.A. Smith, 1885)
Pl. 4, Fig. H
Shinyang, Pyoseon, Bomok, Seogundo, Gangjeong,
Jungmun,
Yerae,
Daepyeong,
Hwasoon,
Sagyeri,
Synonyms:
Parallelodon
obliquatus
Yokoyama,
1920;
Hamo, Daejong, Gwakji, Wimi; other records: Lee J.-S.
Cucullaria orientalis Yokoyama, 1922; Parallelodon chihliensis
(2013) (Seongsan, Shinyang, Sagyeri); Je J.-G. et al.
Chao, 1928.
(2002) (Udo); Noseworthy and Choi K.-S. (2010)
(Seogundo).
Local distribution: Hado, Ojo-ri, Seongsan, Hwasoon,
General distribution: Widely distributed Indo-Pacific
Sagyeri,
species: Korea, Japan, and China to tropical West
Biyangdo, Gwideok, Gwakji; other records: Lee J.J.
Songaksan,
Gapado,
Marado,
Chagwi-do,
Pacific and eastern Africa (Higo et al., 1999; Min D.-K.
(1990; as Pseudogrammatodon dalli) (northern coast);
et al., 2004; Qi, 2004; Huber, 2010; Lee J.-S., 2013);
Lee J.J. (1991; as Pseudogrammatodon dalli) (Gapado,
tropical-subtropical.
Marado); Lee J.J. and Hyun J.M. (1992) (eastern coast;
Habitat: Attached byssally to rocks, stones, or coral
Udo); Lee J.J. and Kim J.C. (1993) (Biyangdo); Lee
rubble, in crevices, from the intertidal zone to depth of
J.J. et al. (1995) (Jagwi-do); Je J.-G. et al. (2004a)
68 m (Huber, 2010).
(Munseom); Lee J.-J. and Hyun J.-M. (1997) (Chujado);
Comments: A possible syntype is figured by Higo et al.
Lee J.J. et al. (2001) (Hwasoon); Je J.-G. et al. (2002)
(2001, p. 151, fig. B245).
(Munseom, Beomseom); Lee J.J. and Hyun J.M. (2002)
(Songacksan); Lee J.-S. (2013) (Seongsan, Sagye-ri).
- 175 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
General
distribution:
Southern
Sakhalin
Island
Comments:
Matsukuma
(1979)
mentioned
that
(Russia, Moneron Island), Japan (southern Hokkaido
Glycymeris vestita (Dunker, 1977) may be a junior
and southwards, East/Japan Sea, Seto Inland Sea,
synonym of G. aspersa but later he always used
southern Kyushu), Korea and Yellow Sea (Kafanov,
Dunker’s name (e.g., Matsukuma (1984; 1986a; b);
1991; Higo et al., 1999; Lutaenko and Noseworthy,
Sanada et Matsukuma (1996)). However, subsequent
2012; Min D.-K. et al., 2004; Lee J.-S., 2013; Zhang et
authors adopted Reeve’s name (Higo et al., 1999;
al., 2016); subtropical.
Huber, 2010). The original type locality of G. aspersa,
Habitat: Byssally attached to rocks and stones on
“Sooloo Archipelago” (Sulu, Philippines) (Adams et
gravelly substrate, from depth of 20-300 m (Higo et al.,
Reeve, 1848-1850, p. 76) is considered erroneous and
1999; Huber, 2010).
was corrected to Japan, Honshu (Huber, 2010); the
Comments: A record for the Bering Sea (Higo et al.,
holotype is figured by Higo et al. (2001, p. 152, fig.
1999) is erroneous. The holotype of P. dalli is figured
B264). The holotype of P. vestitus is figured by
by Higo et al. (2001, p. 151, fig. B253); lectotype and
Matsukuma (1986b, pl. 6, fig. 3). “Glycymeris imperialis
paralectotype of P. obliquatus are figured by Oyama
Kuroda,
(1973, pl. 25, figs. 1-4); lectotype and paralectotype of
(Noseworthy et al., 2007) was a misidentification of
1934”
earlier
recorded
for
Jeju
Island
C. orientalis are figured by Oyama (1973, pl. 25, figs.
juvenile specimens of G. aspersa; the former species is
5, 6).
known only from Gangwon-do, Gyeongsangbuk-do and
Chungcheongnam-do in Korea (Lee J.-S., 2013).
29. Glycymeris (Glycymeris) albolineata (Lischke, 1872)
Family Glycymerididae Dall, 1908
Pl. 7, Fig. I
Subfamily Glycymeridinae Dall, 1908
Genus Glycymeris da Costa, 1778
Synonyms: Glycymeris nakamurai Makiyama, 1927; Glycymeris
Subgenus Glycymeris da Costa, 1778
echigoensis Kanehara, 1940.
28. Glycymeris (Glycymeris) aspersa (A. Adams et
Reeve, 1850)
Local distribution: Iho, Udo (Hagosudong), Jongdal-ri,
Pl. 8, Figs. A, B, D-H
Seongsan,
Seopjikoji,
Shinyang,
Wimi,
Bomok,
Synonyms: Pectunculus vestitus Dunker, 1877.
Jungmun, Hwasoon; other records: Noseworthy et al.
Local distribution: Iho, Jeju-shi, Handong, Jongdal-ri,
Je J.-G. et al. (2002); Ko J.-C. et al. (2012) (southern
Seongsan,
coast of Jeju); Lee J.-S. (2013) (Seongsan, Sagyeri).
(2002) (Seongsan); Je J.-G. et al. (1994a) (Munseom);
Seopjikoji,
Shinyang,
Wimi,
Bomok,
Beophwan, Gangjeong, Jungmun, Yerae, Daepyong,
General distribution: Korea (from Gangwon-do to
Hwasoon,
Biyangdo,
Jeju-do); Japan (southern Hokkaido and southwards), ,
Hyeopjae, Gwakji, Daejong; other records: Choi K.-S. et
East China Sea, Taiwan (Matsukuma, 1984; Higo et
Yongmeori,
Sagyei,
Hamo,
al. (2000; as Glycymeris vestita); Suzuki (2010; as
al., 1999; Lee J.S., 2013); subtropical.
Glycymeris vestita) (Jungmun, Hwasoon); Yang H.-S.
Habitat: Fine sand and sandy mud, at depth of 3-50 m
and Choi K.-S. (2011); Lee J.-S. (2013) (Hwasoon).
(Higo et al., 1999; Huber, 2010).
Habitat: Coarse sandy bottom from depth of 3-71 m
Comments: Smaller members of this species may
(Higo et al., 1999; Qi, 2004).
resemble G. aspersa. However, G. albolineata is
General distribution: Japan (southern Hokkaido and
subquadrate, as opposed to the subcircular shape of G.
southwards), Korea (from Gangwon-do to Jeju-do),
aspersa, usually with shades of yellowish-brown with
China: East China and South China seas (Higo et al.,
narrow, incised, white radial lines, and ornamented
1999; Min D.-K. et al., 2004; Qi, 2004; Huber, 2010;
with
Lutaenko and Noseworthy, 2012; Lee J.-S., 2013);
possesses more varied color patterns. Glycymeris
tropical-subtropical.
(Glycymeris) yessoensis (G.B. Sowerby III, 1889) is
- 176 -
fine
punctations.
Also,
G.
aspersa
usually
Korean J. Malacol. 35(2): 149-238 2019
clearly separable from G. albolineata in lacking
Jungmun, Yeongmeori; other records: Je J.-G. et al.
distinct punctations on the outer shell surface and in
(2002; as Oblimopa japonica) (Beomseom, Supseom);
weaker shell convexity; G. albolineata also differs from
Ko J.-C. et al. (2012; as Oblimopa japonica) (southern
G. imperialis in the possesion of a gently-arched, thick
coast of Jeju).
hinge plate and smooth ligamental area; G. imperialis
General distribution: Korea to East China Sea, Japan,
occassionally has small and scattered punctations on
Taiwan (Higo et al., 1999; Huber, 2010); subtropical.
the outer shell surface and brown-tinged adductor
Habitat: Muddy sand at depth of 10-105 m (Huber,
scars, but they are always very distinct in the shell of
2010).
G. albolineata (Matsukuma, 1984). Type material of G.
Comments: This species was often synonymized with
albolineata has not been found (Cosel, 1998). Although
Limopsis forskalii A. Adams, 1863 (e.g., Higo et al.
Matsukuma
to
(1999)), however, Oliver (1981) regards them as
subgenus Veletuceta Iredale, 1931, subsequent authors
separate species. The possible syntypes of both L.
keep it in the subgenus Glycymeris da Costa, 1778
japonica and L. forskalii are illustrated by Higo et al.
(Higo et al., 1999; Huber, 2010; Lee J.-S., 2013).
(2001; p. 152, fig. B261 and B261s, respectively)). As
(1986a;
b)
assigned
this
species
discussed by Huber (2010), the shell of L. forskalii is
more trigonal in outline; the cancellation is stronger
30. Glycymeris (Glycymeris) rotunda (Dunker, 1882)
and regular, and the hinge is more arcuate. The
Pl. 7, Figs. J, K
Synonyms:
Pectunculus
Yokoyama,
nipponicus
1920;
Pectunculus yamakawai Yokoyama, 1922.
ligament pit in L. forskalii is larger, and this species
grows up to 17 mm. Instead, L. japonica is ovate to
oblique, more compressed, shouldered, the cancellation
Local
distribution:
East
of
Seongsan,
south
of
weaker and the hinge plate straighter, and it is usually
Seogwipo, Hwasoon; other records: Lee J.-S. (2013)
smaller than 14.5 mm (Huber, 2010). Oblimopa soyoae
(Jongdal-ri).
Habe, 1953, described from the East/Japan Sea, is a
General distribution: Korea (eastern coast) and Japan
synonym of L. forskalii; the holotype is illustrated by
(Tsugaru Strait and southwards to Okinawa) to East
Higo et al. (2001, p. 152, fig. B261s). An illustration of
China Sea, China (Higo et al., 1999; Qi, 2004; Lee J.S.,
“Oblimopa multistriata (Forskaal, 1775)” from Japan
2013); subtropical.
(Okutani, 2000, pl. 426, fig. 5) represents L. forskalii
Habitat: Fine sand and sandy mud, from depths of
while, in some other works from Japan and China, the
20-300 m (Higo et al., 1999; Huber, 2010). Lectotype
latter species was illustrated as L. japonica. Generally,
and paralectotypes of P. nipponicus are figured by
L. japonica seems to be restricted to the East/Japan
Oyama (1973, pl. 22, fig. 1-4, 7); lectotype and
Sea and to off Taiwan, whereas L. forskalii is more
paralectotypes of P. yamakawai are figured by Oyama
widely distributed, from the South China Sea and
(1973, pl. 22, figs. 5, 6).
Vietnam to Hokkaido, Japan (Huber, 2010). Thus, L.
forskalii, its synonym O. soyoae, and the distinct
Limopsis (Oblimopa) woodwardi Dunker, 1882 (Torres
Superfamily Limopsoidea Dall, 1895
Strait,
Family Limopsidae Dall, 1895
northeastern
Lizard
and
Low
Australia),
Islands,
regarded
Queensland,
previously
as
Genus Limopsis Sassi, 1827
synonyms of L. japonica (e.g., Higo et al., 1999), are
Subgenus Oblimopa Iredale, 1939
now excluded from its Synonyms.
31. Limopsis (Oblimopa) japonica A. Adams, 1863
Subgenus Nipponolimopsis Habe, 1951
Pl. 4, Figs. E, F, I-L
32. Limopsis (Nipponolimopsis) azumana Yokoyama, 1910
Local distribution:
Tongbatarl
(Ojo-ri),
Seogwipo,
Hwasoon, Seongsan,
Sagyei-ri,
Supjikoji,
Synonyms: Limopsis truncata Yokoyama, 1910.
Yerae,
- 177 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Local distribution: Seongsan (Min D.-K. et al., 2004).
Korea, East/Japan Sea including Russian coast (Higo
General distribution: Korea; eastern Japan (Sagami
et al., 1999; Min D.-K. et al., 2004; Huber, 2010;
Bay to Kii Peninsula) (Higo et al., 1999; Okutani,
Lutaenko
et
al.,
2013;
Zhang
et
al.,
2016);
2000); subtropical.
circumboreal-subtropical.
Habitat: In sand, at depth of 100-250 m (Huber, 2010).
Habitat: Byssally attached to rocks, stones, and other
objects, from intertidal zone to depth of 24 m (Huber,
2010).
Order MYTILIDA Férussac, 1822
Comments: The Mediterranean mussel appeared along
Superfamily Mytiloidea Rafinesque, 1815
the Pacific coast of Asia in the 1920-1930s, in Japan,
Family Mytilidae Rafinesque, 1815
presumably either in the area of Kobe or near
Subfamily Mytilinae Rafinesque, 1815
Hiroshima, both on Honshu (Wilkins et al., 1983;
Genus Mytilus L., 1758
Okutani, 2000; Ishida et al., 2005; Furota and
33. Mytilus galloprovincialis Lamarck, 1819
Nakayama, 2010). The first finds of the species in the
East/Japan Sea, according to the museum collections
Pl. 12, Figs. A-D
Synonyms: Mytilus flavus Poli, 1795; Mytilus sagittatus Poli,
in Japan, belong to the post-WWII period (until 1948,
1795; Mytilus hesperianus Lamarck, 1819; Mytilus dilatatus
Fukui Prefecture) (Ishida et al., 2005) At the same
Gray, 1825; Mytilus galloprovincialis angustata Philippi, 1836;
time, M. galloprovincialis appeared on the coast of
Mytilus grunerianus Dunker, 1853; Mytilus succineus Danilo
China, most likely in the 1950s (Lee and Morton,
et Sandri, 1856; Mytilus galloprovicialis falcata Monterosato,
1985; Gosling, 1992; Xu and Zhang, 2008) and also
1884; Mytilus galloprovincialis herculea Monterosato, 1884;
Korea (Yoo M.S., 1992; Je J.-G. et al., 1990; Lee J.-S.
Mytilus lamarckianus Clessin, 1887; Mytilus violaceus Clessin,
et al., 2010). M. galloprovincialis is very close to the
1889
(non
Lamarck,
1819);
galloprovincialis
native western Pacific Mytilus trossulus Gould, 1850
eduliformis Monterosato, 1891; Mytilus orbicularis Pallary,
Mytilus
not only morphologically, but also genetically, and they
1903; Mytilus edulis diegensis Coe, 1945; Mytilus edulis
often form hybrid zones along the Russian coast of the
zhirmunskii Scarlato et Starobogatov, 1979.
East/Japan Sea, Korea, and Japan (Ivanova and
Lutaenko, 1998; Kartavtsev et al., 2005; Brannock et
Local distribution: Gimnyeong, Wolpyeong, Haengwon,
al., 2009). Originally this species was regarded as
Handong,
Yerae,
Mytilus edulis L., 1758 by Asian researchers. A
Hwasoon, Gapado, Marado, Sinchang, Keumneung;
detailed regional Synonyms and distribution records in
other records: Lee J.J. et al. (1989; as Mytilus edulis);
the East/Japan Sea were recently summarized by
Lee J.J. (1991; as Mytilus edulis) (Gapado, Marado);
Lutaenko and Kolpakov (2016).
Sehwa,
Seongsan,
Shinyang,
Lee J.J. and Hyun J.M. (1991; as Mytilus edulis)
(Seogwipo); Je J.-G. et al. (1994a; as Mytilus edulis)
34. Mytilus coruscus Gould, 1861
(Munseom); Lee J.-J. and Hyun J.M. (1997; as Mytilus
edulis) (Chujado); Choi K.-S. et al. (2000); Lee J.J. et
Text-fig. 6A, B
Synonyms: Mytilus unguiculatus Valenciennes, 1858; Mytilus
al. (2001; as Mytilus edulis) (Hwasoon); Je J.-G. et al.
latus Nordmann, 1862 (non Dillwyn, 1817); Mytilus crassitesta
(2002;
Lischke, 1868.
as
Mytilus
edulis)
(Hyopjae,
Yeongmeori,
Munseom, Beomseom); Suzuki (2010) (Hyopjae); Lee
J.-S. (2013); Lee J.U. et al. (2017).
Local distribution: Seogwipo; other records: Kim H.S.
General distribution: Native to the Mediterranean Sea
and Rho B.J. (1971) (Moseulpo, Seongsan, Seogwipo);
and Black Sea; introduced to South Africa, eastern and
Je J.-G. et al. (1994a) (Munseom); Choi K.-S. et al.
western North America, Hawaii, and north-eastern
(2000); Je J.-G. et al. (2002); Lee J.-S. (2013)
Asia;
(Seongsan, Donggwi).
Japan
(from
Hokkaido
to
Okinawa
and
Ogasawara islands), China (Yellow Sea to Hong Kong),
General distribution: Korea; Japan (southern Hokkaido
- 178 -
Korean J. Malacol. 35(2): 149-238 2019
and southwards, East/Japan Sea, East China Sea,
2010) and was illustrated in two works (Lee J.-S. and
Yellow Sea (from Liaoning to Fujian provinces of
Min, 2002, fig. 278; Min et al., 2004, fig. 1193) but
China), Primorye (Russia: north-western East/Japan
then, in a review of Korean Pteriomorphia, it was not
Sea) (Scarlato, 1981; Higo et al., 1999; Qi, 2004; Yang
cited for the Korean fauna (Lee J.-S., 2013). It is
H.-S. and Choi K.-S. (2011); Kolpakov, 2012; Lutaenko
known in Japan in Yamagata Prefecture (East/Japan
and Noseworthy, 2012; Lee J.-S., 2013; Zhang et al.,
Sea), Pacific coast from Chiba to Kagoshima Prefecture,
2016); subtropical.
Tokyo Bay, northern Kyushu, and western Ryukyu
Habitat: Byssally attached to rocks or stones, from
Archipelago (Higo et al., 1999; Lutaenko et al., 2013).
intertidal zone to depth of 20 m (Scarlato, 1981; Higo
et al., 1999; Huber, 2010).
Comments: The widely used name M. coruscus, a
Subfamily Lithophaginae H. et A. Adams, 1857
common mussel in north-east Asia, appears to be a
Genus Lithophaga Röding, 1798
junior synonym of Mytilus unguiculatus Valenciennes,
Subgenus Leiosolenus Carpenter, 1857
1858, an earlier name. According to Article 23.9.1.1 of
36. Lithophaga (Leiosolenus) lischkei (Huber, 2010)
Text-fig. 5B; Pl. 9, Figs. K, L, O, P
ICZN, the name M. coruscus was qualified as nomen
protectum to continue its further usage despite the
Synonyms: Lithophagus curtus Lischke, 1874 (non Lithodomus
priority of M. unguiculatus (Lutaenko, 2005). The
curta Stoliczka, 1870).
holotype of M. coruscus is figured by Lutaenko (2005,
p. 96, fig. 1I-J); a syntype of M. unguiculatus is
Local
figured by Lutaenko (2005, p. 96, figs. A, B).
Daepyeong, Bukchon-ri, Ojo-ri, Seogwipo, Munseom,
Occurrence of this species in Alaska (Lee J.-S., 2013)
Bumseom, Yeongmeori, Sagyeri, Songaksan, Chagwido,
distribution:
Dong-gwi,
Jeju-shi,
Hamdeok,
is erroneous. A rather fragile shell for its size that
Keumneung, Biyangdo, Gwideok; other records: Lee
often cracks when dried.
J.J. (1990; as Lithophaga curta) (northern coast); Kim
Genus Perna Philipsson, 1788
Hyun J.M. (1992; as Lithophaga curta and Leiosolenus
35. Perna viridis (L., 1758)
curta) (eastern coast; Udo); Lee J.J. and Kim J.C.
H.S. et al. (1992; as Lithophaga curta); Lee J.J. and
Pl. 12, Fig. G
(1993; as Lithophaga curta) (Biyangdo); Je J.-G. et al.
Synonyms: Mytilus smaragdinus Gmelin, 1791; Mytilus opalus
(1994a; as Lithophaga curta) (Munseom); Lee J.J. et al.
Lamarck, 1819.
(1995; as Lithophaga curta) (Jagwi-do); Lee J.-J. and
Hyun J.-M. (1997; as Lithophaga curta) (Chujado); Je
Local distribution: Shinyang, Hwasoon.
J.-G. et al. (2002; as Lithophaga curta) (Munseom,
General distribution: Southern Korea and Japan
Beomseom, Supseom, Jigwido); Lee J.J. and Hyun J.M.
(introduced), East China Sea, Taiwan, South China
(2002; as Lithophaga curta) (Songacksan); Ko J.-C. et
Sea,
Indonesia;
al. (2011; as Lithophaga cura) (Daepo, Sincheon,
Indo-Pacific; U.S.A. and Caribbean (introduced) (Higo
Aewol, Woljeong, Jungmun, Hacheon, Haengwon);
et al., 1999; Min D.-K. et al., 2004; Huber, 2010);
Yang H.-S. and Choi K.-S. (2011; as Lithophaga curta);
tropical-subtropical.
Lee J.-S. (2013; as Lithophaga curta) (Seogwipo,
Habitat: Byssally attached to rocks, rocky beaches,
Beomdo, Munseom, Jagwido, Marado); Cho I.-Y. et al.
timbers, also sheltered areas in estuaries; from
(2014; as Lithophaga curta) (Beomseom, Munseom);
Singapore,
Philippines,
and
intertidal zone to depth of 20 m (Higo et al., 1999;
Denis et al. (2014) (Biyangdo, Jigwido, Seogwopo);
Huber, 2010).
Noseworthy et al. (2016) (Keumneung).
Comments: Dead shells only were collected; P. viridis
General distribution: Korea, Japan (including Okinawa
may not be living in Jeju Island. P. viridis is regarded
and Ogasawara Islands) to South China Sea and
as an introduced species to Korea (Lee J.-S. et al.,
Vietnam (Huber, 2010); tropical-subtropical.
- 179 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Habitat: Chemically boring into soft rocks, corals
(Chujado); Je J.-G. et al. (2002; as Modiolus modiolus
(Montipora), limestone, and other shells, from the
difficilis) (Munseom, Beomseom, Jigwido); Lee J.J. and
intertidal zone to depth of 20 m (Huber, 2010); in Jeju,
Hyun J.M. (2002; as Modiolus modiolus difficilis)
this
coral
(Songacksan); Ko J.-C. et al. (2011) (Haengwon); Lee
Psammocora albopicta Benzoni, 2006, in a depth range
J.-S. (2013) (Chagwido, Munseom); Noseworthy and
of 5-10 m (Denis et al., 2014).
Choi K.-S. (2018) (Hamo).
species
lives
in
the
scleractinian
General distribution: Korea, Japan (Tokyo Bay and
37. Lithophaga (Leiosolenus) malaccana (Reeve, 1857)
northwards,
northern
East/Japan
Sea
Hokkaido),
Yellow Sea, northwestern East/Japan Sea (Russia),
Pl. 9, Figs. M, N
Synonyms: Lithophaga reticulata Dunker, 1882; Dactylus
southern Sakhalin, Kurile Islands (Scarlato, 1981;
fauroti Jousseaume, 1888; Lithophaga calcifer Iredale, 1939.
Higo et al., 1999; Min D.-K. et al., 2004; Lee J.-S.,
Local distribution: Munseom, Keumneung, Biyangdo,
Habitat: Attached by byssus to rocks, from the
Jongdal-ri; other records: Noseworthy et al. (2016)
intertidal zone to depth of 85 m (Kurile Islands) or
(Keumneung).
even to 150 m (Scarlato, 1981; Higo et al., 1999).
2013; Zhang et al., 2016).
General distribution: Widely distributed Indo-Pacific
Comments: Volsella difficilis Kuroda et Habe, 1950 is
species: Korea and southern Japan to Indo-Pacific, Red
a junior secondary homonym of Modiola difficilis
Sea,
Australia
(Kleemann,
1984;
Huber,
2010);
Deshayes in Maillard, 1863, and M. kurilensis is a
replacement name (Bernard,
tropical-subtropical.
1983).
Although M.
Habitat: Chemically boring mainly in dead corals but
kurilensis was synonymized with Modiolus modiolus
also in calcareous rocks and shells, from the intertidal
L., 1758 without clear substantiation by many authors
zone to depth of 40 m (Huber, 2010).
(e.g., Coan et al. (2000); Xu and Zhang (2008); Huber
Comments: This species exhibits variability in shape
(2010); Zhang et al. (2016)), mitochondrial DNA data
from rather narrow to broader. L. malaccana occurs
show that M. kurilensis is a valid species (Liu et al.,
occasionally with almost smooth incrustations; however,
2014). Huber (l.c.) mentioned that Russian authors
typically, these are strongly wrinkled (Huber, 2010). A
synonymized these species; however, most Russian
possible syntype of L. malaccana is figured by Higo et
authors did not do that (Scarlato, 1981; Kafanov, 1991;
al. (2001, p. 149, fig. B179).
Lutaenko and Noseworthy, 2012; etc.).
39. Modiolus (Modiolus) auriculatus (Krauss, 1848)
Pl. 9, Figs. A, B
Subfamily Modiolinae G. et H. Termier, 1950
Genus Modiolus Lamarck, 1799
Synonyms: Mytilus (Modiola) hepaticus Gould, 1850; Modiola
Subgenus Modiolus Lamarck, 1799
cymbula Preston, 1908; Modilous auriculatus var. aurantius
Jousseaume in Lamy, 1919; Modiola rufanensis Turton, 1932;
38. Modiolus (Modiolus) kurilensis Bernard, 1983
Volsella agripetus Iredale, 1939.
Pl. 9, Figs. D-F
Synonyms: Volsella difficilis Kuroda et Habe, 1950 (non
Modiola difficilis Deshayes in Maillard, 1863).
Local
distribution:
Jungmun,
Yerae,
Hwasoon,
Yongmeori, Weoljeong, Hamdok, Seongsan, Sagyei,
Local
distribution:
Hamdeok,
Seongsan,
Yerae,
Hamo, Gwakji; other records: Je J.-G. et al. (2002); Ko
Yeongmeori, Bomok, Supseom, Jungmun, Hwasoon,
J.-C. et al. (2011) (Sincheon, Woljeong, Hacheon); Lee
Sagyei, Hamo, Gwideok, Gwakji; other records: Kim
J.-S. (2013); Cho I.-Y. et al. (2014) (Munseom).
H.S. et al., 1992 (as Modiolus difficilis); Je J.-G. et al.
General distribution: Widely distributed Indo-Pacific
(1994a; as Modilous difficilis) (Munseom); Lee J.-J.
species: from Red Sea and eastern Africa to Indonesia,
and Hyun J.M. (1997; as Modiolus modiolus difficilis)
Melanesia, Australia, Micronesia, Polynesia, Philippines,
- 180 -
Korean J. Malacol. 35(2): 149-238 2019
South and East
China seas,
Japan and Korea;
al., 1971; Habe, 1973). Both species were later
Mediterranean Sea (introduction) (Higo et al., 1999; Min
synonymized (M. agripetus and M. nipponicus) with M.
D.-K. et al., 2004; Huber, 2010); tropical-subtropical.
auriculatus (Habe, 1977), and finally M. nipponicus
Habitat: Byssally attached to rocks, in tidepools and in
was again recognized as valid (Habe, 1981; Habe et al.,
crevices, also on dead coral blocks and among rubble,
1986) considered M. agripetus conspecific with M.
from the intertidal zone to depth of 35 m (Huber,
auriculatus. For detailed Synonyms and taxonomic
2010).
discussion, see Lutaenko and Kepel (2017).
40. Modiolus (Modiolus) nipponicus (Oyama, 1950)
41. Modiolus (Modiolus) comptus G.B. Sowerby III, 1915
Pl. 9, Figs. C, G, H
Local distribution: Munseom, Beomseom (Je J.-G. et
Local distribution: Iho, Handong, Hamdeok, Udo
al., 2002); Jeju (Lee J.-S., 2013).
(Sanhosa), Jongdal-ri, Seongsan, Seopjikoji, Shinyang,
General distribution: Korea and Japan, South China
Supseom,
Bumseom,
Yongmeori,
Weoljeong,
Jungmun,
Yerae,
Hwasoon,
Sea, East China Sea, Yellow Sea (Higo et al., 1999;
Sagyei,
Hamo,
Biyangdo,
Huber, 2010; Lee J.-S., 2013); subtropical.
Gwakji; other records: Je J.-G. et al. (2002; as
Habitat: Subtidal, at depth of 2-87 m, sandy mud and
Modiolus agripetus) (Munseom); Noseworthy et al.
sand, also byssally attached to rocks (Huber, 2010).
(2002; as Modiolus difficilis) (Seongsan); Ko J.-C. et al.
The holotype is figured by Higo et al. (2001, p. 148, fig.
(2011; as Modiolus agripetus) (Haengwon); Lee J.-S.
B108).
(2013) (Munseom); Cho I.-Y. et al. (2014) (Beomseom).
General
distribution:
Korea,
Japan
to
Taiwan,
Indonesia, Philippines, South China Sea (Huber, 2010);
Subfamily Brachidontinae Nordsieck, 1969
tropical-subtropical.
Genus Brachidontes Swainson, 1840
Habitat: Byssally attached to rocks and other hard
42. Brachidontes mutabilis (Gould, 1861)
substrates, from the intertidal zone to depth of 40 m
Text-fig. 5G, H; Pl. 10, Figs. A, B, D, E
(Huber, 2010).
Synonyms: Mytilus curvatus Dunker, 1857; Hormomya sinensis
Comments: This species differs from the closely related
Wang, 1983.
M. auriculatus and M. kurilensis by more dorso-ventrally
expanded shell, with almost triangular shape, a
Local
narrower byssal gap, and reddish-pink shades in color,
Hamdeok,
both interior and exterior, and additionally from M.
Seongsan,
kurilensis by the splits at the ends of the periostracal
Gangjeong, Jungmun, Yerae, Daepyeong, Hwasoon,
hairs. J.-S. Lee (2013) indicates that the hairs of the
Yeongmeori,
periostracium in M. auriculatus are split into 3-4
Keumneung, Hyeopjae, Gwakji, Soessogak, Wimi; other
branches whereas, in M. nipponicus, 7-8 branches.
records: Je J.-G. et al. (1994a; as Hormomya mutabilis)
Higo
(1999)
regarded
distribution:
Udo
Jeju-shi,
Handong,
(Hagosudong),
Pyoseon,
Sagyei,
Bomok,
Udo
Seogwipo,
Songaksan,
Sehwa,
(Sanhosa),
Seogundo,
Hamo,
Gosan,
agripetus
(Munseom); Lee J.J. et al. (2001; as Hormomya)
(Iredale, 1939), described from Queensland, Australia,
(Hwasoon); Je J.-G. et al. (2002; as Hormomya)
as a distinct species, and M. nipponicus as its synonym,
(Beophwan, Bomok); Noseworthy and Choi K.-S. (2010;
and indicated also that some authors combine M.
as Hormomya) (Seogundo); Ko J.-C. et al. (2011)
et
al.
Modiolus
agripetus and a close species, M. auriculatus. This
(Aewol, Woljeong, Hacheon, Haengwon); Yang H.-S.
confusion appears to have arisen when Japanese
and Choi K.-S. (2011; as Hormomya); Lee J.-S. (2013;
malacologists first considered M. nipponicus as a
as Hormomya) (Hwasoon, Seongsan, Sagye-ri); Lee
separate species (Habe, 1951; Yamamoto and Habe,
J.U. et al. (2017).
1958), then as a synonym of M. agripetus (Kuroda et
General distribution: Korea and Japan, Philippines,
- 181 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
South China Sea, Vietnam, East China Sea, Okinawa
Jungmun, Yerae, Daepyeong, Hwasoon, Yongmeori,
(Higo et al., 1999; Min D.-K. et al., 2004; Huber, 2010);
Sagyeri, Songaksan, Gapado, Marado, Hamo, Gosan,
tropical-subtropical.
Chagwi-do,
Habitat: Byssally attached to rock, shells, and stones
records: Lee J.J. et al. (1989); Lee J.J. (1990) (northern
in intertidal zone (Huber, 2010).
coast); Lee J.J. (1991) (Gapado); Lee J.J. and Hyun
Keumneung,
Biyangdo,
Gwakji;
other
Comments: This species has been observed using its
J.M. (1991) (Seogwipo); Kim H.S. et al., 1992; Lee J.J.
byssal threads to trap muricid predators (Ishida and
and Hyun J.M. (1992) (eastern coast; Udo); Lee J.J. and
Iwasaki, 1999).
Kim J.C. (1993) (Biyangdo); Je J.-G. et al. (1994a)
(Munseom); Lee J.J. et al. (1995) (Chagwi-do); Lee J.-J.
and Hyun J.M. (1997) (Chujado); Choi K.-S. et al.
Subfamily Septiferinae Scarlato et Starobogatov, 1979
(2000); Je J.-G. et al. (2002) (Youngunchon, Yeongmeori,
Genus Septifer Récluz, 1848
Beophwan,
Subgenus Septifer Récluz, 1848
Beomseom, Supseom, Jigwido); Lee J.J. and Hyun J.M.
43. Septifer (Septifer) bilocularis (L., 1758)
(2002)
Biyangdo,
Udo,
(Songacksan);
Sinchon,
Suzuki
(2010)
Munseom,
(Jungmun,
Hwasoon, Gwakji); Yang H.-S. and Choi K.-S. (2011);
Pl. 11, Figs. A-E
Synonyms: Mytilus nicobaricus Röding, 1798; Mytilus septulifer
Cha J.-H. et al. (2013) (north-eastern coast of Jeju);
Menke, 1830; Tichogonia wiegmannii Küster, 1841; Tichogonia
Lee
kraussii Küster, 1841; Modiola subtriangularis Turton, 1932.
Munseom, Seongsan, Hwasoon); Cho I.-Y. et al. (2014)
J.-S.
(2013)
(Jeju
Harbour,
Udo,
Jagwido,
(Gapado, Beomseom, Munseom); Lee J.U. et al. (2017).
Local distribution: Udo (Hagosudong), Udo (Sanhosa),
General distribution: Korea and Japan to Vietnam and
Udo (Geomeollae), Seongsan, Wimi, Jungmun, Yerae,
South China Sea (Higo et al., 1999; Min D.-K. et al.,
Hwasoon,
Yongmeori,
Sagyei,
Songaksan,
Gwakji,
2004; Huber, 2010); tropical-subtropical.
Hamo; other records: Lee J.-S. (2013) (Seongsan).
Habitat: Byssally attached to rocks, often in clumps,
General distribution: Widely distributed Indo-Pacific
on exposed rocky shores in high intertidal zone
species, from southern and eastern Africa to Indonesia,
(Huber, 2010).
Melanesia, Australia, Micronesia, Philippines, South
China Sea, Vietnam, East China Sea, Okinawa, Korea
45. Septifer (Mytilisepta) keenae Nomura, 1936
Pl. 11, Figs. F-I
and Japan (Higo et al., 1999; Min D.-K. et al., 2004;
Huber, 2010); tropical-subtropical.
Habitat: Byssally attached to rocks among algae, near
Local distribution: Dong-gwi, Iho, Hwabuk, Hamdeok,
low tide mark, from intertidal zone to depth of 35 m
Bukchon-ri,
(Huber, 2010).
Handong, Hado, Udo (Hagosudong), Udo (Sanhosa), Udo
Gimnyeong,
(Geomeollae),
Jongdal-ri,
Wolpyeong,
Daejong, Ojo-ri,
Haengwon,
Seongsan,
Subgenus Mytilisepta Habe, 1951
Seopjikoji, Shinyang, Pyoseon, Namwon, Wimi, Bomok,
44. Septifer (Mytilisepta) virgatus (Wiegmann, 1837)
Seogwipo,
Beophwan,
Gangjeong,
Daepo,
Jungmun,
Yerae, Hwasoon, Yongmeori, Sagyei, Songaksan, Gapado,
Text-fig. 5A, C, D; Pl. 10, Figs. C, F-I
Synonyms: Dreissena purpurascens Benson in Cantor, 1842;
Marado,
Hamo,
Chagwido,
Keumneung,
Biyangdo,
Septifer crassus Dunker, 1853; Septifer herrmannseni Dunker,
Hyeopjae, Gwideok, Weoljeong, Gwakji; other records:
1853.
Lee J.J. and Jwa Y.W. (1988); Lee J.J. et al. (1989);
Lee J.J. (1990) (northern coast); Lee J.J. (1991)
Local distribution: Iho, Jeju-shi, Samyang, Hamdeok,
(Gapado, Marado); Lee J.J. and Hyun J.M. (1991)
Bukchon-ri,
Jongdal-ri,
(Seogwipo); Kim H.S. et al., 1992; Lee J.J. and Hyun
Seongsan, Ojo-ri, Shinyang, Pyoseon, Wimi, Bomok,
J.M. (1992) (eastern coast; Udo); Lee J.J. and Kim J.C.
Supseom, Seogwipo, Munseom, Beomseom, Gangjeong,
(1993) (Biyangdo); Je J.-G. et al. (1994a) (Munseom);
Hado,
Udo
(Sanhosa),
- 182 -
Korean J. Malacol. 35(2): 149-238 2019
Lee J.J. et al. (1995) (Chagwido); Lee J.-J. and Hyun
subtropical.
J.M. (1997) (Chujado); Je J.-G. et al. (2002; including
Habitat: Sand and gravel, at depth of 100-300 m
excisus) (Youngunchon, Yeongmeori, Shinyang, Sanho,
(Huber, 2010).
Hado, Gimyong, Munseom, Beomseom, Supseom); Lee
Comments: This species was not listed for the Korean
J.J. and Hyun J.M. (2002) (Songacksan); Noseworthy
fauna by J.-S. Lee (2013).
et al. (2002) (Seongsan); Noseworthy and Choi K.-S.
(2010; as Septifer excisus) (Seogundo); Suzuki (2010)
(Hwasoon); Ko J.-C. et al. (2011); Yang H.-S. and Choi
Subfamily Musculinae Iredale, 1939
K.-S. (2011); Lee J.-S. (2013) (Seogwipo, Sehwa,
Genus Gregariella Monterosato, 1884
Shinyang, Seongsan, Sagye-ri, Hwasoon); Cho I.-Y. et
47. Gregariella barbata (Reeve, 1858)
al. (2014) (Beomseom); Noseworthy et al. (2016)
Synonyms: Modiolaria arcuata Gould, 1861; Lithodomus laniger
(Keumneung); Noseworthy and Choi K.-S. (2018)
Reeve, 1858; Modiolaria barbata Angas, 1868.
(Hamo).
Sea,
Local distribution: Hwasoon (Min D.-K. et al., 2004; as
East/Japan (north to Hokkaido and Peter the Great
Trichomusculus semigranatus); Daepo, Aewol, Hacheon
General
distribution:
Korea,
Japan,
Yellow
Bay), East China Sea (Scarlato, 1981; Higo et al.,
(Ko J.-C. et al., 2011; as Trichomusculus semigranatus);
1999; Min D.-K. et al., 2004; Huber, 2010; Zhang et al.,
Hangaechang,
2016); subtropical.
Trichomusculus semigranatus).
Habitat: Byssally attached to rocks, from intertidal
General distribution: Korea and Japan to Thailand,
zone to depth of 30 m (Huber, 2010).
Philippines, South China Sea, Melanesia, Australia,
Comments: J.-S. Lee (2013) stated that all records of
and New Zealand (Higo et al., 1999; Min D.-K. et al.,
Septifer
excisus
(Wiegmann,
1837)
from
Korea
Munseom
(Lee
J.-S.,
2013;
as
2004; Huber, 2010); tropical-subtropical.
including Min D.-K. et al. (2004) are instead S. keenae.
Habitat: Under stones, on rocks, nestling in cavities of
The shell of S. excisus appears to have narrow but
larger shells, also embedded in sandy clay and mud,
distinct radial, often bifurcating, ribs which exhibit
from intertidal zone to depth of 90 m (Huber, 2010).
various degrees of beading; the edge of the septum
Comments: This species was known in regional literature
contains a concavity (notch) of varying depth; although
(e.g., Min D-K. et al., 2004; Lee J.-S. (2013)) as
some illustrations of this species show no notch; the
“Trichomusculus
anterior margin is more rounded than in S. keenae,
Gregariella semigranata (Reeve, 1858), a Mediterranean-
and the ribs of the latter are generally somewhat
Atlantic species).
semigranatus
(Reeve,
1858)”
(=
wider and smoother, and the anterior margin is more
angular. The edge of the septum in S. keenae exhibits
48. Gregariella coralliophaga (Gmelin, 1791)
no
Synonyms: Modiola semen Lamarck, 1819; Modiola divaricata
concavity.
Jeju
records
of
S.
excisus
(e.g.,
Philippi, 1847; Lithodomus gossei Reeve, 1858; Lithodomus
Noseworthy et al. (2007)) are actually S. keenae.
argenteus Reeve, 1858; Gregariella opifex Dall, 1889; Tibialectus
otteri Iredale, 1939.
Subfamily Crenellinae Gray, 1840
Genus Exosiperna Iredale, 1929
Local distribution: Seongsan (Min D.-K. et al., 2004);
46. Exosiperna kuroharai Habe, 1961
Jeju (Lee J.-S., 2013).
General distribution: Widely distributed Indo-Pacific
Local distribution: Seongsan (Min D.-K. et al., 2004);
species (eastern Africa to Australia, Philippines, South
Jeju (Lee J.U. et al., 2017; as Exsiperna kuroharai).
China Sea, East China Sea, Yellow Sea, Korea, Japan)
General distribution: Korea, Japan to Okinawa (Higo
and also Atlantic (North Carolina to Brazil) (Huber,
et al., 1999; Min D.-K. et al., 2004; Huber, 2010);
2010); tropical-subtropical.
- 183 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Habitat: Chemically boring in corals (Porites sp.),
from intertidal zone to depth of 70 m (Huber, 2010).
limestone, and shells; from the intertidal zone to depth
Comments: “Musculus nanus (Dunker, 1857)” known in
of 20 m (Huber, 2010).
regional literature (e.g., Okutani, 2000; Min D.-K. et
Comments: This species exhibits rare, “cosmopolitan”
al., 2004; Lee J.-S., 2013), and exhibiting some
distribution but it needs a molecular confirmation
variability in shape, is actually M. viridulus; M. nanus
(Huber, 2010).
is restricted to southern Australia (Huber, 2010); a
syntype of the latter is figured by Moolenbeek (2009,
figs. 8, 9).
Genus Musculus Röding, 1798
Subgenus Musculus Röding, 1798
Subgenus Modiolarca Gray, 1842
49. Musculus (Musculus) pusio (A. Adams, 1862)
51. Musculus (Modiolarca) cupreus (Gould, 1861)
Text-fig. 6C; Pl. 8, Figs. I-K
Synonyms: Modiolaria quadrula Gould, 1861; Musculus
Local
distribution:
Udo
(Hagosudong,
Sanhosa),
neglectus Kuroda, 1941.
Seongsan, Wolpyeong, Bomok, Munseom, Hwasoon,
Yeongmeori, Gwakji, Woljeong; other records: Je J.-G.
Local distribution: Munseom (Je J.-G. et al., 2002);
et al. (2002) (Munseom, Beomseom); Noseworthy and
Seongsan (Min D.-K. et al., 2004).
Choi K.-S. (2010; as Musculus nanus) (Seogundo); Ko
General distribution: Korea and Japan, East China
J.-C. et al. (2011; as Musculus viridulus) (Sincheon,
Sea, Taiwan, Yellow Sea (Huber, 2010); subtropical.
Aewol, Haengwon); Lee J.-S. (2013) (Hwasoon; as
Habitat: Subtidal, at depth of 1-50 or 150 m, byssally
Musculus viridulus); Lee J.U. et al. (2017); Noseworthy
attached among ascidians or in rock crevices (Huber,
and Choi K.-S. (2018; as Musculus nanus) (Hamo).
2010).
General distribution: Japan and Korea, East China
Comments: “Musculus cumingiana (Reeve, 1857)” (Min
Sea, Yellow Sea (Huber, 2010); subtropical.
D.-K. et al. (2004)) is M. cupreus (Lee J.-S., 2013);
Habitat: Byssally attached to rock, gravel and seaweed
however, the latter author did not list Jeju among the
in soft mud, from intertidal zone to depth of 20 m
localities of M. cupreus in Korea. The holotype is
(Huber, 2010).
figured by Higo et al. (2001, p. 149, fig. B156).
Comments: J.-S. Lee (2013) regarded a Jeju record of
this species (Min D.-K. et al., 2004, fig. 1218) as a
misidentification of Musculus viridulus (H. Adams,
Subfamily Arcuatulinae Scarlato et Starobogatov, 1979
1871). Alernatively, the illustration in Min D.-K. et al.
Genus Arcuatula Jousseaume in Lamy, 1919
(l.c., fig. 1219) of “M. virudulus” is M. pusio.
52. Arcuatula senhousia (Benson in Cantor, 1842)
Synonyms: Modiola bellardiana Tapparone-Canefri, 1874;
Modiola aquarius Grabau et King, 1928.
50. Musculus (Musculus) viridulus (H. Adams, 1871)
Synonyms: Modiolaria miranda Smith, 1884.
Local distribution: Shinyang, Gangjeong, Jungmun,
Local distribution: Hwasoon (Lee J.-S., 2013; as
Hwasoon,
Musculus nanus).
Chagwido, Keumneung, Biyangdo; other records: Lee
General distribution: Widely distributed Indo-Pacific
J.J. et al. (1989; as Musculus senhausia); Lee J.J. and
species, from Red Sea to Indonesia, Australia, Melanesia,
Hyun J.M. (1992) (eastern coast; Udo); Lee J.J. and
Polynesia, Philippines, Thailand, South China Sea,
Kim J.C. (1993; as Musculus senhausia) (Biyangdo); Je
Vietnam, East China Sea, Korea and Japan (Min D.-K. et
J.-G. et al. (1994a; as Musculus senhousia) (Munseom);
Songaksan,
Lee
(Chagwi-do); Lee J.-J. and Hyun J.-M. (1997; as
sand grains, also nestling on rocks and on large shells,
Musculus senhousia) (Chujado); Lee J.J. et al. (2001;
al.
as
Gosan,
Habitat: Rocky and sandy areas, byssally attached to
et
(1995;
Gapado,
al., 2004; Huber, 2010); tropical-subtropical.
- 184 -
J.J.
Sagyei,
Musculus
senhausia)
Korean J. Malacol. 35(2): 149-238 2019
as Musculus senhausia) (Hwasoon); Je J.-G. et al.
by
Kimura
(1996);
according
to
him,
its
shell
(2002; as Musculus senhousia) (Youngunchon, Hado);
morphology is similar to Xenostrobus balani Ockelmann,
Lee J.J. and Hyun J.M. (2002; as Musculus senhausia)
1983 and Xenostrobus mangle Ockelmann, 1983 from
(Songacksan); Lee J.-S. (2013).
south-eastern
General distribution: North-western East/Japan Sea,
association of X. atratus, as well as other allied
Sea of Okhotsk (Aniva Bay: Sakhalin Island), Japan,
species, with the genus Limnoperna Rochebrune, 1862
Yellow Sea, Korea, China, Vietnam, Thailand, and
(Beu, 2006) was later abandoned (Beu, 2012). X.
Asia
(Ockelmann,
1983).
Previous
Singapore; invasive species in the Mediterranean Sea,
atratus is the type species of Vignadula Kuroda et
Australia, New Zealand and the Pacific coast of North
Habe in Kuroda, Habe et Oyama, 1971 which is
America (Scarlato, 1981; Coan et al., 2000; Min D.-K.
believed to be a synonym of Xenostrobus Wilson, 1967
et al., 2004; Huber, 2010); subtropical-lowboreal.
(Ockelmann, 1983; Kimura, 1996; Beu, 2006).
Habitat: Sheltered bays, burrowing in soft mud,
muddy
gravel,
also
among
eelgrass,
occasionally
building nests by byssal threads, from intertidal zone
Order OSTREIDA Férussac, 1822
to depth of 20 m (Scarlato, 1981; Huber, 2010).
Superfamily Ostreoidea Rafinesque, 1815
Comments: A syntype of A. senhousia is figured by
Family Ostreidae Rafinesque, 1815
Higo et al. (2001, p. 149, fig. B159).
Subfamily Ostreinae Rafinesque, 1815
Genus Ostrea L., 1758
54. Ostrea denselamellosa Lischke, 1869
Genus Xenostrobus Wilson, 1967
Pl. 13, Fig. I
53. Xenostrobus atratus (Lischke, 1871)
Synonyms: Ostrea auriculata G.B. Sowerby II, 1871; Ostrea
Pl. 9, Figs. I, J
Synonyms: Modiola aterrima Dall, 1871.
multicostata G.B. Sowerby II, 1871 (non Deshayes, 1832);
Ostrea multiradiata G.B. Sowerby II, 1871; Ostrea tubercularis
Local
distribution:
Hamdeok,
Handong,
Ojo-ri,
Yongmeori, Hyeopjae; other records: Lee J.-S. (2013)
G.B. Sowerby II, 1871; Ostrea cristatella Lamy, 1929; Ostrea
jubata Lamy, 1929.
(Udo).
General distribution: South Korea (all provinces except
Local distribution: Hamdeok, Hado, Jongdal-ri, Ojo-ri,
for Gangwon-do), Japan (northeastern Honshu and
Seongsan, Seogwipo, Jungmun, Hwasoon, Gapado,
southwards, western Kyushu), Yellow Sea, northern
Marado, Gosan, Chagwido, Biyangdo, Gwideok; other
South China Sea (Higo et al., 1999; Okutani, 2000; Qi,
records: Lee J.-J. et al. (1989); Lee J.J. (1990)
2004;
(northern coast); Lee J.J. (1991) (Gapado); Lee J.J. and
Min
et
al.,
2004;
Zhang
et
al.,
2016);
tropical-subtropical.
Hyun J.M (1991) (Seogwipo); Lee J.J. and Hyun J.M.
Habitat: Marine-estuarine, sheltered situations; byssally
(1992) (eastern coast); Lee J.J. and Kim J.C. (1993)
attached to rocks or shells, from the high intertidal
(Biyangdo); Je J.-G. et al. (1994a) (Munseom); Lee J.J.
zone to depth of 1 m (Huber, 2010).
et al. (1995) (Chagwido); Lee J.-J. et al. (1997)
Comments: The holotype of M. aterrima is figured by
(Chujado); Lee J.J. et al. (2001) (Hwasoon); Je J.-G. et
Higo et al. (2001, p. 148, fig. B127s), and its
al. (2002) (Munseom, Beomseom, Supseom); Lee J.J.
morphology does correspond to X. atratus; syntypes of
and Hyun J.M. (2002) (Songacksan).
X. atratus were figured by Lutaenko and Chaban
General distribution: Japan (Mutsu Bay, Honshu and
(2016, figs. 1A-D; 2A-C). The latter name has priority
southwards), Yellow Sea, Korea, East China Sea,
over M. aterrima, as Lischke’s (1871) paper was
China (to Taiwan and Hong Kong) to Philippines,
published in January, while Dall’s (1871) paper was
Indonesia, Vietnam and Thailand (Higo et al., 1999;
published on November 2, 1871 (Coan et al., 2000). The
Min D.-K. et al., 2004; Qi, 2004; Huber, 2010; Zhang et
morphology and anatomy of X. atratus were described
al., 2016); tropical-subtropical.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Habitat: Left valve cemented to rocks and pebbles, on
Synonyms: Lopha theca Röding, 1798; Ostrea haliotidaea
sand and gravel bottoms from the intertidal zone to
Lamarck, 1819; Ostrea erucella Lamarck, 1819; Ostrea
depth of 35 m (Higo et al., 1999; Huber, 2010).
fucorum Lamarck, 1819; Ostrea labrella Lamarck, 1819;
Comments: A possible syntype of O. denselamellosa is
?Ostrea sinuata Lamarck, 1819; Ostrea affinis G.B. Sowerby
figured by Higo et al. (2001, p. 155, fig. B395).
II, 1871; Ostrea dubia G.B. Sowerby II, 1871; Ostrea
55. Ostrea circumpicta Pilsbry, 1904
Lamarck, 1819); Ostrea dalli Lamy, 1930 (nom. nov. pro
cognata Jousseaume, 1888; Ostrea serra Dall, 1914 (non
Ostrea serra Dall, 1914 non Lamarck, 1819); Ostrea bresia
Pl. 13, Figs. A-D
Iredale, 1939.
Local distribution: Iho, Jeju-shi, Jongdal-ri, Seongsan,
Supseom, Seogwipo, Munseom, Bumseom, Jungmun,
Local distribution: Marado (Qi and Choe, 2000);
Daepyeong, Hwasoon, Sagyei, Keumneung, Biyangdo,
Seongsan (Min D.-K. et al., 2004).
Yerae, Gwakji, west of Pyeoson, Soesoggak; other
General distribution: Kii Penisula, Japan, Korea to
records: Choi K.-S. et al. (2000); Lee J.J. et al. (2001)
Taiwan,
(Hwasoon); Je J.-G. et al. (2002) (Munseom, Supseom);
Australia, the Philippines, Indonesia to the Indo-West
Hainan,
South
and
East
China
seas,
Noseworthy et al. (2002) (Seongsan); Cho I.-Y. et al.
Pacific and eastern Pacific (from Mexico to Colombia
(2014) (Bumseom, Munseom); Noseworthy et al. (2016;
and Galapagos Islands), eastern Mediterranean (Qi,
as Striostrea) (Keumneung).
2004; Lam and Morton, 2009; Huber, 2010; Coan and
General distribution: Korea and Japan (Mutsu Bay,
Valentich-Scott, 2012); tropical-subtropical.
Honshu and southwards), East China Sea, Taiwan
Habitat: Cemented by finger-like processes to stems of
(Higo et al., 1999; Inaba et al., 2004; Lee J.-S., 2016);
gorgonians, sea fans, rocks or corals, from depths of
subtropical.
5-50 m (Huber, 2010).
Habitat: Exposed shores; attached to non-calcareous
Comments: Biogeographically, it is likely that D.
rocks, forming dense banks, from intertidal zone to
folium immigrated through the Suez Channel and is
depth of 30 m (Huber, 2010).
now established in the eastern Mediterranean (Huber,
Comments: Salvi and Mariottini (2016), based on
molecular
analysis,
re-classified
this
species
2010).
as
belonging to the genus Ostrea L., 1758 instead of
57. Dendostrea sandvichensis (G.B. Sowerby II, 1871)
previous association with Striostrea Vialov, 1936
Synonyms:
(Harry, 1985; Higo et al., 1999; Huber, 2010). This is
crenulifera G.B. Sowerby II, 1871; Ostrea albatra Jousseaume
in accord with the viewpoint of Inaba et al. (2004). O.
in Lamy, 1925; Ostrea kupua Dall, Bartsch et Rehder, 1938;
circumpicta is a quite heavy, solid species, with the
Ostrea nomades Iredale, 1939.
Ostrea
semiplicata
Küster,
1868;
Ostrea
typical metallic-iridescent internal sheen, often in
whitish-grey/green, often reddish around the borders;
Local distribution: Jeju (Min D.-K. et al., 2004); Ko
the noduled upper border denticulations are also
J.-C. et al. (2011; as Dendostrea crenulifera) (Daepo,
typical; juveniles have a particularly large scar; in well
Aewol);
preserved specimens, irregularly divergent ribs are
crenulifera) (Bumseom, Munseom).
Cho
I-Y.
et
al.
(2014;
as
Dendostrea
found (Huber, 2010). A syntype is figured by Higo et
General distribution: Japan to South China Sea,
al. (2001, p. 155, fig. B417).
Indonesia,
Vietnam
and
western
Indo-Pacific
(to
eastern Africa, Red Sea and Persian Gulf), eastern
Mediterranean; Hawaii, ?Mexico (Gulf of California)
Subfamily Lophinae Vyalov, 1936
(Higo et al., 1999; Galil, 2007; Huber, 2010; Coan and
Genus Dendrostrea Swainson, 1835
Valentich-Scott, 2012); tropical-subtropical.
56. Dendostrea folium (L., 1758)
Habitat: Attached to other shells, on pilings, rocky
- 186 -
Korean J. Malacol. 35(2): 149-238 2019
substrate, or coral rubble, from the intertidal zone to
Mediterranean Sea, Canada, Brazil, Argentina, Angola,
depth of 40 m (Huber, 2010).
Namibia), Australia, New Zealand, Hawai (Scarlato,
Comments: This species has been long known in
1981; Higo et al., 1999; Coan et al., 2000; Min D.-K. et
literature as Dendostrea crenulifera (G.B. Sowerby II,
al.,
1871) (see extensive Synonyms in Inaba et al. (2004)).
Valentich-Scott,
2004;
Qi,
2004;
2012;
Huber,
2010;
Lutaenko
and
Coan
and
Noseworthy,
The holotype of O. crenulifera is figured by Higo et al.
2012); subtropical-lowboreal.
(2001, p. 155, fig. B403).
Habitat: Sheltered, often brackish, bays with low
salinity; on rocks, often in mud, from intertidal zone to
depth of 30 m (Higo et al., 1999; Huber, 2010).
Morphological identification of
species
Subfamily Crassostreinae Scarlato et Starobogatov, 1979
Comments:
Genus Crassostrea Sacco, 1897
belonging to this genus is complicated and should be
Subgenus Magallana Salvi et Mariottini, 2016
complemented by molecular methods. Close species to
58. Crassostrea (Magallana) gigas (Thunberg, 1793)
C. gigas are Crassostrea lugubris (Sowerby II, 1871)
(blackscar
Pl. 15, Figs. A-F
Synonyms:
Ostrea
laperousii
Schrenck,
1861;
Ostrea
oyster),
C.
angulata
(Lamarck,
1819)
(Portuguese oyster), C. hongkongensis Lam et Morton,
talienwhanensis Crosse, 1862; Ostrea rostralis G.B. Sowerby
2003 (Hong Kong cultured oyster), C. ariakensis
II, 1871 non Lamarck, 1819; Ostrea cymbaeformis G.B.
Fujita, 1913 (Suminoe oyster) and C. nippona (Seki,
Sowerby II, 1871; Ostrea virginica var. lusitanica Osorio,
1934) (Japanese oyster). Lam and Morton (2004) noted
1916;
that
Ostrea
gravitesta
Yokoyama,
1926;
Dioeciostrea
C.
angulata
can
not
be
morphologically
hispaniola Orton, 1928; Ostrea chemnitzii var. elongata
differentiated from the closest C. gigas based on
Grabau et King, 1928; ?Ostrea gigas var. tientsiensis
anatomy and shell whereas Batista et al. (2009)
Grabau et King, 1928; Ostrea posjetica Razin, 1934; Ostrea
observed differences in shell shape and muscle scar
complanata Fenaux, 1944; Lopha (Ostreola) posjetica beringi
pigmentation between adults of the two taxa. However,
Vyalov, 1946; Lopha (Ostreola) posjetica zawoikoi Vyalov,
the existence of C. angulata in waters of northern
1946; Lopha (Ostreola) posjetica newelskyi Vyalov, 1946.
China, and thus in Korea, is doubtful and, most likely,
this species does not live north of the Yangtze River
Local distribution: Hamdeok, Wolpyeong, Jongdal-ri,
estuary but is widespread from Zhejiang Province to
Seopjikoji,
Beophwan,
Hainan Island (Wang et al., 2008, 2010). Later, Sekino
Jungmun, Yerae, Daepyeong, Hwasoon, Yongmeori,
and Yamashita (2013) and Sekino et al. (2016)
Shinyang,
Wimi,
Bomok,
Sagyei, Hamo, Hyeopjae, Gwideok, Gwakji, Gangeong
revealed the presence of a hitherto-unrecognized
Harbour, Soesoggak; other records: Lee J.J. and Jwa
population of the Portuguese oyster C. angulata in
Y.W. (1988); Lee J.-J. et al. (1989); Lee J.J. (1990)
Okinawa, Japan and on the coast of Shikoku Island,
(northern coast); Lee J.J. and Hyun J.M. (1991)
Japan. In the latter location, they also confirmed the
(Seogwipo); Kim H.S. et al. (1992); Je J.-G. et al.
distribution of a recently recognized oyster species,
(1994a) (Munseom); Choi K.-S. et al. (2000); Je J.-G. et
Crassostrea dianbaiensis Xia, Wu, Xiao et Yu, 2014)
al. (2002); Suzuki (2010) (Hamdeok, Jungmun); Cha
which makes it possible to regard its distributional
J.-H. et al. (2013) (north-eastern coast of Jeju); Lee
range as wider being a tropical-subtropical species.
J.U. et al. (2017).
Wang et al. (2010) found that C. gigas and C. angulata
General distribution: Native range is from Russian
are easily hybridized and genetically are closely
coast of the East/Japan Sea (north to Amur River
related, so they suggest accepting C. angulata as a
estuary), Sakhalin Island, south Kurile Islands, Japan
subspecies of C. gigas. In other words, the problem
(Hokkaido and southwards), Korea, Yellow Sea, China
with the validity of C. angulata is not resolved even
to South China Sea; introduced to the Pacific coast of
after
North
without further examination, regard this name as
America
and
Atlantic
Ocean
(Europe,
- 187 -
molecular
studies,
whereas
other
authors,
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
valid (Boudry et al., 1998; O’Foighil et al., 1998; Lam
as distinct genera as they form two highly supported
and Morton, 2004; Batista et al., 2009; Hsiao et al.,
monophyletic clades in all phylogenetic studies; they
2016). Inaba et al. (2004) and Huber (2010) treated C.
have a strictly allopatric distribution in different
angulata as a synonym of C. gigas. Recent analyses of
oceans. The genetic divergence between them at
mitochondrial cytochrome oxidase I (COI) sequences
mitochondrial and nuclear genes is similar or even
from a large number of specimens of Crassostrea
higher than genetic distance observed between genera
collected from 12 locations along Taiwanese and
belonging
southern Chinese coastlines confirm that all samples
Ostreinae and Lophinae genera. According to divergent
were the Portuguese oyster, C. angulata, rather than
time
to
estimates
different
based
subfamilies,
on
i.e.
mitogenome
between
data,
the
C. gigas (Hsiao et al., 2016). As for other species of the
divergence between Asian and American Crassostrea is
complex, C. lugubris (a synonym of Crassostrea
as ancient as 83 million years. Based on this and other
bilineata (Röding, 1798), according to Huber (2010)) is
molecular evidence, they proposed a new genus
known from China, Philippines, Thailand (Lam and
Magallana Salvi et Mariottini, 2016 that includes
Morton, 2004) or very widely in the Indo-Pacific (from
Asian species previously assigned to Crassostrea (Salvi
Aden and Madagascar to China and Vietnam) (Huber,
and Mariottini, 2016); however, we prefer to regard it
2010). C. hongkongkensis, in southern China (Lam and
as a subgenus of Crassostrea. The genus name
Morton, 2004; Huber, 2010); C. ariakensis, from
Magallana Salvi, Macali et Mariottini, 2014 (Salvi et
Vietnam to Yellow Sea, Korea, and Japan (western
al., 2014) is not available under ICZN rules (Art. 13.1).
Kyushu, Ariake Bay, Osaka Bay, Honshu (Chiba
Prefecture)) (Higo et al., 1999; Huber, 2010). C.
59. Crassostrea (Magallana) nippona (Seki, 1934)
ariakensis, C. hongkongensis, and C. nippona are
closely related species based on molecular data (Reece
Local distribution: Jungmun (Min D.-K et al., 2004);
et al., 2008). The presence of more than two species of
Munseom (Je
J.-G.
et
al.,
2002;
as Crassosrea
Crassostrea in Jeju Island or more than three species
“nipponica”); Jeju (Lee J.U. et al., 2017; as Crassosrea
in Korea (Lee J.-S., 2016) is possible. Hong J.-S. et al.
“nipponica”).
(2012) identified the Kumamoto oyster, Crassostrea
General distribution: Korea, Japan (Honshu: Mutsu
sikamea (Amemiya, 1928) a sibling species of C. gigas,
Bay and southwards to southern Kyushu), Yellow Sea,
in Suncheon Bay, located in the southern Korean
(Higo et al., 1999; Lee J.-S. and Min, 2002; Zhang et
Peninsula, This species which ranges from the South
al., 2016]; subtropical.
China Sea to Japan, and has been introduced to
Habitat: Left valve cemented to rocks, from the
Oregon and Australia (Huber, 2010), occurred along
intertidal zone to depth of 20 m (Higo et al., 1999).
with a C. sikamea / C. gigas hybrid which was detected
Comments: “Crassostrea nipponica” (e.g., Habe (1977);
from mitochondrial and nuclear DNA markers; there
Higo et al. (1999); Lee J.-S. and Min (2002); Min D.-K. et
were no significant differences in shell morphology
al. (2004)) is a misspelling. The close relationship of C.
between the two species.
nippona and C. hongkongensis has been demonstrated
The type material of Ostrea chemnitzii var. elongata
based on mitochondrial genome comparison (Yu and Li,
Grabau et King, 1928 (Yellow Sea) included specimens
2012).
of both C. gigas and of Talonostrea talonata Li et Qi,
1994; however, designation of the lectotype settles its
status as a synonym of C. gigas (Coan et al., 2015). A
Subfamily Saccostreinae Salvi et Mariottini, 2016
syntype of O. laperousii is figured by Lutaenko (2016,
Genus Saccostrea Dollfus et Dautzenberg, 1920
p. 128, figs. 14A, B).
60. Saccostrea echinata (Quoy et Gaimard, 1835)
Salvi et al. (2014) has shown that two groups of
Crassostrea, American and Asian, should be designed
Pl. 14, Figs. E-H
Synonyms: Ostrea mytiloides Lamarck, 1819 (non Hermann,
- 188 -
Korean J. Malacol. 35(2): 149-238 2019
1781 nec Gmelin, 1791); ?Ostrea spathulata Lamarck, 1919;
height to 90-123 mm, wide commissural shelf, and the
Ostrea spinosa Deshayes, 1836; Ostrea arakanensis G.B.
presence of violet squamae or lamellae on the outer
Sowerby II, 1871; Ostrea nigromarginata G.B. Sowerby II,
right valve (Sekino and Yamashita, 2016). This
1871; Saxostrea gradiva Iredale, 1939.
nominal species has long been neglected; then Huber
(2010) regarded it as valid, so, we now accept S.
Local distribution: Soessogak; other records: Lee J.J.
spathulata as a possible synonym of S. echinata. A
and Jwa Y.W. (1988; as Saxostrea); Lee J.-J. et al.
synonym
(1989; as Saxostrea); Lee J.J. and Hyun J.M (1991; as
barclayana G.B. Sowerby II, 1871 (type locality:
Saxostrea) (Seogwipo); Lee J.J. and Hyun J.M. (1992)
Mauritius)
(eastern coast; Udo); Lee J.J. and Kim J.C. (1993)
anatomy of S. echinata was described in detail by
(Biyangdo); Je J.-G. et al. (1994a) (Munseom); Lee J.J.
Amaral and Simone (2016).
of
S.
is,
spathulata
(Huber,
2010).
Shell
in
turn,
Ostrea
morphology
and
et al. (1995) (Jagwido); Lee J.-J. et al. (1997)
(Chujado); Choi K.-S. et al. (2000; as Crassostrea); Lee
61. Saccostrea kegaki Torigoe et Inaba, 1981
Text-fig. 6D; Pl. 14, Figs. A-D
J.J. et al. (2001) (Hwasoon); Je J.-G. et al. (2002; as
Crassostrea); Min D.-K. et al. (2004) (Munseom,
Hwasoon).
Local distribution: Iho, Samyang, Munseom, Hwasoon,
General distribution: Korea, Okinawa, Amami Islands,
Hamo, Gwakji, Pyeoson, south-west of Shinyang,
East China Sea to Indonesia, Vietnam, South China
Soessogak, Daejeong; other records: Yang H.-S. and
Sea and Indo-West Pacific to eastern Africa (Inaba et
Choi
al.,
2004;
Huber,
2010;
Lee
J.-S.,
2016);
K.-S.
(2011);
Cha
J.-H.
et
al.
(2013)
(north-eastern coast of Jeju); Lee H.-J. et al. (2013)
tropical-subtropical.
(Gimnyung Harbor); Lee J.U. et al. (2017).
Habitat: Marine-estuarine; attached to rocks, mangrove
General distribution:
roots, and trees, in the intertidal zone to 20 m
Honshu and southwards to Amami Islands), East
(Okutani, 2000; Huber, 2010).
China Sea (Higo et al., 1999; Min D.-K. et al., 2004;
Comments: The taxonomy of Saccostrea species is
Huber, 2010); subtropical.
extremely difficult because ot their shell plasticity
Habitat: Attached to rocks and other hard substrate in
(Lam and Morton, 2004; 2006). Lam and Morton
the intertidal zone (Okutani, 2000; Huber, 2010).
(2004)
Korea,
Japan
(Mutsu
Bay,
of
Comments: This is a usually strongly spinose, small,
Saccostrea cucullata (Born, 1778) and, based on
and rather fragile species morphologically close to S.
mitochondiral DNA sequence analysis, showed that S.
echinata (Huber, 2010) but well-developed hyote spines
cucullata is a superspecies with Saccostrea glomerata
allow relatively easy separation of S. kegaki from other
regarded
S.
echinata
as
a
synonym
(Gould, 1850) as a component species and S. echinata
lineages of Saccostrea (Sekino and Yamashita, 2016).
as a morpho- or ecotype. Huber (2010) regarded S.
Moreover, S. kegaki is genetically distinct (Lam and
cucullata, S. glomerata and S. echinata as distinct.
Morton, 2006; Sekino and Yamashita, 2016). Compared
Sekino
Japanese
to S. echinata which also occurs in mangrove areas or
and
Yamashita
(2016)
divided
Saccostrea species into seven mitochondrial lineages,
mud flats, S. kegaki has only been found intertidally
however, for most of the lineages, it was difficult to
attached to stones on rocky shores (Huber, 2010).
identify
thereby
Okutani (2000) states that juvenile individuals of S.
preventing a decisive link from being established
echinata may resemble S. kegaki; however, the shell of
between the observed lineages and morphology-based
S. echinata becomes much larger and elongate-oval, as
species names. It seems that S. echinata sensu Inaba
opposed to the generally subcircular shape of S.
et al. (2004) included lineages A, B or H (with hyote
kegaki. Torigoe and Inaba (1981) provided additional
spines) and another oyster, Saccostrea spathulata
distinguishing characters between S. kegaki and S.
(Lamarck, 1919) (lineage J) because of its large size,
echinata; the holotype and two paratypes are figured
a
lineage-specific
shell
feature,
- 189 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
in the same paper (l.c., pl. 1, figs. 1-7; pl. 2, figs. 1-6).
Family Gryphaeidae Vialov, 1936
62. Saccostrea scyphophilla (Peron et Lesueur, 1807)
Genus Hyotissa Stenzel, 1971
Synonyms: Ostrea mordax Gould 1850; Ostrea mordax var.
63. Hyotissa hyotis (L., 1758)
Subfamily Pycnodonteinae Stenzel, 1959
cornucopiaeformis Saville-Kent, 1893 (nom. nov. pro Ostrea
Pl. 13, Figs. E, F
cornucopia Saville-Kent, 1891 non Gmelin, 1791); Ostrea
Synonyms: Ostrea radiata Lamarck, 1819 (non Gmelin,
forskäli var. sueli Jousseaume in Lamy, 1925; Saxostrea
1791).
amasa Iredale, 1939.
Local
Local distribution: Taepyongri (Qi and Choe, 2000).
Korea
(Jeju),
Japan
distribution:
Jeju-shi,
Seongsan,
Seogwipo,
Munseom, Bumseom, Yerae, Hwasoon, Keumneung,
(Kii
Soesoggak,
Jongdal-ri,
Peninsula and southwards, Okinawa), East China Sea,
Jungmun;
other records:
Taiwan, China to Philippines, Malaysia, Thailand and
(Munseom, Supseom); Ko J.-C. et al. (2011) (Gwakji,
Indo-West Pacific (to eastern Africa) and Hawaii
Daepo, Aewol); Cho I-Y. et al. (2014) (Bumseom);
(introduced)
Noseworthy et al. (2016) (Keumneung).
General
distribution:
(Higo
et
al.,
1999;
Huber,
2010);
Daepyeong,
Je
South
J.-G.
Yeongmeori,
et
Korea,
(2002)
al.
Japan
(Kii
tropical-subtropical.
General
Habitat: Often grgarious on intertidal rocks and
Peninsula and southwards), East China Sea, South
stones, cemented by left valve (Huber, 2010).
China
Comments: This species has been long known as
Philippines, Australia to Indo-West Pacific and eastern
Saccostrea mordax (Gould, 1850) but it appears that it
Africa; Clipperton Island; introduced to Florida (Higo
was
Ostrea
et al., 1999; Min D.-K. et al., 2004; Qi, 2004; Kaiser,
scyphophilla Peron et Lesueur, 1807 (Huber, 2010).
2007; Mikkelsen and Bieler, 2008; Huber, 2010);
Lam and Morton (2006; as S. mordax) suggested that
tropical-subtropical.
this species occurs only on exposed, wholly marine rocky
Habitat: Coral and rocky reefs, often attached to rocks
shores whereas the other Saccostrea lineages occupy a
and corals, occasionally free living, at depths from 2-55
wide range of habitats from brackish mangroves to
m (Qi, 2004; Kaiser, 2007; Huber, 2010).
somewhat less exposed marine shores.
Comments: Harry (1985) considered this genus as
This species is missing in the book on Korean mollusks
monotypic, with H. hyotis extending from Madagascar
by D.-K. Min et al. (2004) and a catalogue of J.-S. Lee
to the Tuamotu Islands and occurring also in the
earlier
described
from
Australia
as
distribution:
Sea,
Taiwan,
Hainan
Island,
Vietnam,
and Min (2002), however, it is mentioned as “Saccostrea
tropical eastern Pacific. Huber (2010) recognized
cucullata (Born, 1778)” by Lee J.-S. (2015; 2016).
additionally Hyotissa fisheri (Dall, 1914) (Panamic
According to Huber (2010), S. scyphophilla is internally
area: Mexico to Panama, Ecuado,r and Colombia) and
glossier than S. cucullata, often with a raised, deep
Hyotissa sinensis (Gmelin, 1791) (Indo-Pacific: from
black scar and glossy black margins, solid, generally
Andaman Sea, South China Sea, Thailand, and Borneo
with the typical strong radially crumbled sculpture;
to Polynesia). Coan and Valentich-Scott (2012) regard
un-eroded S. scyphophilla individuals are distinct from
the former as a synonym of H. hyotis. Huber (2010)
other
described H. sinensis as thick and solid, but not, or only
species
of
Saccostrea
in
terms
of
the
regularly-spaced grooves radiating from the umbo to the
weakly,
ventral margin of the right valve, its triangular shell
blackish-purple
shape, and finely, regularly, “m-shaped” plicated valve
compressed, the muscle scar not elevated, typically deep
margin (Lam and Morton, 2009). Moreover, S. cucullata
purple, occasionally yellowish brown. The easiest way to
is limited to both eastern and western Africa (Huber,
disginguish between the two Indo-Pacific species are the
2010), and there is no evidence for Pacific distribution
muscle scars in adults which are yellowish-brown and
of this species (Sekino and Yamashita, 2016).
markedly elevated in H. hyotis and generally smaller,
- 190 -
plicate,
with
borders,
almost
smooth,
generally
flatter,
usually
more
Korean J. Malacol. 35(2): 149-238 2019
dark purple-black and flat in H. sinensis.
Ostrea lentiginosa G.B. Sowerby II, 1871; Ostrea cerata G.B.
Genus Parahyotissa Harry, 1985
1871; Ostrea hanleyana G.B. Sowerby II, 1871; Ostrea
Sowerby II, 1871; Ostrea inaequivalvis G.B. Sowerby II,
Subgenus Parahyotissa Harry, 1985
thaanumi Dall, Bartsch et Rehder, 1938; Ostrea procles
64. Parahyotissa (Parahyotissa) inermis (G.B. Sowerby
Iredale, 1939.
II, 1871)
Pl. 12, Figs. E, F; Pl. 13, Fig. J
Local distribution: Bumseom (Qi and Choe, 2000; as
Synonyms: Ostrea imbricata Lamarck, 1819 (non Gmelin,
Parahyotissa inaequivalvis).
1791); Ostrea nobilis G.B. Sowerby II, 1871; Ostrea sellaformis
General distribution: Korea (?Busan, Ullung-do and
Saville-Kent, 1891 (non Conrad, 1832); Ostrea quirites Iredale,
Jeju-do), Japan (Amami Islands and southwards), East
1939 (nom. nov. pro Ostrea sellaformis Saville-Kent, 1891 non
China Sea, China, Vietnam, Philippines to Indo-West
Conrad, 1832).
Pacific and eastern coast of Africa, Australia, Hawaii
(Higo et al., 1999; Min D.-K. et al., 2004; Xu and
Local distribution: Seongsan, Munseom, Hwasoon,
Zhang, 2008; Huber, 2010); tropical-subtropical.
Sagyei, Yerae; other records: Kim H.S. and Rho B.J.
Habitat: Left valve cemented to rubble, stones, corals,
(1971; as Pretostrea imbricata) (Seogwipo); Je J.-G. et
on rocks, coral reefs, under rocks and coral blocks,
al. (1994a; as Hyotissa imbricata) (Munseom); Je J.-G.
intertidal to 20 m (Huber, 2010).
et
al.
(2002;
as
Hyotissa
hyotis
imbricata
and
Pretostrea imbricata) (Bumseom, Supseom, Munseom).
Subgenus Pliohyotissa Harry, 1985
General distribution: Southern Korea and Japan (Boso
66. Parahyotissa (Pliohyotissa) sp.
Peninsula
and
southwards),
South
China
Pl. 13, Figs. G, H
Sea,
Vietnam, Philippines to Indo-Pacific and eastern
Supseom,
Jagwido,
Munseom,
Africa, Australia; Hawaii (introduced), Easter Island
Local
(Higo et al., 1999; Inaba et al., 2004; Min D.-K. et al.,
Bumseom, Tongilli (Daejeong?), Marado (Qi and Choe
2004;
(2000; as Parahyotissa chemnitzi); Beomseom (Je J.-G.
Huber,
2010;
Raines
and
Huber,
2012);
distribution:
tropical-subtropical.
et al., 2002; as Pretostrea chemnitzii); Seongsan,
Habitat: Cemented by left valve to rocks and corals,
Munseom,
rocky bottoms, from the intertidal zone to depth of 30
Parahyotissa chemnitzii).
m (Higo et al., 1999; Huber, 2010).
General distribution: Korea and Japan (Boso Peninsula
Yerae
(Min
D.-K.
et
al.,
2004;
as
Comments: Hota, Chiba Prefecture, Honshu, Japan was
and southwards) to northern Borneo (Higo et al., 1999;
designated as the type locality for this species (Huber,
Huber, 2010); tropical-subtropical.
2010). P. inermis is the most commonly found gryphaeid
Habitat: Upper subtidal zone.
in the Indo-Pacific, and extremely variable in colors
Comments:
(white, yellow, brown, grey, golden, silvery, red, purple,
present in Borneo, South China Sea to Sagami Bay,
and in all shades in between) and, in shape, usually
Honshu,
rather flat and ovate, but is quite easily differentiated
chemnitzii auctt. (non Hanley, 1846). This species has
by its spongy shell structure, usually best seen on the
a more elongate muscle scar than P. inermis but is
inside borders in fresh specimens, and the round,
similarly colored as the eastern Pacific Parahyotissa
whitish muscle scar (Huber, 2010). The holotype of P.
quercinus (G.B. Sowerby II, 1871) (syntype figured by
inermis is figured by Higo et al. (2001, p. 155, fig. B389).
Coan and Valentich-Scott (2012, pl. 82)), and was cited
A
Parahyotissa
Japan,
was
often
(Pliohyotissa)
named
species,
Parahyotissa
for Japan and China by Inaba et al. (2004); Ostrea
Subgenus Numismoida Harry, 1985
chemnitzii Hanley, 1846 is a synonym of Dendostrea
65. Parahyotissa (Numismoida) numisma (Lamarck, 1819)
rosacea (Deshayes, 1836). Coan and Valentich-Scott
Synonyms: Ostrea violacea Deshayes in Maillard, 1863;
(2012) synonymized both Parahyotissa and Pliohyotissa
- 191 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
with Hyotissa. Records of “Pretostrea rosacea” (in
vitrea Gmelin, 1791; Pinna nuttallii Conrad, 1837; Pinna
Gryphaeidae) and “Dendostrea rosacea” (in Ostreidae)
elongata Reeve, 1858 (non Röding, 1798); Streptopinna
in Jeju Island (Je J.-G. et al., 1994a; Je J.-G. et al.,
saccata inusitata Iredale, 1927.
2002) are perhaps this species.
Local distribution: Jungmun, “all around” (Min D.-K. et
al., 2004).
Superfamily Pinnoidea Leach, 1819
General distribution: Japan (Boso Peninsula and
Family Pinnidae Leach, 1819
southwards, southern Japan/East Sea) and Korea to
Genus Pinna L., 1758
East
Subgenus Quantulopinna Iredale, 1939
Philippines, and Indo-Pacific (to eastern and southern
67. Pinna (Quantulopinna) muricata L., 1758
Africa),
China
and
Hawaii,
South
Australia,
China
seas,
Micronesia,
Indonesia,
Polynesia,
Marquesas, Isla del Coco, Costa Rica; Clipperton
Pl. 18, Figs. A-D
Synonymss: Pinna cancellata Mawe, 1823; Pinna semicostata
Island (pan-Pacific) (Wang, 1964; Higo et al., 1999;
Conrad, 1837; Pinna zebuensis Reeve, 1858; Pinna aequilatera
Min D.-K. et al., 2004; Xu and Zhang, 2008; Schultz
Martens in Möbius, 1880 (non Weinkauff, 1867); Pinna
and Huber, 2013); tropical-subtropical.
hawaiensis Dall, Bartsch et Rehder, 1938; Quantulopinna
Habitat: Byssally attached to coral blocks or within
delsa Iredale, 1939; Quantulopinna delsa howensis Iredale,
branches of living corals, on gravel, under or among
1939.
rocks, subtidal from 1-25 m (Schultz and Huber, 2013).
Comments: S. saccata is the only species currently
Local distribution: Udo (Sanhosa), Yerae, Yeongmeori,
included in the genus Streptopinna.
Hwasoon.
General distribution: Southern Korea and southern
Genus Atrina Gray, 1842
Japan (Pacific coast of southern Honshu, Shikoku,
Subgenus Servatrina Iredale, 1939
Kyushu
69. Atrina (Servatrina) japonica (Reeve, 1858)
and
Ogasawara
southwards,
Islands)
Izu
to
Islands,
Vietnam,
northern
Philippines,
Indonesia, and Indo-Pacific (to eastern and southern
Local distribution: Jeju (Choi K.-S. et al., 2000; as
Africa,
Atrina pectinata).
Red
Sea
and
Persian
Gulf),
Australia,
Easter
General distribution: Korea, Japan, East China Sea,
Island (Wang, 1964; Higo et al., 1999; Min D.-K. et al.,
Yellow Sea (Huber, 2010; Schultz and Huber, 2013);
Melanesia,
Micronesia,
Polynesia,
Hawaii,
2004; Huber, 2010, 2015; Schultz and Huber, 2013);
subtropical.
tropical-subtropical.
Habitat: Deeply buried in fine and sandy mud,
Habitat: Sandy mud among gravel or rocks, sandflats,
interidal to 30 m (Schultz and Huber, 2013).
lagoons, from the intertidal zone to depth of 60 m
Comments: “Atrina (Servatrina) pectinata (L., 1767)”
(Schultz and Huber, 2013).
often
Comments: According to Huber (2010), no marked
(Okutani, 2000; Min D.-K. et al., 2004; Qi, 2004; Zhang
differences were detected between eastern African,
et al., 2016) is instead Atrina (Servatrina) japonica
Indian,
this
(Reeve, 1858) (Schultz and Huber, 2013). A. pectinata
species is very widely distributed and one of the most
occurs in the Indian Ocean, from Seychelles and
common Indo-Pacific pinnids.
Arabia, and India to Indonesia. It is tan to brownish,
Australian,
and
Hawaiian
material;
recorded
from
Japan,
Korea,
and
China
rather fragile, narrower and weakly ribbed, compared
Genus Streptopinna Martens in Möbius, 1880
to the broader, more inflated, almost smooth A.
68. Streptopinna saccata L., 1758
japonica. The latter species grows much larger; it is
olive green, sometimes greenish-black (Huber, 2010;
Pl. 18, Fig. E
Synonyms: Pinna picta Forsskål in Niebuhr, 1775; Pinna
Schultz and Huber, 2013). Another species, Atrina
- 192 -
Korean J. Malacol. 35(2): 149-238 2019
(Servatrina) chinensis (Deshayes in Cuvier, 1841),
al. (2013)) for P. heteroptera. According to Huber
known from China and Japan, is in general smaller,
(2010), P. avicular is generally smaller, more colorful,
with stronger radial sculpture, occassionally spiny,
iridescent,
rather trigonal shape,, more compressed, and has a
heteroptera has a brownish-red, duller, generally larger
deeper habitat (to 200 m) (Schultz and Huber, 2013).
shell, more quadrate as adults, often with strongly
It seems that these two species were recognized both
twisted wings. We follow this interpretation. Kilburn
morphologically (as non-scaly and scaly forms) and
(1975)
genetically in Japan as two forms of “A. pectinata”
described from Java, may be conspecific with P.
(Yokogawa, 1996). This species was not listed by
brevialata
and
strongly
mentioned
but
that
the
oblique
P.
latter
as
adults;
originally
tortirostris,
appears
P.
to
be
less
Noseworthy et al. (2007) although it was previously
inaequivalve, with a somewhat less twisted anterior
found in Jeju and illustrated by Choi K.-S. et al.
wing and deep chestnut coloration, although these
(2000).
characters are quite variable. According to Huber
(2010),
P.
tortirostris
appears
biogeographically
restricted to Indonesian and Philippine waters, and is
Superfamily Pterioidea Gray, 1847
uncommon. No specimens close to P. tortirostris were
Family Pteriidae Gray, 1847
collected in Japan, eastern Africa, or the Red Sea.
Subfamily Pteriinae Gray, 1847
Genus Pteria Scopoli, 1777
71. Pteria gregata (Reeve, 1857)
70. Pteria heteroptera (Lamarck, 1819)
Synonyms: Avicula libella Reeve, 1857; Avicula lovéni
Dunker, 1879; Pteria sibogae Prashad, 1932; Pteria howensis
Pl. 17, Figs. E, F
Synonyms: Avicula rufa Dunker, 1849; Avicula inquinata
Reeve,
1857;
Avicula
castanea
Reeve,
1857;
Lamprell et Healy, 1997; ?Pteria bulliformis Wang, 2001.
Avicula
aquatilis Reeve, 1857; Avicula brevialata Dunker, 1872;
Local distribution: Seogwipo, Munseom (Min D.-K. et
Austropteria levitata Iredale, 1939.
al., 2004).
General distribution: Southern Korea and Japan (Boso
Local distribution: Munseom; other records: Cho I.-Y.
Peninsula and southwards including Japan/East Sea)
et al. (2014; as Pteria brevialata) (Beomseom); Ko J.-C.
to East China Sea, Taiwan, Indonesia, Vietnam,
et al. (2011; as Pteria brevialata) (Daepo).
Philippines, South China Sea, and the Indo-West
General distribution: Southern Korea and Japan (Boso
Pacific, Australia, Polynesia, Melanesia (Higo et al.,
Peninsula and southwards) to East China Sea, Taiwan,
1999; Min D.-K. et al., 2004; Lee J.-S., 2016; Huber,
Philippines, Thailand, Indonesia, and Indo-West Pacific
2010); tropical-subtropical.
(to eastern and southern Africa and Red Sea),
Habitat: Reef slopes, byssally attached to gorgonians,
Australia, northern New Zealand (Higo et al., 1999; as
from depth of 2-144 m (Huber, 2010).
P. brevialata; Min D.-K. et al., 2004; as P. brevialata;
Comments: This species was commonly known as
Huber, 2010); tropical-subtropical.
Pteria loveni (Dunker, 1879) (e.g., Higo et al. [1999];
Habitat: Byssally attached to gorgonians, from depth
Okutani [2000]; Min D.-K. et al. [2004]; Qi [2004]; Xu
of 9-180 m (Huber, 2010).
and Zhang [2008]) until Huber (2010) demonstrate
Comments: This species belongs to the complicated
that it is a synonym of P. gregata (type locality:
complex “Pteria avicular (Holten, 1802)-Pteria tortirostris
Samoa). Characteristic of this species is a fine, dense
(Dunker, 1849)-Pteria heteroptera (Lamarck, 1819)”
commarginal
(Huber, 2010). Asian authors have long used name
periostracal lamellae (Huber, 2010).
microsculpture
overlaid
Pteria brevialata (Dunker, 1872) (e.g., Wang (1978);
Habe (1977); Higo et al. (1999); Okutani (2000); Qi
72. Pteria avicular (Holten, 1802)
(2004); Dharma (2005); Xu and Zhang (2008); Yang et
- 193 -
Pl. 17, Figs. G, H
with
fine
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Synonyms: Avicula chinensis Leach, 1814; Avicula crocea
Local distribution: Hwasoon (our data; Min D.-K. et
Lamarck, 1819; ?Avicula semisagitta var. b Lamarck, 1819;
al., 2004).
?Avicula maura Reeve, 1857; Avicula iridescens Reeve,
General distribution: Korea, Japan (Suruga Bay and
1857; Avicula producta Reeve, 1857; Avicula trochilus
southwards) to East China Sea, Taiwan, Indonesia,
Reeve, 1857; Avicula coturnix Dunker, 1879; Austropteria
Malaysia, Philippines and Indo-Pacific (to eastern
antelata Iredale, 1939.
Africa and Red Sea), Melanesia, Micronesia, Australia
(Wang, 1978; Higo et al., 1999; as P. nigra; Huber,
Local distribution: Munseom, Hwasoon, “all around”
2010; as both P. albina and P. nigra; Lee J.-S., 2016;
(Min D.-K. et al., 2004).
as both P. albina and P. nigra); tropical-subtropical.
General distribution: Southern Korea and Japan (Kii
Habitat: Coral reefs, sand, gravel, byssally attached to
Peninsula and southwards) to East China Sea, Taiwan,
rocks, among stones or gorgonians, intertidal to 35 m
Vietnam, Indonesia, Philippines, and Indo-West Pacific
(Huber, 2010).
(to Red Sea, eastern Africa and Madagascar), Australia
Comments: According to Tëmkin (2010), P. albina and
(Wang, 1978; Higo et al., 1999; Min D.-K. et al., 2004;
Pinctada nigra (Gould, 1850), generally distinguished
as P. coturnix; Huber, 2010); tropical-subtropical.
by shell color, represent two color morphs of the same
Habitat: Byssally attached to alcyonarians, gorgonians,
species. However, Huber (2010) regards P. albina as
or shell debris from depth of 1-20 m (Huber, 2010).
restricted to western Australia only and he stresses
Comments: Huber (2010) regards Avicula maura
that various “P. nigra” were doubtfully identified; no
Reeve, 1857 as a valid species, however, this was long
specimens from the type locality have been genetically
known as Pteria coturnix (Dunker, 1879 (type locality;
analyzed, and this synonymization is not substantiated
Japan) (e.g., Higo et al. (1999); Okutani (2000); Min
(Huber, 2015). Proviionally following Tëmkin [l.c.], the
D.-K et al. (2004); Qi (2004)). P. coturnix appears to be
record of P. nigra from Jeju Island (Noseworthy et al.,
a slightly more prosocline, and possibly juvenile, form
2007) should belong to this species.
of P. avicular (I. Tëmkin, e-mail, March 26, 2015).
Syntypes of A. coturnix are figured by Tëmkin et al.
74. Pinctada imbricata Roding, 1798
Pl. 16, Figs. A-C; Pl. 17, Figs. A, B
(2009, figs. 84, 85). In sense of Huber (2010), P.
avicular is variable in shape and color, but often
Synonyms: Perlamater vulgaris Schumacher, 1817; Avicula
radially striped and with a glossy, golden hue. This is
squamulosa Lamarck, 1819; Margarita crocata Swainson
a comparatively small and fragile species.
1831; Avicula nebulosa Conrad, 1837; Avicula pallida Conrad,
1837;
Philippi,
1849;
?Avicula
pica
Avicula lacunata Reeve, 1857; Avicula aerata Reeve, 1857;
Pl. 16, Figs. F-I
Synonyms: Avicula (Meleagrina) nigra Gould, 1850; Avicula
Dunker,
laticauda
Gould, 1850; Avicula (Meleagrina) citrina Dunker, 1852;
73. Pinctada albina (Lamarck, 1819)
turdus
?Avicula
Philippi, 1849; Avicula fucata Gould, 1850; Avicula lurida
Genus Pinctada Röding, 1798
Philippi,
1851;
1852;
Avicula
Avicula
(Meleagrina)
anomioides
Reeve,
atropurpurea
1857;
Avicula
Avicula perviridis Reeve, 1857; Avicula (Meleagrina) concinna
Dunker, 1872; Avicula (Meleagrina) grisea Dunker, 1872;
Avicula
(Meleagrina)
prasina
Dunker,
1872;
Avicula
fimbriata Reeve, 1857; Avicula imbricata Reeve, 1857; Avicula
(Meleagrina)
irradians Reeve, 1857; Avicula placunoides Reeve, 1857; Avicula
martensii Dunker, 1880 (nom. nov. pro Avicula (Meleagrina)
radula Reeve, 1857; Avicula sugillata Reeve, 1857; Avicula
japonica Dunker, 1879 non Dunker, 1852).
varia
Dunker,
1872;
Avicula
(Meleagrina)
flexuosa Reeve, 1857 (non Orbigny, 1849); Avicula (Meleagrina)
concinna Dunker, 1872; Avicula (Meleagrina) scheepmakeri
Local distribution: Hwasoon, Biyangdo, Hyeopjae, “all
Dunker, 1872; Avicula (Meleagrina) tristis Dunker, 1872; Avicula
around” (Min D.-K. et al., 2004); other records: Je J.-G.
(Meleagrina)
Pteria (Margaritifera)
et al. (1994a; as Pinctada fucata martensii) (Munseom);
carchariarum Jameson, 1901; Pinctada perrutila Iredale, 1939.
Je J.-G. et al. (2002; as Pinctada fucata martensii)
reentsii
Dunker,
1872;
- 194 -
Korean J. Malacol. 35(2): 149-238 2019
(Munseom, Beomseom, Supseom); Cho I.-Y. et al. (2014;
holotype of Avicula fucata Gould, 1850 is figured at:
as Pinctada fucata) (Beomseom).
http://collections.mnh.si.edu/search/iz/?qn=Avicula+fucata.
General distribution: Nearly “cosmopolitan” (globally
distributed but limited to tropical and subtropical
75. Pinctada margaritifera (L., 1758)
Text-fig. 6F; Pl. 16, Figs. D, E
zones); Atlantic and Mediterranean; Korea and Japan
to Indo-Pacific (Wada and Tëmkin, 2008; Huber, 2010);
Synonyms: Margaritiphora communis Megerle von Mühlfeld,
tropical-subtropical.
1811; Margarita sinensis Leach, 1814; Avicula hystrix Reeve,
corals,
1857; ?Avicula chamoides Reeve, 1857; Meleagrina nigromarginata
occasionally estuarine attached to mangrove roots; in
Saville-Kent, 1893; Pteria (Margaritifera) margaritifera typica
muddy areas attached to seagrass or shells, from the
Jameson, 1901; Pteria (Margaritifera) margaritifera erythraeensis
intertidal zone to depth of 60 m (Huber, 2010).
Jameson, 1901; Pteria (Margaritifera) margaritifera persica
Comments: This species belongs to the complicated
Jameson, 1901; Pteria (Margaritifera) margaritifera zanzibarensis
species complex “Pinctada fucata (Gould, 1850)-P.
Jameson, 1901.
Habitat:
Byssally
attached
to
rocks
or
martensii (Dunker, 1880)-P. radiata (Leach, 1814)-P.
imbricata Röding, 1798” (Akoya pearl oyster). Its
Local distribution: Beomseom (Oh et al., 2017).
taxonomic status long remained unsettled due to
General distribution: Widely distributed Indo-Pacific
extensive variation in shell characters within and
species extending from Red Sea, eastern Africa,
among
Persian
populations,
distribution,
extremely
transport
geographical
hybridization
Gulf
to
Australia,
Indonesia,
Melanesia,
Micronesia, Polynesia, Malaysia, Thailand, Philippines,
humans (Wada and Tëmkin, 2008; Tëmkin, 2010;
South China and East China seas, Taiwan, Okinawa,
Cunha et al., 2011). Traditionally, three distinct
and Japan (Wang, 1978; Higo et al., 1999; Huber,
species
2010); tropical-subtropical.
recognized
and
wide
by
were
and
corresponding
to
three
biogeographic regions: P. imbricata in the western
Habitat: Subtidal, at depth of 1-66 m, coral reefs,
Atlantic region, P. radiata in the eastern Indian Ocean
byssally attached to coral slabs or rocks, live and dead
and the Red Sea regions, and P. fucata in the
corals, also on rocky and gravelly bottoms, occur either
Indo-Pacific region. The Japanese populations were
singly or in large clusters (Wada and Tëmkin, 2008;
regarded as a distinct species, P. martensii, or a
Huber, 2010).
subspecies, P. fucata martensii (Wada and Tëmkin,
Comments: The intraspecific variation of this species
2008). A recent molecular study suggests that P.
have raised a possibility that it might, in fact,
imbricata, P. fucata, and P. radiata are distinct
represent
genealogical units and it is desirable to provisionally
Polynesian and Hawaiian populations have sometimes
recognize them at a subspecies level under the senior
been referred to as separate species or subspecies:
synonym, P. imbricata (as Pinctada imbricata imbricata,
Pinctada cumingii (Reeve, 1857) and Pinctada galtsoffi
P. imbricata fucata, and P. imbricata radiata) (Tëmkin,
Bartsch, 1931, respectively (Wada and Tëmkin, 2008).
2010). However, Cunha et al. (2011), based on molecular
Huber (2010) considered them both as valid species.
more
than
a
single
species.
French
studies, regard P. radiata and P. imbricata as distinct
The COI gene of P. margaritifera collected from Jeju
species. Until this mess is resolved, we prefer to use the
was grouped to P. margaritifera, and the species was
oldest name, P. imbricata, sharing Huber’s (2010) opinion
most closely related to the species collected from
but we exclude synonyms based on specimens from the
Chuuk
Atlantic; P. radiata is likely based on Carribean material
Polynesia with 99.4% similarity; 28S rDNA sequence
and this binomen should be avoided when we consider
of
Asian species (I. Tëmkin, e-mail, February 25, 2015). A
margaritifera which were most similar to the pearl
syntype of Avicula (Meleagrina) martensii Dunker, 1880
oysters collected from Micronesia, Japan, and French
is figured by Tëmkin et al. (2009, figs. 45, 46). The
Polynesia with 99.8% similarities (Oh et al., 2017).
- 195 -
Lagoon,
specimens
Micronesia,
from Jeju is
and
also
Takaroa
Atoll,
grouped to
P.
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Genus Pterelectroma Iredale, 1939
Melanesia, Polynesia, Philippines, Thailand, South
76. Pterelectroma physoides (Lamarck, 1819)
China Sea, Taiwan and Amami Islands, Hawaii,
Synonyms: Avicula zebra Reeve, 1857; Pteria tomlini
Mexico, Panama, Galapagos Islands (Huber, 2010);
Prashad, 1932.
tropical.
Habitat: Intertidal and subtidal, to depth of 70 m,
Local distribution: Seogwipo, Munseom (Min D.-K. et
byssally attached to or under rocks, stones, other
al., 2004).
shells, and in coral crevices and calcareous algae
General distribution: Southern Korea and Japan (Boso
(Huber, 2010).
Peninsula and southwards) to East China Sea, Taiwan,
Comments: Jeju specimens exhibit varying degrees of
Vietnam, Philippines, Indonesia, and Indo-West Pacific
concentric
(to Red Sea, eastern and southern Africa), Australia,
straighter and lighter in color, usually whitish or
lamellae,
being
smaller,
somewhat
Melanesia (Wang, 1978; Higo et al., 1999; Min D.-K. et
brownish, with irregular concentric lamellae on the
al., 2004; Huber, 2010); tropical-subtropical.
exterior nacreous area, and blister-like laminations on
Habitat: Coral reefs, byssally attached to whip corals
the
at depth of 3-150 m (Huber, 2010).
morphological form of the species is close to “Malleus
Comments: This species was previously associated with
maculosus Reeve, 1958”, which is one of the synonyms
Vulsellidae but a molecular study showed it is related
of M. regula (Noseworthy et al., 2016b).
calcareous
extension.
It
appears
that
this
to Pteriidae (Tsubaki et al., 2011; Huber, 2015).
Possible syntypes of A. zebra are figured by figured by
Subfamily Isognomoninae Woodring, 1925
Higo et al. (2001, p. 153, fig. B298).
Genus Isogonomon Lightfoot, 1786
78. Isognomon isognomum (L., 1758)
Pl. 19, Figs. A, B, E-J
Family Malleidae Lamarck, 1818
Subfamily Malleinae Lamarck, 1818
Synonyms: Ostrea isogonum L., 1767; Isognoma lignea Lightfoot,
Genus Malleus Lamarck, 1799
1786; Isogonum norma Röding, 1798; Isogonum gnomon Röding,
77. Malleus regula (Forsskål in Niebuhr, 1775)
1798; Perna isogona Bosc, 1801; Isognomon brevirostre Link,
1807; Perna tranquebarensis Leach, 1815; Perna femoralis
Pl. 17, Figs. C, D
Synonyms: Ostrea valsella Gmelin, 1791; Himantopoda
Lamarck, 1819; Perna avicularis Lamarck, 1819; Perna canina
truncata Schumacher, 1817; Malleus vulsellatus Lamarck,
Lamarck, 1819; Perna fimbriata Reeve, 1858; Perna patibulum
1819; Malleus decurtatus Lamarck, 1819; Vulsella nuttalli
Reeve, 1858; Perna rudis Reeve, 1858; Perna vulsella Reeve,
Conrad, 1837; Malleus maculosus Reeve, 1858; Malleus
1858 (non Lamarck, 1819); Perna attenuata Reeve, 1858; Perna
aquatilis Reeve, 1858; Malleus vesiculatus Reeve, 1858;
vespertilio Reeve, 1858; Perna novohollandiae Clessin, 1890;
Malleus rufipunctatus Reeve, 1858; Malleus solitarius Reeve,
Perna aquila Clessin, 1890; Perna rollei Clessin, 1890; Perna
1858; Malleus panamensis Mörch, 1861; Malleus obvolutus
flava Clessin, 1890; Perna planulata Clessin, 1890; Perna
Folin in Folin et Périer, 1867; Fundella lioyi Gregorio, 1884;
isognomon var. obliqua Martens, 1894; Perna isognomon var.
Parimalleus cursator Iredale, 1931; Pinna aenigmatica
brachyptera Martens, 1894.
Turton, 1932; Pinna saccata Turton, 1932; Pinna saccata
similis Turton, 1932; Parimalleus rex Iredale, 1939.
Local
distribution:
Munseom,
Seongsan,
Seogundo,
Jungmun,
Bomok,
Seogwipo,
Yerae,
Hwasoon,
Local distribution: Gwakji, Gwideok (Noseworthy et al.,
Yongmeori,
2016b).
Keumneung; other records: ?Je J.-G. et al. (1994a; as
General
distribution:
Tropical
pan-Pacific
species
Sagyeri,
Songaksan,
Soesoggak,
Isognomon legumen) (Munseom); ?Je J.-G. et al. (2002;
known from eastern Mediterranean, Red Sea, Persian
as
Gulf,
Jigwido); Cho I.-Y. et al. (2014) (as Isognomon
southern and eastern Africa to
Australia,
- 196 -
Isognomon
legumem)
(Munseom,
Beomseom,
Korean J. Malacol. 35(2): 149-238 2019
ephippium) (Beomseom, Munseom); Noseworthy et al.
Superfamily Pectinoidea Rafinesque, 1815
(2016; as Isognomon ephippium) (Keumneung).
General
South
distribution:
Korea
and
Order PECTINIDA Gray, 1854
Japan
Family Pectinidae Rafinesque, 1815
(southern Kyushu and southwards, Ogasawara Islands)
Subfamily Pectininae Rafinesque, 1815
to East China and South China seas, Malaysia,
Genus Pecten Müller, 1776
Indonesia, Thailand, Philippines, and Indo-West Pacific
80. Pecten albicans (Schröter, 1802)
Pl. 20, Figs. F-H
(to eastern Africa), Melanesia, Polynesia, Australia
(Wang, 1980; Higo et al., 1999; Xu and Zhang, 2008;
Synonyms: Pecten laqueatus G.B. Sowerby II, 1842; Pecten
Huber, 2010); tropical-subtropical.
antonii Philippi, 1844; Pecten naganumanus Yokoyama,
Habitat: Byssally attached to rocks or shells from the
1920.
intertidal zone to depth of 10 m (Huber, 2010).
variability,
Local distribution: Sehwa, Udo (Hagosudong), Udo
especially in shell shape, and can be subdivided into
(Sanhosa), Jongdal-ri, Seongsan, Seopjikoji, Beophwan,
several forms (e.g., a form with broad trigonal wing
Gangjeong,
and a wingless form with short and narrow hinge line
Yeongmeori; other records: Kim H.S. and Rho B.J.
(Huber, 2010)). However, no genetic data available for
(1971) (Seogwipo); Je J.-G. et al. (1994a) (Munseom);
separation into several species. Records of “Isognomon
Choi K.-S. et al. (2000); Je J.-G. et al. (2002); Ko J.-C.
ephippium (L., 1758)” from Jeju Island (Noseworthy et
et al. (2012) (southern coast of Jeju).
Comments:
2007)
This
species
belongs
to
shows
this
high
species.
Whether
Jungmun,
Yerae,
Hwasoon,
Gwakji,
I.
General distribution: Korea, Japan (southern Hokkaido
ephippium exists in Korea (Min D.-K. et al., 2004) is
to Kyushu), Yellow and East China seas, Taiwan (Min
questionable.
D.-K. et al., 2004; Xu and Zhang, 2008; Huber, 2010;);
al.,
subtropical.
79. Isognomon nucleus (Lamarck, 1819)
Habitat: Subidal, at depth of 10-150 m, sandy and
Pl. 19, Figs. C, D
sandy mud bottoms (Huber, 2010).
Synonyms: Perna rupella Dufo, 1840; Perna planorbis Dufo,
Comments:
1840; Perna dentifera Krauss, 1848; Perna nana Gould
naganumanus are figured by Oyama (1973, pl. 33, figs.
1850; Perna pectinata Reeve, 1858; Perna quadrangularis
8, 10).
Reeve,
1858;
Perna
lobata
Reeve,
1858;
Lectotype
and
paralectotype
of
P.
Isognomon
acutirostris Otuka, 1936; Parviperna francisensis Cotton et
81. Pecten excavatus Anton, 1838
Godfrey, 1938; Parviperna perexigua Iredale, 1939.
Pl. 19, Fig. K; Pl. 20, Fig. E
Seongsan,
Udo
(Hagosudong),
Local distribution: Yeongmeori.
Local
General distribution: Korea, Japan (Honshu (Kii
Shinyang, Jungmun, Hwasoon, Yongmeori, Gwakji;
Peninsula and southwards)), Ogasawara Islands) to
other records:
South China Sea, Philippines, Vietnam, Thailand and
(Seogwipo); Je J.-G. et al. (2002); Ko J.-C. et al. (2012;
Indo-Pacific (to eastern and southern Africa and Red
as Pecten sinensis puncticulatus) (southern coast of
Sea),
Jeju).
Australia, Polynesia,
Melanesia, Marquesas,
distribution:
Kim
H.S.
and
Rho
B.J.
(1971)
Easter Island (Higo et al., 1999; Xu and Zhang, 2008;
General distribution: Korea and Japan to East China
Huber, 2010); tropical-subtropical.
Sea, Okinawa, China (Min et al., 2004; Xu and Zhang,
Habitat: Byssally attached to rock crevices, under
2008; Huber, 2010); subtropical.
stones and among corals, intertidal to 25 m (Huber,
Habitat: Subtidal, at depth of 100-50 m, sandy bottom
2010).
(Huber, 2010).
Comments: First record for Korea and Jeju Island.
Comments: In regional literature, there was much
confusion concerning P. excavatus and allied Pecten
- 197 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
puncticulatus Dunker, 1877 (Japan and Korea) and
Caledonia, Melanesia) and Y. japonicum belong to a
Pecten sinensis G.B. Sowerby II, 1842 (South China
separate genus based on molecular and morphological
Sea-Okinawa) (see Huber (2010)). P. excavatus exhibits
data. Ylistrum can be separated from Amusium based
strong radial ribbing, also in the interstices of the
on size, number of internal ribs and coloration:
right valve, not found in the closely-related species P.
Ylistrum species are typically larger, reaching 120 mm
puncticulatus and P. sinensis. Also, it has a stronger
in height as compared to 100 mm in Amusium;
radially-grooved structure, flatter squarish ribs on the
Ylistrum has a significantly greater number of internal
left valve, and is much more excavated than the other
ribs on both valves, with 30-45 and 36-51 ribs on left
two species, with a larger umbo, and is only known in
and right valves, respectively, whereas Amusium bears
a dark reddish-white color with a brown border inside
between 19-27 internal ribs on the left valve and 20-30
(Huber, 2010). The similar Australian species, Pecten
ribs on the right (Mynhardt et al., 2014). Huber (2015)
dijkstrai Duncan et Wilson, 2012 is smaller, longer,
additionally listed Amusium taiwanicum Habe, 1992
and less robust than typical Japanese specimens of P.
(nom. nov. pro Amusium japonicum formosum Habe,
excavatus; the radial ribs are weaker (lower) and less
1964 non Melvill et Standen,1907) with a synonym
angular than in P. excavates, and the left valve of P.
Amusium japonicum forma taiwanense Dijkstra, 1988
dijkstrai may be more deeply concave in some
(unavailable,
specimens; the color is paler than in P. excavatus
relationship to Ylistrum will be revealed by molecular
invalid
infrasubspecific
name).
Its
methods although Mynhardt et al. (2014) regarded this
(Duncan and Wilson, 2012).
name as a synonym of Y. japonicum. Jeju populations
Genus Ylistrum Mynhardt et alejandrino
of Y. japonicum are too small to be commercially
in Mynhardt, Alejandrino, Puslednik, Corrales et Serb, 2014
viable.
82. Ylistrum japonicum (Gmelin, 1791)
Pl. 20, Figs. A-D
Subfamily Decatopectininae Waller, 1986
Local
distribution:
Beophwan,
Wimi,
Gangjeong,
Seogwipo,
Jungmun,
Munseom,
Hwasoon;
other
Genus Decatopecten G.B. Sowerby II, 1839
83. Decatopecten plica (L., 1758)
Pl. 22, Figs. D, E
records: Kim H.S. and Rho B.J. (1971; as Amusium
japonicum) (Seogwipo); Je J.-G. et al. (1994a; as
Synonyms: Pallium striatum Schumacher 1817; ?Pecten
Amusium japonicum) (Munseom); Choi K.-S. (1996; as
quadriradiatus Lamarck, 1819; Pecten plicata Swainson,
Amusium japonicum japonicum) (Seogwipo); Choi K.-S.
1840; Pecten subplicatus G.B. Sowerby II, 1842; Pecten
et al. (2000; as Amusium japonicum japonicum); Je
velutinus G.B. Sowerby II, 1842.
J.-G. et al. (2002; as Amusium japonicum); Ko J.-C. et
al. (2012; as Amusium japonicum) (southern coast of
Local distribution: Jungmun, Seongsan, Yongmeori,
Jeju).
Gwakji;
General distribution: Korea, Japan, East China Sea
Decatopecten striatus) (Seogwipo).
other
records:
Choi
K.-S.
(1996;
as
(Higo et al., 1999; Min D.-K. et al., 2004; Xu and
General distribution: Widely distributed Indo-Pacific
Zhang, 2008; Huber, 2010;); subtropical.
species-from Red Sea, Yemen, Oman and Pakistan to
Habitat: Subtidal, at depth of 10-100 m, fine sand
the Philippines, Indonesia, Vietnam, Thailand, Taiwan,
(Huber, 2010); in Jeju, this species was trawled on
China, Japan and Korea (Min D.-K. et al., 2004;
sand flats at a depth of 20-30 m (Choi K.-S., 1996).
Huber, 2010); tropical-subtropical.
Comments: Mynhardt et al. (2014) demonstrated that
Habitat: Intertidal to subtidal zone, at depth of 50 m,
Amusium Röding, 1798 is monotypic with Amusium
mud, muddy sand, or sandy bottoms (Huber, 2010); in
pleuronectes
(L.,
1758)
whereas
Ylistrum
balloti
(Bernardi, 1861) (Indonesia to Australia and New
Jeju, this species was trawled on sand flats, at depth
of 20-30 m (Choi K.-S., 1996).
- 198 -
Korean J. Malacol. 35(2): 149-238 2019
Comments: Japanese (e.g., Habe, 1977; Higo et al.,
J.-M. (1997; as Chlamys farreri farreri) (Chujado); Choi
1999) and Korean workers (Min D.-K. et al., 2004)
K.-S. et al. (2000; as Chlamys farreri farreri); Je J.-G.
regard Decatopecten striatus (Schumacher, 1817) as a
et al. (2002) (Munseom, Beomseom); Ko J.-C. et al.
separate species. This is a narrow, heavy, thick, and
(2011;
strongly convex form (Huber, 2010); however, Dijkstra
Aewol, Woljeong, Haengwon).
including
(1999; 2013) and Huber (2010) regard it as a synonym
General distribution: Korea, Japan, Taiwan, China,
of D. plica. Decatopecten strangei (Reeve, 1852) from
Yellow Sea, and East/Japan Sea, to Peter the Great
Chlamys
nipponensis)
(Sincheon,
Australia is closely allied to D. plica and it differs
Bay (Higo et al., 1999; Min D.-K. et al., 2004; Xu and
from the latter by its flattened left valve, rather
Zhang, 2008; Huber, 2010; Lutaenko and Noseworthy,
constant number of ribs (5 against 3-7 ribs in D. plica),
2012); subtropical.
higher and broader ribs, and less bright coloration. D.
Habitat: Subtidal, at depth of 1-60 m, juveniles may
strangei may be only a southern geographical variant of
live intertidally, byssally attached to rocks and gravel,
D. plica, because their differences in shape, size and
able to swim (Scarlato, 1981; Lutaenko, 2005b; Huber,
sculpture are not substantial (Dijkstra and Beu, 2018).
2010).
A similar species, D. amiculum (Philippi, 1851) (Red
Comments:
Separation
of
Ch.
into
farreri
two
Sea and eastern Africa to Japan) Higo et al. (1999);
subspecies in Japanese literature is not supported by
Huber (2010)), with more radial plicae (8-11), and living
other workers (Kafanov and Lutaenko, 1998; Wang,
in the same habitat as D. plica, may also belong to the
2002; Huber, 2010). This species does not inhabit the
range of variation of D. plica (Dijkstra, 2013); Huber
Kurile Islands (Scarlato, 1981), as stated by Huber
(2010) regards the former as a separate species. A
(2010) but it is known, however, from northern
record
Hokkaido (Sea of Okhotsk) (Kitamura et al., 2002). A
of
Gloripallium
pallium
(Linnaeus
1758)
(Decatopectininae) recorded from Seogwipo (Kim H.S.
syntype of Ch. farreri is figured by Higo et al. (2001, p.
and Rho B.J., 1971) and Munseom (Je J.-G. et al.,
156, fig. B453).
1994a; Je J.-G. et al., 2002) and previously not known
from Korea (Lee J.-S. and Min D.-K., 2002; Min D.-K.
85. Chlamys (Scaeochlamys) squamea Dijkstra et
et al., 2004) may be a misidentification of D. plica;
Maestrati, 2009
Pl. 21, Figs. A-J
however, G. pallium was recently listed from Korea by
Lee J.-S. (2016).
Local
Jeju-shi,
distribution:
Seongsan,
Bomok,
Subfamily Chlamydinae Teppner, 1922
Hyungjaeseom,
Udo
Jungmun,
Biyangdo,
(Hagosudong),
Hwasoon,
Gwakji,
Sagyei,
Bukchon-ri,
Genus Chlamys Röding, 1798
Gimnyeong, Wolpyeong, Jongdal-ri, Shinyang, Seogwipo,
Subgenus Scaeochlamys Iredale, 1929
Munseom,
84. Chlamys (Scaeochlamys) farreri (Jones et Preston, 1904)
Biyangdo, Daejong; other records: Noseworthy et al.
Yerae,
Yongmeori,
Hamo,
Keumneung,
(2002; as Chlamys irregularis) (Seongsan); ?Suzuki
Pl. 22, Figs. A, B
Synonyms: Pecten laetus Gould, 1861 (non Gould, 1850);
(2010; as Chlamys irregularis) (Jungmun, Hwasoon,
Chlamys farreri nipponensis Kuroda, 1932; Chlamys farreri
Gwakji); Ko J.-C. et al. (2011; as Chlamys irregularis)
akazara Kuroda, 1932; Chlamys nantiensis Bernard, Cai et
(Sincheon, Aewol, Woljeong, Haengwon); Noseworthy et
Morton, 1993 (nom. nov. pro Pecten laetus Gould, 1861).
al. (2016; as Laevichlamys cuneata) (Keumneung).
General distribution:
Korea,
Japan,
East
China,
Local distribution: Seongsan, Beophwan, Gangjeong,
Taiwan, and the Philippines to Australia (Dijkstra and
Jungmun, Gwakji, Yeongmeori, Daejong; other records:
Beu, 2018).
Lee J.J. (1990) (northern coast); Kim H.S. et al. (1992);
Habitat: Subtidal, at depth of 5-50 m, shell is often
Je J.-G. et al. (1994a) (Munseom); Lee J.-J. and Hyun
deformed as a result of its coral-dwelling niche
- 199 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
(Djikstra, 2013; Dijkstra and Beu, 2018; Dijkstra and
Maestrati, 2009).
Comments:
This
(northern coast); Je J.-G. et al. (1994a; as Chlamys)
(Munseom); Je J.-G. et al. (2002; as Chlamys); Min
species has
been
mistaken
for
D.-K et al. (2004).
Laevichlamys lemniscata (Reeve, 1853) known only
General distribution: Korea and Japan, South China
from the south-western Indian Ocean: S. squamea has
Sea,
coarser radial macrosculpture on the right valve with
tropical-subtropical.
fewer radial riblets (30-36) than L. lemniscata (up to 50)
Habitat: Subtidal-bathyal, at depth of 50-600 m, coarse
and lacks antimarginal microsculpture on the right valve,
sand and stones (Huber, 2010).
Ch. squamea has persistant shagreen microsculpture on
Comments: The holotype of Ch. empressae is figured
the left valve lacking in L. lemniscata, as in all other
by Higo et al. (2001, p. 156, fig. B444).
East
China
Sea,
Taiwan
(Huber,
2010);
species of Laevichlamys (Djikstra, 2013; Dijkstra and
Beu,
2018).
Chlamys
(Scaeochlamys)
squamata
(Gmelin, 1791), known from the western Pacific, differs
Family Spondylidae J. Gray, 1826
from Ch. squamea in having fewer prominent radial
Subfamily Spondylinae J. Gray, 1826
costae (5-6) on the left valve (compared with 8-9 ribs
Genus Spondylus L., 1758
in Ch. squamea), weaker early radial sculpture on the
87. Spondylus varius G.B. Sowerby I, 1827
right valve, and slightly more radial costae (6-7) on
Pl. 23, Fig. I
the anterior auricle of the right valve (compared with
Synonyms: Spondylus delessertii Chenu, 1844; Spondylus
4 in Ch. squamea) and shagreen microsculpture
striatospinosus
throughout on both valves (Ch. squamea only on the
Sowerby II, 1847 (?lapsus calami).
Chenu,
1844;
Spondylus
varians
G.B.
left valve early in ontogeny) (Djikstra, 2013; Dijkstra
and Beu, 2018). Specimens from Korea depicted as
Local distribution: Seongsan, Jungmun, Hwasoon,
“Chlamys (Scaeochlamys) irregularis (G.B. Sowerby II,
Yongmeori, Hamo; other records: Ko J.-C. et al. (2011)
1842)” (Min D.-K. et al., 2004, fig. 1306; Lutaenko and
(Sincheon).
Noseworthy, 2012, pl. 20, figs. E, F), “Chlamys
General distribution: Widely distributed Indo-Pacific
(Coralichlamys) empressae Kuroda et Habe, 1951 (Min
species extending from eastern Africa to Australia,
D.-K. et al., 2004, fig. 1308), “Chlamys (Azumapecten)
Melanesia, Polynesia, Micronesia, Indonesia, Philippines,
larvata (Reeve, 1853)” (Min D.-K. et al., 2004, fig.
South China Sea, Taiwan, China, Amami Islands and
1313) and “Chlamys (Azumapecten) lemniscata (Reeve,
Korea (Xu and Zhang, 2008; Huber, 2010); tropical-
1853)” (Min D.-K. et al., 2004, fig. 1314; Lutaenko and
subtropical.
Noseworthy, 2012, pl. 21, figs. A, B) are all instead
Habitat: Subtidal, to depth of 2-60 m, attached to dead
Ch. squamea (H. Dijikstra, pers. comm., February 3
corals, rocks, debris, or living free (Huber, 2010).
and 9, 2015). Huber (2015) synonymizes Ch. squamea
Comments: This is the largest species of the family
with Ch. squamata indicating that these species
reaching 400 mm in height (Huber, 2010). In Korea, is
intergrade in number of dorsal ribs, in ribs on auricles,
is also known from Ullung Island in the East/Japan
and in reticulate microsculpture. Separation of these
Sea (Choe B.L. et al., 1994).
species is much in need of molecular study.
88. Spondylus squamosus Schreibers, 1793
Genus Veprichlamys Iredale, 1929
Pl. 23, Figs. A, B, E, F
86. Veprichlamys jousseaumei (Bavay, 1904)
Synonyms: Spondylus spathuliferus Lamarck, 1819; Spondylus
Synonyms: Chlamys (Chlamys) empressae Kuroda et Habe
barbatus Reeve, 1856; Spondylus mus Reeve, 1856; Spondylus
in Kuroda, Oyama et Habe, 1971.
cruentus Lischke, 1868; Spondylus japonicus Kuroda, 1932
(nom. nov. pro Spondylus sinensis G.B. Sowerby II, 1847 non
Local distribution:
Lee
J.J.
(1990;
as
Chlamys)
Schreibers, 1793).
- 200 -
Korean J. Malacol. 35(2): 149-238 2019
Local distribution: Iho, Haengwon, Handong, Hado,
Udo
(Hagosudong),
Seongsan,
Sagye-ri,
Shinyang,
Munseom,
Udo
(Sanhosa),
Jongdal-ri,
Pyoseon,
Bomok,
Seogwipo,
Jungmun,
Yerae,
Hwasoon,
Yerae,
Hwasoon,
Daejong;
Yongmeori,
other records:
Je
Hamo,
J.-G.
Keumneung,
et
al.
(2002)
(Supseom); Noseworthy et al. (2002) (Seongsan); Cho
I.-Y. et al. (2014) (Beomseom); Noseworthy et al. (2016)
Yongmeori, Hamo, Keumneung, Biyangdo, Hyeopjae,
(Keumneung).
Gwakji, Daepyong, Weoljeong; other records: Kim H.S.
General distribution: Widely distributed Indo-Pacific
et al. (1992; as Spondylus barbatus); Je J.-G. et al.
species
(eastern
Africa
to
Australia,
Melanesia,
(1994a; as Spondylus barbatus) (Munseom); Choi K.-S.
Micronesia, Indonesia, Philippines Thailand, South
et al. (2000; as Spondylus barbatus); Je J.-G. et al.
China Sea, East China Sea, Korea and Japan (Min
(2002) (Munseom, Beomseom); Noseworthy et al. (2002;
D.-K. et al., 2004; Xu and Zhang, 2008; Huber, 2010);
as Spondylus barbatus cruentus) (Seongsan); Suzuki
tropical-subtropical.
(2010; as Spondylus cruentus) (Hamdeok, Sehwa,
Habitat: Intertidal and subtidal, to depth of 38 m, in
Ojo-ri, Shinyang, Hyopjae, Kwakji); Noseworthy et al.
coral reefs, protected areas, attached to corals (Huber,
(2016) (Keumneung).
2010).
General distribution: Korea and Japan, Australia,
Comments: This species was figured as Spondylus
Melanesia, Philippines, Thailand, South China Sea,
longitudinalis Lamarck, 1819 in Korean literature
East China Sea, Taiwan (Min D.-K. et al., 2004; Xu
(Min D.-K. et al., 2004, fig. 1329). Huber (2010)
and Zhang, 2008; Huber, 2010); tropical-subtropical.
regards this species as dubious. However, a probable
Habitat: Subtidal, to depth of 1-100 m, attached to
holotype of S. longitudinalis was found and illustrated
corals and rocky bottom (Huber, 2010).
(Finet and Lamprell, 2008, pl. 20, figs. 91-93), and
Comments: This species was recorded and illustrated
these authors consider this species as a synonym of
for Korea as Spondylus balbatus (sic!) cruentus
Spondylus americanus Hermann, 1781. The latter is a
Lischke, 1868 (Min D.-K. et al., 2004, fig. 1328) and
Caribbean species known from North Carolina and
Spondylus cruentus Lischke, 1868 (Lutaenko and
Florida to Brazil (Huber, 2010; Mikkelsen and Bieler,
Noseworthy, 2012, pl. 23, figs. A, B). Lamprell and
2008) whereas Spondylus americanus Schreibers, 1793
Willan (2000) designated and figured lectotypes of S.
(non Hermann, 1791) is a synonym of S. butleri. S.
barbatus and S. squamosus and found them conspecific.
butleri
is closely
related to
Spondylus
versicolor
S. squamosus is similar is shape to Spondylus sinensis
Schreibers, 1793 (Japan to Australia), differing in
Schreibers, 1793 but the former lacks the well-defined
having thicker, sometimes spathulate spines (Lamprell,
and uniform radial rows of ribs and uniform spines,
2006).
radial rows of black and white in the interstices, and
spotting at the
umbonal
area (Lamprell,
2006).
Lectotype of Spondylus spathuliferus Lamarck, 1819,
Superfamily Entolioidea Teppner, 1922
another synonym, was figured and discussed by
Family Propeamussiidae Abbott, 1954
Lamprell and Willan (2000) and Finet and Lamprell
Genus Parvamussium Sacco, 1897
(2008). Two syntypes of S. cruentus were figured by
90. Parvamussium intuscostatum (Yokoyama, 1920)
Higo et al. (2001, p. 158, fig. B536).
Synonyms: Pecten tenuicostatus Yokoyama, 1922.
89. Spondylus butleri Reeve, 1856
Local distribution: Seongsan (Min D.-K. et al., 2004).
Pl. 23, Figs. C, D, G, H
General distribution: Korea, East China Sea, Okinawa,
Synonyms: Spondylus americanus Schreibers, 1793 (non
Japan (Kyushu and Honshu) (Higo et al., 1999; Huber,
Hermann, 1781).
2010).
Habitat: Subtidal-bathyal, at depth of 50-500 m, fine
Local distribution: Seongsan, Munseom, Jungmun,
sand and mud, coarse sand and gravel (Huber, 2010).
- 201 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Comments. The lectotype is figured by Oyama (1973,
red, rose, orange, brownish and cream to white
(Huber, 2010). The true identity of Anomia plana
pl. 33, fig. 1).
Grabau et King, 1928 is somewhat doubtful (Huber,
2010), type material was not found (Coan et al., 2015).
Superfamily Anomioidea Rafinesque, 1815
“A. chinensis” from Korea (Min D.-K. et al., 2004, fig.
Family Anomiidae Rafinesque, 1815
1330; Lutaenko and Noseworthy, 2012, pl. 23, fig. G,
Genus Anomia L., 1758
H) is instead A. cytaeum. Monia umbonata (Gould,
91. Anomia cytaeum Gray, 1850
1861) (now in Isomonia Dautzenberg et H. Fischer,
1897) previously listed for Jeju (Noseworthy et al., 2007;
Pl. 15, Figs. G-I
Synonyms: Anomia placentella Reeve, 1859; Anomia rubella
Cho I.-Y. et al., 2014) is a doubtful record. A syntype of
Philippi in Küster, 1868; Anomia lischkei Dautzenberg et H.
A. cytaeum is figured by Higo et al. (2001, p. 159, fig.
Fischer, 1907; Anomia nipponensis Yokoyama, 1920; Anomia
B553s); lectotype and paralectotypes of A. nipponensis
cuticulum Grabau et King, 1928; Anomia nipponensis var.
are figured by Oyama (1973, pl. 28, figs. 2, 6).
obsoletocostata Grabau et King, 1928; Anomia plana Grabau
et King, 1928.
Superfamily Dimyoidea P. Fischer, 1886
Local
distribution:
Udo
(Hagosudong),
Seongsan,
Family Dimyidae P. Fischer, 1886
Bomok, Seogwipo, Munseom, Seogundo, Jungmun,
Genus Dimya Rouault, 1850
Yongmeori, Sagyei, Keumneung, Biyangdo, Soesoggak,
92. Dimya japonica Habe, 1971
Hyeopjae, Gwakji; other records: Kim H.S. et al., 1992
Synonyms: Dimya lima Habe, 1958 (non Bartsch, 1913).
(as Anomia chinensis); Je J.-G. et al. (1994a; as
Anomia chinensis) (Munseom); Je J.-G. et al. (2002; as
Local distribution: Hwasoon (Min D.-K. et al., 2004).
Anomia chinensis) (Beomseom, Jigwido); Noseworthy
General distribution: Korea and Japan, East China
and Choi K.-S. (2010; as Monia umbonata) (Seogundo);
Sea, (Higo et al., 1999; Min D.-K. et al., 2004; Huber,
Ko J.-C. et al. (2011; as Anomia chinensis) (Sincheon);
2010; Lutaenko and Noseworthy, 2012).
?Cho I.-Y. et al. (2014) (as Monia umbonata) (Munseom);
Habitat: Subtidal-bathyal, at depth of 20-600 m, attached
Noseworthy et al.
to other shells or gravel (Habe, 1971; Huber, 2010).
(2016;
as Isomonia umbonata)
(Keumneung).
Comments: The holotype and paratypes are figured by
General distribution: Korea and Japan, Indo-Pacific
Habe (1971, figs. 9-19).
(widely distributed species known from Melanesia,
New Caledonia, Singapore to Yellow Sea), (Huber,
2010); tropical-subtropical.
Superfamily Plicatuloidea Gray, 1854
Habitat: Intertidal to 30 or 116 m, attached by
Family Plicatulidae Gray, 1854
calcareous byssus to rocks, timber, or shells (Huber,
Genus Plicatula Lamarck, 1801
2010).
93. Plicatula australis Lamarck, 1819
Comments: A. cytaeum was originally described from
Pl. 22, Fig. C
China and this is the earliest name for this common
Synonyms: Plicatula depressa Lamarck, 1819; Plicatula
species, although it was often misidentified as “Anomia
lineata Récluz, 1851; Plicatula multiplicata Deshayes in
chinensis Philippi, 1849” which is also rather widely
Maillard, 1863; Plicatulostrea onca Simone et Amaral, 2008.
distributed in the Indo-Pacific including Philippines,
Thailand, Vietnam, China, and Japan (Huber, 2010).
Local
The former species has a rougher, irregularly wrinkled
Hwasoon, Sagyei, Yeongmeori.
sculpture as compared to A. chinensis, the upper valve
General distribution: Widely distributed Indo-Pacific
is moderately inflated, with a variety of colors-deep
species extending from eastern Africa and India to
- 202 -
distribution:
Shinyang,
Bomok,
Seogwipo,
Korean J. Malacol. 35(2): 149-238 2019
Australia, Micronesia, Guam, Philippines, Indonesia,
General distribution: Korea, Japan (Honshu, Sagami
Vietnam, East China Sea, Okinawa, Korea and Japan
and Toyama bays) (Higo et al., 1999; Min D.-K. et al.,
(Min D.-K. et al., 2004; Xu and Zhang, 2008; Huber,
2004; Huber, 2010); subtropical.
2010); tropical-subtropical.
Habitat: Subtidal, at depth of 50-150 m (Huber, 2010).
Habitat: Intertidal and subtidal, to depth of 150 m,
Comments: “Lima vulgata” aucct. is misspelling. “Lima
coral reefs, flatly attached to rocks, dead corals and
vulgaris Link” of Je J.-G. et al. (1994a) from Munseom
gravel (Huber, 2010).
and “Lima sowerbyi” of Lee J.-J. and Hyun J.-M.
(1997) from Chujado and of Lee J.J. (1991) from
Gapado may be this species; Lima sowerbyi Deshayes
94. Plicatula horrida Dunker, 1882
in Maillard, 1863 is a synonym of Lima vulgaris Link,
Pl. 22, Fig. F
Synonyms: Plicatula irregularis Dunker, 1882 (nom. nov.
1807; the latter taxon is a distinct Indo-Pacific species
pro Plicatula rugosa Dunker, 1877 non Lamarck, 1819).
extending from eastern Africa to Japan (Huber, 2010).
The lectotype and paralectotype are figured by Oyama
Local
distribution:
Seongsan,
Yeongmeori;
other
(1973, pl. 36, figs. 1, 2).
records: Min D.-K. et al. (2004) (Sagyeri).
General distribution: Korea and Japan, East China
97. Lima (Allolima) fujitai Oyama, 1943
Sea, China, (Min D.-K. et al., 2004; Huber, 2010);
subtropical.
Local distribution: Hwasoon (Min D.-K. et al., 2004).
Habitat: Subtidal, at depth of 10-20 m, rubble and
General distribution: Korea, Japan, and South China
corals (Huber, 2010).
Sea, Philippines, East China Sea, Taiwan (Min D.-K.
et al., 2004; Huber, 2010); tropical-subtropical.
Habitat: Subtidal-bathyal, at depth of 40-300 m,
Superfamily Limoidea Rafinesque, 1815
byssally attached to sand and gravel (Huber, 2010).
Family Limidae Rafinesque, 1815
Comments: Some authors (e.g., Higo et al. (1999))
Subfamily Liminae Rafinesque, 1815
synonymized L. fujitai with L. vulgatula. However, L.
Genus Lima Bruguière, 1797
fujitai is smaller and more convex, with more ribs
Subgenus Lima Bruguière, 1797
(28-35) as compared to L. vulgatula; the latter has 22
95. Lima (Lima) zushiensis Yokoyama, 1920
broad, flattish ribs (Huber, 2010).
Pl. 24, Figs. A, B
Genus Ctenoides Mörch, 1853
Local distribution: Seongsan, Yeongmeori, Daejong,
98. Ctenoides annulatus (Lamarck, 1819)
Hwasoon, Daepyeong.
Pl. 24, Figs. E, F
General distribution: Korea, Japan (Kyushu, Shikoku,
Synonyms: Lima brunnea Cooke, 1886; Ctenoides ferescabra
Honshu, Hokkaido), East/Japan Sea (Higo et al., 1999;
Iredale, 1939; Ctenoides minimus Hayami et Kase, 1993.
Min D.-K. et al., 2004; Huber, 2010); subtropical.
Habitat: Subtidal, at depth of 30-200 m, byssally
Local distribution: Iho, Seongsan, Shinyang, Bomok,
attached to rock and stones (Huber, 2010).
Jungmun,
Yerae,
Hwasoon,
Yeongmeori,
Sagyei,
Comments: The lectotype is figured by Oyama (1973,
Gwakji,
pl. 36, fig. 6).
(Jungmun, Hwasoon, Gwakji); Ko J.-C. et al. (2012)
Daejong;
other
records:
Suzuki
(2010)
(southern coast of Jeju).
Subgenus Allolima Stuardo, 1968
General distribution: Widely distributed Indo-Pacific
96. Lima (Allolima) vulgatula Yokoyama, 1922
species extending from Red Sea, eastern Africa, Oman
to
Local distribution: Hwasoon, Gwakji.
Indonesia,
Australia,
Melanesia,
Micronesia,
Polynesia, Philippines, South China Sea, East China
- 203 -
Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Sea, Korea and Japan (Min D.-K. et al., 2004; Huber,
Je J.-G. et al. (1994a) (Munseom); Je J.-G. et al. (2002)
2010); tropical-subtropical.
(Munseom).
Habitat: Subtidal at depth of 5-200 m, on coral reefs,
General distribution: Korea and Japan, South China
byssally attached to corals, in crevices and dead coral
and East China seas (Higo et al., 1999; Min D.-K. et
heads, also among coral rubble and in subtidal caves
al., 2004; Huber, 2010); subtropical.
(Huber, 2010). The holotype of C. minimus is figured
Habitat: Subtidal, at depth of 1-50 m, among pebbles,
by Hayami and Kase (1993, figs. 244-246).
on rock and gravel bottoms (Higo et al., 1999; Huber,
2010).
Comments: In Jeju Island, empty shells are usually
99. Ctenoides lischkei (Lamy, 1930)
found dead on beaches after storms. This species has a
Pl. 24, Fig. G
Synonyms: Lima japonica Dunker, 1877 (non A. Adams,
very fragile shell. A syntype is figured by Higo et al.
1864); Lima dunkeri Smith, 1885 (non Hagenow, 1842;
(2001, p. 154, fig. B367).
nom. nov. pro Lima japonica Dunker, 1877); Ctenoides
corallicola Iredale, 1939.
Subfamily Limatulinae Kasum-Zade, 2003
Local
distribution:
Iho,
Gimnyeong,
Wolpyeong,
Genus Limatula Wood, 1839
Seongsan, Shinyang, Gangjeong, Jungmun, Yerae,
Subgenus Limatula Wood, 1839
Hwasoon, Yongmeori, Sagyei,
101. Limatula (Limatula) vladivostokensis (Scarlato, 1955)
Hamo, Keumneung,
Biyangdo, Gwakji; other records: Noseworthy et al.
(2016) (Keumneung).
Local distribution: Hwasoon (Min D.-K. et al., 2004).
General distribution: Korea and Japan, Indonesia,
General distribution: East/Japan Sea including Korean
Papua New Guinea, Australia, Philippines, South
and Russian coasts, western Sakhalin, Japan, Kurile
China Sea, East China Sea, Okinawa (Higo et al.,
Islands, eastern Kamchatka (Scarlato, 1981; Higo et
1999;
al., 1999; Min D.-K. et al., 2004); subtropical-boreal.
Min
D.-K.
et
al.,
2004;
Huber,
2010);
tropical-subtropical.
Habitat: Subtidal-bathyal, at depth of 75-664 m, in
Habitat: Subtidal, at depth of 2-160 m, attached to
fine sand, muddy sand, rarely mud, sandy gravel
rocks and stones, gravelly bottom (Huber, 2010).
(Scarlato, 1981; Higo et al., 1999).
Comments: The main differences between C. annulatus
and C. lischkei are larger size, with fewer and
102. Limatula (Limatula) nippona Habe, 1960
prickly-scaled ribs in the former, whereas the latter
has more and smoother ribs and it is smaller, C.
Local distribution: Supseom (Je J.-G. et al., 2002);
annulatus is generally much more common (Huber,
Shinyang, Hwasoon (Min D.-K. et al., 2004).
2010). This species is often quite common on beaches
General distribution: Korea and Japan, East China
of Jeju Island as dead, single valves. This may be
Sea, Amami Islands (Higo et al., 1999; Min D.-K. et
“Ctenoides lamyi (Smith)” (Kwon et al., 1993; Je J.-G.
al., 2004; Huber, 2010); subtropical.
et al., 2002; Cho I.-Y. et al., 2014) from Beomseom and
Habitat: Subtidal, at depth of 5-30 or 80 m, fine sand
Sinchon, a non-existent name.
and sandy mud (Higo et al., 1999; Huber, 2010).
Comments: “Limatula nipponica” aucct. is a mispelling.
Genus Limaria Link, 1807
A possible type specimen is figured by Higo et al.
Subgenus Platilimaria Habe, 1977
(2001, p. 154, fig. B375).
100. Limaria (Platilimaria) hirasei (Pilsbry, 1901)
103. Limatula (?Limatula) kurodai Oyama, 1943
Pl. 24, Figs. C, D
Local distribution: Seongsan, Munseom, Beomseom,
Jungmun, Hwasoon, Hyeopjae, Gwakji; other records:
Local distribution: Hwasoon (Min D.-K. et al., 2004).
- 204 -
Korean J. Malacol. 35(2): 149-238 2019
General distribution: Korea and Japan (Kyushu,
study. Dr. Tsukasa Waki, Meguro Parasitological
Shikoku, Honshu, Hokkaido including East/Japan Sea)
Museum, Tokyo, Japan, kindly provided Japanese
(Higo et al., 1999; Huber, 2010); subtropical.
literature references. Thanks go also to Harry Lee,
Habitat: Subtidal-upper bathyal, at depth of 10-300 m,
Florida, U. S. A., for taxonomic assistance, and the
in fine sand (Higo et al., 1999).
late Richard E. (Dick) Petit, South Carolina, U.S.A.,
who helped to unravel some challenging bivalve
Subgenus Limatuletta Fleming, 1978
nomenclature. Many knowledgeable members of the
104. Limatula (Limatuletta) japonica A. Adams, 1864
Conch-L listserve were always ready to answer queries
concerning bivalve research. Thanks go also to Joy
Local distribution: Shinyang (Min D.-K. et al., 2004).
Noseworthy, the second author's wife, who was of
General distribution: Korea, Japan, and Aleutian
great assistance during many field trips. Also, Irina E.
Islands (Coan et al., 2000; Huber, 2010); subtropical-
Volvenko, Zoological Museum, Science and Educational
boreal.
Museum of the Far Eastern Federal University,
Habitat: Subtidal, at depth of 10-300 m, in fine sand
Vladivostok, Russia, did a great deal of work taking all
and sandy mud (Huber, 2010). Two syntypes are
the shell photographs and assembling them in plates;
figured by Higo et al. (2001, p. 154, fig. B383).
she also provided much help while working with the
museum collection in her possesion. Dr. Henk Dijkstra,
Subgenus Stabilima Iredale, 1939
Naturalis
105. Limatula (Stabilima) bullata (Born, 1778)
Netherlands, and Dr. Ilya Tëmkin, American Museum
Biodiversity
Center,
Leiden,
the
Synonyms: Limatula strangei Oyama, 1943 (non G.B.
of Natural History, N.Y., U.S.A., kindly checked our
Sowerby II, 1872).
identifications of Pectinidae, Malleidae, and Pteriidae,
and provided many comments on their taxonomy. Prof.
Local distribution: Seongsan, Shinyang, Hwasoon,
Byung-Lae
Choe,
formerly,
Sung
Kyung
Kwan
Sagyei.
University, Suwon, Republic of Korea, Dr. Jong-Geel
General distribution: Korea and Japan, Philippines,
Je, formerly, Korea Institute of Ocean Science and
South China Sea, Borneo, East China Sea, Okinawa
Technology, Ansan, Republic of Korea, and Dr. Lee
(Huber, 2010); tropical-subtropical.
Jung-Sang, Gangwon National Iniversity, Chuncheon,
Habitat: Subtidal, at depth of 8-100 m, in coral sand
Republic of Korea, provided literature.
and fine sand (Huber, 2010).
Comments: The corrected type locality for this species
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Acknowledgements
This work would not have possible without the
generous support of many people. We would like to
thank the students at the Shellfish and Aquaculture
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Kang
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Korean J. Malacol. 35(2): 149-238 2019
Plate 1. Fig. A Didimacar tenebrica (Reeve, 1844): Udo, Sanhosa, ZMFU no. 41412/Bv-6111, shell length 14.6 mm; Figs. B, E
Nucula (Nucula) paulula A. Adams, 1856: northern coast of Jeju-do, 33°32′32″ N, 126°49′50″ E, ZMFU no. 41392/Bv-6096,
shell length 4.1 mm (B), 3.3 mm (E); Figs. C, D Striarca symmetrica (Reeve, 1844): Jeju-do, Seongsan, ZMFU no.
41408/Bv-6107, shell length 12.8 mm; Fig. F Acar plicata (Dillwyn, 1817): Jeju-do, Gwakji, ZMFU no. 41410/Bv-6109, shell
length 17.6 mm; Figs. G, F Arca (Arca) avellana Lamarck, 1819: Jeju-do, Yeongmeori, ZMFU no. 41416/Bv-6115, shell
length 23.2 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 2. Figs. A-C Anadara (Scapharca) sp.: Jeju-do, Yeongmeori, ZMFU no. 41404/Bv-6106, shell length 31.9 mm (A);
Hwasoon, ZMFU no. 41409/Bv-6108, shell length 36.8 mm (B); Seongsan, ZMFU 41413/Bv-6112, shell length 30.1 mm (C);
Fig. D Anadara (Scapharca) talmiensis Kalishevich, 1976: Jeju-do, JNU coll., shell length 51.9 mm.
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Korean J. Malacol. 35(2): 149-238 2019
Plate 3. Fig. A Anadara (Scapharca) kagoshimensis (Tokunaga, 1906): Jeju-do, Yeongmeori, ZMFU no. 41403/Bv-6105, shell
length 29.4 mm; Fig. B A. (S.) kagoshimensis (Tokunaga, 1906): Jeju-do, JNU coll., shell length 45.2 mm; Fig. C Anadara
(Tegillarca) granosa (L., 1758): Jeju-do, Gumullae, ZMFU no. 41411/Bv-6110, shell length 32.7 mm; Fig. D A. (T.) granosa
(L., 1758): Jeju-do, JNU coll., shell length 45.3 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 4. Figs. A, B Anadara (Scapharca) broughtonii (Schrenck, 1867): southern coast of Jeju-do, 2 km south-west of Wimi,
ZMFU no. 52558/Bv-8291, shell length 68.3 mm; Figs. C, D Barbatia (Abarbatia) decussata (G.B. Sowerby I, 1833):
Jeju-do, Hwasoon, ZMFU no. 43363/Bv-6599; Figs. E, F, I-L Limopsis (Oblimopa) japonica A. Adams, 1863: Jeju-do, Yerae,
ZMFU no. 40559/Bv-5957, shell length 13.8 (E, F), 15.0 mm (I-L); Fig. G Barbatia (Mesocibota) bistrigata (Dunker, 1866):
Jeju-do, Seongsan, ZMFU no. 52562/Bv-8295; Fig. H Porterius dalli (E.A. Smith, 1885): Jeju-do, Seongsan, ZMFU no.
41414/Bv-6113, shell length 38.3 mm.
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Korean J. Malacol. 35(2): 149-238 2019
Plate 5. Figs. A, B Barbatia (Abarbatia) trapezina (Lamarck, 1819): Jeju-do, ZMFU no. 25365/Bv-4245, shell length 43.7 mm;
Fig. C B. (A.) trapezina (Lamarck, 1819): southern coast of Jeju-do, Daepyeong Village, ZMFU no. 43364/Bv-6600, shell
length 39.0 mm; Fig. D, E B. (A.) trapezina (Lamarck, 1819): Jeju-do, Hwasoon, ZMFU no. 43365/Bv-6601, shell length 36.2
mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 6. Figs. A, B Barbatia (Savignyarca) virescens (Reeve, 1844): Jeju-do, Hwasoon, ZMFU no. 43358/Bv-6594, shell length
31.7 mm; Figs. C, D B. (S.) virescens (Reeve, 1844): Jeju-do, Daepyeong, ZMFU no. 43359/Bv-6595, shell length 21.2 mm;
Figs. E, F B. (S.) virescens (Reeve, 1844): Jeju-do, Seongsan, ZMFU no. 43357/Bv-6593, shell length 34.2 mm.
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Korean J. Malacol. 35(2): 149-238 2019
Plate 7. Figs. A, B Barbatia (Savignyarca) virescens (Reeve, 1844): Jeju-do, Yeongmeori, ZMFU no. 43360/Bv-6596, shell
length 33.1 mm; Figs. C-H Barbatia (Ustularca) stearnsii (Pilsbry, 1895): Jeju-do, Yeongmeori, ZMFU no. 43362/Bv-6598,
shell length 11.5 mm (C), 14.4 mm (D), 16.5 mm (E), 19.6 mm (F, G), 16.1 mm (H); Fig. I Glycymeris (Glycymeris) cf.
albolineata (Lischke, 1872): Jeju-do, JNU coll., shell length 62.9 mm; Fig. J Glycymeris (Glycymeris) cf. rotunda (Dunker,
1882): Jeju-do, JNU coll., shell length 40.9 mm; Fig. K G. (G.) rotunda (Dunker, 1882): Jeju-do, JNU coll., shell length 22.6
mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 8. Figs. A, B Glycymeris (Glycymeris) aspersa (A. Adams et Reeve, 1850): Jeju-do, Jungmun, ZMFU no. 41368/Bv-6100,
shell length 21.0 mm; Fig. C Glycymeris sp.: Jeju-do, Jungmun, ZMFU no. 41399/Bv-6101, shell length 24.9 mm; Fig. D G.
(G.) aspersa (A. Adams et Reeve, 1850): Jeju-do, Hwasoon, ZMFU no. 40577/Bv-5973, shell length 51.0 mm; Fig. E G.
(G.) cf. aspersa (A. Adams et Reeve, 1850): Jeju-do, Jungmun, ZMFU no. 40578/Bv-Bv-5974, shell length 31.0 mm; Figs.
F-H G. (G.) aspersa (A. Adams et Reeve, 1850): Jeju-do, Jungmun, ZMFU no. 40578/Bv-Bv-5974, shell length 25.0 mm (F),
18.0 mm (G), 20.0 mm (H); Figs. I, J Musculus (Musculus) pusio (A. Adams, 1862): Jeju-do, Seongsan, ZMFU no.
41400/Bv-6102, shell length 4.8 mm; Fig. K M. (M.) pusio (A. Adams, 1862): Jeju-do, Gwakji, ZMFU no. 41402/Bv-4104,
shell length 7.8 mm.
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Korean J. Malacol. 35(2): 149-238 2019
Plate 9. Figs. A, B Modiolus (Modiolus) auriculatus (Krauss, 1848): Jeju-do, ZMFU no. 25055/Bv-4121, shell length 28.0 mm (A),
15.1 mm (B); Fig. C Modiolus (Modilous) nipponicus (Oyama, 1950): Jeju-do, Yeongmeori, ZMFU no. 41361/Bv-6088, shell
length 30.1 mm; Figs. D, E Modiolus (Modiolus) kurilensis Bernard, 1983: Jeju-do, Yerae, ZMFU no. 52557/Bv-8290, shell
length 18.6 mm; Fig. F M. (M.) kurilensis Bernard, 1983: Jeju-do, Yeongmeori, ZMFU no. 52556/Bv-8289, shell length 32.0
mm; Figs. G, H M. (M.) nipponicus (Oyama, 1950): Jeju-do, Gwakji, ZMFU no. 52555/Bv-8228, shell length 17.8 mm; Figs.
I, J Xenostrobus atratus (Lischke, 1871): Jeju-do, Ojo-ri, ZMFU no. 52554/Bv-8287, shell length 9.3 mm; Figs. K, L
Lithophaga (Leiosolenus) lischkei (Huber, 2010): Jeju-do, JNU coll., shell length 32.3 mm; Figs. M, N Lithophaga
(Leiosolenus) cf. malaccana (Reeve, 1857): Jeju-do, Jongdal-ri, ZMFU no. 41389/Bv-6093, shell length 18. mm (M), 6.9 mm
(N); Figs. O, P L. (L.) lischkei (Huber, 2010): Jeju-do, Daepyeong, ZMFU no. 52552/Bv-8265, shell length 19. 1 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 10. Figs. A, B, D, E Brachidontes mutabilis (Gould, 1861): Jeju-do, Yeongmeori, ZMFU no. 41396/Bv-6098, shell length
14.5 mm (A, B), 12.5 mm (D, E); Figs. C, H, I Septifer (Mytilisepta) virgatus (Wiegmann, 1837): Jeju-do, Gwakji, ZMFU no.
41236/Bv-6048, shell length 37.7 mm (C), 51.6 mm (H, I); Fig. F, G S. (M.) virgatus (Wiegmann, 1837): Jeju-do, Hwasoon,
ZMFU no. 41237/Bv-6049, shell length 44.0 mm.
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Korean J. Malacol. 35(2): 149-238 2019
Plate 11. Figs. A-E Septifer (Septifer) bilocularis (L., 1758): Jeju-do, Yeongmeori, ZMFU no. 41401/Bv-6103, shell length 30.6
mm (A, B), 23.7 mm (C-E); Figs. F, G Septifer (Mytilisepta) keenae Nomura, 1936: Jeju-do, Yerae, ZMFU no.
41350/Bv-6077, shell length 28.3 mm; Figs. H, I S. (M.) keenae Nomura, 1936: Jeju-do, Seongsan, ZMFU no.
41352/Bv-6079, shell length 19.0 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 12. Figs. A, B Mytilus galloprovincialis Lamarck, 1819: Jeju-do, Hwasoon, ZMFU no. 40900/Bv-6012, shell length 73.4
mm; Figs. C, D M. galloprovincialis Lamarck, 1819: Jeju-do, Yeongmeori, ZMFU no. 41238/Bv-6050, shell length 57.5 mm;
Figs. E, F Parahyotissa (Parahyotissa) cf. inermis (G.B. Sowerby II, 1871): Jeju-do, Munseom, depth 10 m, ZMFU no.
52572/Bv-8305, shell height 98.7 mm; Fig. G Perna viridis (L., 1758): Jeju-do, ZMFU no. 52553/Bv-8286, shell length 104.
mm.
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Plate 13. Figs. A, B Ostrea circumpicta Pilsbry, 1904: Jeju-do, Pyeoson, ZMFU no. 45477/Bv-7295, shell height 64.4 mm (A),
69.9 (B); Figs. C, D Ostrea circumpicta Pilsbry, 1904: Jeju-do, ZMFU no. 45478/Bv-7296, shell height 54.6 mm; Figs. E, F
Hyotissa hyotis (L., 1758): Jeju-do, Daepyeong, ZMFU no. 45772/Bv-7290, shell height 59.5 mm; Figs. G, H Parahyotissa
(Pliohyotissa) sp.: Jeju-do, JNU coll., shell height 45.9 mm; Fig. I Ostrea cf. denselamellosa Lischke, 1869: Jeju-do, Hwasoon,
ZMFU no. 45475/Bv-7293, shell height 42.4 mm; Fig. J Parahyotissa (Parahyotissa) cf. inermis (G.B. Sowerby II, 1871):
Jeju-do, Yerae, ZMFU no. 45480/Bv-7298, shell height 31.5 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 14. Figs. A, B Saccostrea kegaki Torigoe et Inaba, 1981: Jeju-do, JNU coll., shell height 21.7 mm (A), 38.3 mm (B); Figs.
C, D S. kegaki Torigoe et Inaba, 1981: Jeju-do, Shinyang, ZMFU no. 45476/Bv-7294, shell height 28.0 mm (C), 34.0 (D);
Figs. E, F Saccostrea echinata (Quoy et Gaimard, 1835): Jeju-do, Seongsan, ZMFU no. 45479/Bv-7297, shell height 41.5
mm; Figs. G, H S. echinata (Quoy et Gaimard, 1835): Jeju-do, Hwasoon, ZMFU no. 52564/Bv-8297, shell height 23.6 mm
(G), 26. 1 mm (H).
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Plate 15. Figs. A-D Crassostrea (Magallana) gigas (Thunberg, 1793): Jeju-do, Hwasoon, ZMFU no. 45473/Bv-7291, shell height
60.3 mm (A, B), 70.7 mm (C, D); Figs. E, F C. (M.) gigas (Thunberg, 1793): Jeju-do, Hwasoon, ZMFU no. 45474/Bv-7292,
shell height 68.7 mm; Fig. G Anomia cytaeum Gray, 1850: Jeju-do, Jungmun, ZMFU no. 43418/Bv-6652, shell height 18.1
mm; Figs. H, I Anomia cytaeum Gray, 1850: Jeju-do, Yeongmeori, ZMFU no. 43417/Bv-6651, shell height 35.7 (H), 16.7 (I).
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 16. Figs. A-C Pinctada imbricata Roding, 1798: Jeju-do, Hwasoon, ZMFU no. 43548/Bv-6728, shell length 61.8 mm (A),
34.3 (B), 61.8 (C); Figs. D, E Pinctada margaritifera (L., 1758): Jeju, Beomseom, JNU coll., shell length 105. 7 mm; Figs.
F-G Pinctada albina (Lamarck, 1819): Jeju-do, JNU coll., shell length 33.6 mm; Figs. H, I P. albina (Lamarck, 1819): Jeju-do,
Hwasoon, RGN coll., shell length 20.6 mm.
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Plate 17. Figs. A, B Pinctada imbricata Roding, 1798: Jeju-do, Seongsan, JNU coll., shell height 66.9 mm; Figs. C, D Malleus
regula (Forsskål in Niebuhr, 1775): Jeju-do, Gwideok, JNU coll., shell height 21-22 mm, after Noseworthy et al. (2015, fig.
2); Figs. E, F Pteria heteroptera (Lamarck, 1819): Jeju-do, JNU coll., shell length 60.1 mm; Figs. G, H Pteria avicular
(Holten, 1802): Jeju-do, Hwasoon, ZMFU no. 43547/Bv-6727, shell length 69.7 mm (two different valves).
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 18. Figs. A-D Pinna (Quantulopinna) muricata L., 1758: Jeju-do, JNU coll., shell height 91.7 mm (A, B); 44.6 mm (C, D);
Fig. E Streptopinna saccata L., 1758: Jeju-do, Jungmun, ZMFU no. 41397/Bv-6099, shell height 42.9 mm.
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Plate 19. Figs. A, B Isognomon isognomum (L., 1758): Jeju-do, Hwasoon, ZMFU no. 43550/Bv-6730, shell length 18.7 mm;
Figs. C, D Isognomon nucleus (Lamarck, 1819): Jeju-do, Yeongmeori, ZMFU no. 43549/Bv-6729, shell length 12.5 mm;
Figs. E-F I. isognomum (L., 1758): Jeju-do, Yerae, ZMFU no. 43551/Bv-6731, shell height 25.6 mm; Figs. G-J I. cf.
isognomum (L., 1758): Jeju-do, JNU coll., shell height 81.4 mm (G-I), 53.1 mm (J); Fig. K Pecten excavatus Anton, 1838:
Jeju-do, Udo (Hagosudong), ZMFU no. 43468/Bv-6702, shell length 40.7 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 20. Figs. A, B Ylistrum japonicum (Gmelin, 1791): Jeju-do, Hwasoon, ZMFU no. 43419/Bv-6653, shell length 49.3 mm;
Figs. C, D Y. japonicum (Gmelin, 1791): Jeju-do, ZMFU 22616/Bv-3572, shell length 98.6; Fig. E Pecten excavatus Anton,
1838: Jeju-do, JNU coll., shell length 80.7 mm; Fig. F Pecten albicans (Schröter, 1802): Jeju-do, Hwasoon, ZMFU no.
43467/Bv-6701, shell length 34.9 mm; Fig. G P. albicans (Schröter, 1802): Jeju-do, Jungmun, ZMFU no. 43466/Bv-6700,
shell length 29.0 mm; Fig. H P. albicans (Schröter, 1802): Jeju-do, JNU coll., shell length 55.2 mm.
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Plate 21. Fig. A Chlamys (Scaeochlamys) squamea Dijkstra et Maestrati, 2009: Jeju-do, Gwakji, ZMFU no. 43462/Bv-6696, shell
length 16.9 mm; Figs. B-D Ch. (S.) squamea Dijkstra et Maestrati, 2009: Jeju-do, Jungmun, ZMFU no. 43463/Bv-6697, shell
length 21.9 mm (B), 18.4 mm (C), 16.2 mm (D); Figs. E, F, I, J Ch. (S.) squamea Dijkstra et Maestrati, 2009: Jeju-do,
Jungmun, ZMFU no. 43465/Bv-6699, shell length 18.7 mm (E), 14.3 mm (F), 13.9 mm (I), 13.3 mm (J); Figs. G, H Ch. (S.)
squamea Dijkstra et Maestrati, 2009: Jeju-do, Yeongmeori, ZMFU no. 43464/Bv-6698, shell length 10.9 mm (G), 21.0 (H).
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 22. Figs. A, B Chlamys (Scaeochlamys) farreri (Jones et Preston, 1904): Jeju-do, JNU coll., shell height 35.7 mm; Fig. C
Plicatula australis Lamarck, 1819: Jeju-do, Hwasoon, ZMFU no. 41393/Bv-6097, shell height 14.8 mm; Figs. D, E
Decatopecten plica (L., 1758): Jeju-do, JNU coll., shell height 32.9 mm (D), shell length 40. 7 mm (E); Fig. F Plicatula
horrida Dunker, 1882: Jeju-do, Seongsan, ZMFU no. 40920/Bv-6017, shell height 24.0 mm.
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Plate 23. Figs. A, B Spondylus squamosus Schreibers, 1793: Jeju-do, JNU coll., shell length 53.8 mm; Figs. C, D Spondylus
butleri Reeve, 1856: Jeju-do, JNU coll., shell length 64.2 mm; Figs. E, F S. squamosus Schreibers, 1793: Jeju-do,
Hwasoon, ZMFU no. 43553/Bv-6733, shell length 32.6 mm; Figs. G, H S. cf. butleri Reeve, 1856: Jeju-do, Hwasoon, ZMFU
coll., shell length 35.4 mm; Fig. I Spondylus varius G.B. Sowerby I, 1827: Jeju-do, Seongsan, ZMFU no. 52577/Bv-8310,
shell length 27.6 mm.
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Marine Bivalve Mollusks of Jeju Island (Korea). Part 1
Plate 24. Fig. A Lima (Lima) zushiensis Yokoyama, 1920: Jeju-do, Daepyeong, ZMFU no. 51190/Bv-8089, shell height 30.3 mm;
Fig. B L. (L.) zushiensis Yokoyama, 1920: Jeju-do, Yeongmeori, ZMFU no. 51189/Bv-8088, shell height 40.3 mm; Figs. C, D
Limaria (Platilimaria) hirasei (Pilsbry, 1901): Jeju-do, Hwasoon, ZMFU no. 51194/Bv-8093, shell height 35.3 mm; Fig. E
Ctenoides annulatus (Lamarck, 1819): Jeju-do, Yeongmeori, ZMFU no. 51192/Bv-8091, shell height 28.1 mm; Fig. F C.
annulatus (Lamarck, 1819): Jeju-do, Yerae, ZMFU no. 51191/Bv-8090, shell height 25.1 mm; Fig. G Ctenoides cf. lischkei
(Lamy, 1930): Jeju-do, Seongsan, ZMFU no. 51188/Bv-8087, shell height 26.6 mm.
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