Contribution to the Knowledge of the Marine Bivalve Mollusk Fauna of Gangwon Province, Korea

Abstract

Marine bivalve mollusks were obtained from 13 collecting sites, intertidally on beaches and from fishing nets in ports, along the coast of Gangwon Province, Korea. A total of 70 species belonging to 27 families were encountered; all species are illustrated with photographs. Among them, 17 species are recorded for the first time for Gangwon-do, and two species are new records for Korea: Gari chinensis Deshayes, 1855 (Psammobiidae) and Clinocardium likharevi (Kafanov in Scarlato, 1981) (Cardiidae). Taxonomic and distributional comments on 13 species are provided, and the biogeography of the area is discussed. Some warm-water species of bivalves (e.g., Anadara talmiensis (Kalishevich, 1976), Trapezium liratum (Reeve, 1843), and Meretrix lusoria (Röding, 1798)), found in Gangwon-do for the first time, are remnants of the Holocene warming in the Russian sector of the East Sea (Sea of Japan) thus indicating the significance of Korean waters as a transitional zone for past molluscan migrations.

Full text

Original Article
Contribution to the knowledge of the marine bivalve mollusk fauna of
Gangwon Province, Korea
Konstantin A. Lutaenko
a
,
*
, Ronald George Noseworthy
b
a
National Scientific Center of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690041, Russia
b
Faculty of Marine Biomedical Sciences, Jeju National University, Jeju-si 690-756, Republic of Korea
article info
Article history:
Received 25 January 2018
Received in revised form
9 May 2018
Accepted 23 July 2018
Available online 7 August 2018
Keywords:
Biogeography
Bivalves
East Sea (Sea of Japan)
Korea
New records
abstract
Marine bivalve mollusks were obtained from 13 collecting sites, intertidally on beaches and from fishing
nets in ports, along the coast of Gangwon Province, Korea. A total of 70 species belonging to 27 families
were encountered; all species are illustrated with photographs. Among them, 17 species are recorded for
the first time for Gangwon-do, and two species are new records for Korea: Gari chinensis Deshayes, 1855
(Psammobiidae) and Clinocardium likharevi (Kafanov in Scarlato, 1981) (Cardiidae). Taxonomic and
distributional comments on 13 species are provided, and the biogeography of the area is discussed. Some
warm-water species of bivalves [e.g. Anadara talmiensis (Kalishevich, 1976), Trapezium liratum (Reeve,
1843), and Meretrix lusoria (Röding, 1798)], found in Gangwon-do for the first time, are remnants of the
Holocene warming in the Russian sector of the East Sea (Sea of Japan), thus indicating the significance of
Korean waters as a transitional zone for past molluscan migrations.
Ó2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing
Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
Introduction
Marine bivalve mollusks of the Republic of Korea are generally
well studied with publication of several books, atlases, taxonomic
monographs, and comprehensive national and regional checklists
and accounts (Kwon and Lee 1999; Kwon et al 1993, 2001; Lee
1956, 1958, 2013, 2014, 2015, 2016; Lee and Min 2002; Lutaenko
2014; Lutaenko et al 2003; Lutaenko and Noseworthy 2012; Min
et al 2004; Noseworthy et al 2007). However, detailed data on
the regional faunas are still scarce. For that reason, we undertook a
faunal study of bivalve mollusks of the most northern province of
South Korea, on the East Sea (Sea of Japan) coast of Gangwon
Province, or Gangwon-do. Gangwon Province is divided between
North and South Korea; therefore, the southern part was studied.
Previous data on marine molluscan species living in Gangwon-
do were mostly published in the books covering the entire
Korean fauna (Kwon et al 2001; Min et al 2004) in which regional
distribution by provinces was included. We extracted this infor-
mation, taking into account current taxonomy and synonymy of
bivalve mollusks, updated it with our own data, and then published
in English in a catalog of marine bivalves of the continental coast of
the East Sea (Sea of Japan) (Lutaenko and Noseworthy 2012).
Bivalve mollusks, being one of the dominant groups in intertidal
and subtidal communities, have been mentioned as part of various
environmental and ecological studies along the Gangwon-do coast:
Gangneung (Choi et al 2000), Aninjin (Kim et al 1983), Namdae-
chon estuary, Yangyang (Hong et al 2000), and artificial reefs (Kim
et al 2008). Park et al (2011) published a checklist of marine
invertebtrates, including mollusks, of Goseong-gun. Lutaenko et al
(2002) described molluscan beach assemblages in six localities
along the eastern coast of Korea including three (Gangneung,
Jumunjin, and Gallam) in Gangwon-do. A total of 143 species of
marine bivalves based on these combined data are known at pre-
sent from Gangwon-do (Lutaenko and Noseworthy 2012, 2014).
Material and methods
The material for this study was collected in May 2016 on the
coast of Gangwon-do which included several kinds of sampling:
beach collections (beach thanatocoenoses of dead shells), intertidal
sampling, and fish nets near docks in fishing ports. The collecting
was undertaken at 13 localities along the coast (Figure 1): Mangsan,
Gamchu, Anin, Yeomjeon, Sacheon, Yeongeok, Chuam, Maenbang,
Geumjing, and Okgye beaches and Deoksan and Geumjing ports.
Empty shells were washed in fresh water, dried, properly labeled,
*Corresponding author.
E-mail address: lutaenko@mail.ru (K.A. Lutaenko).
Peer review under responsibility of National Science Museum of Korea (NSMK) and
Korea National Arboretum (KNA).
Contents lists available at ScienceDirect
Journal of Asia-Pacific Biodiversity
journal homepage: http://www.elsevier.com/locate/japb
https://doi.org/10.1016/j.japb.2018.07.009
pISSN2287-884X eISSN2287-9544/Ó2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open
access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Journal of Asia-Pacific Biodiversity 12 (2019) 14e44

and then sorted in the laboratory; live specimens were fixed in 70%
alcohol. After identification, selected shells were photographed
using a digital camera, Sony
a
77 (SONY Corp.). In total, 1695 spec-
imens were processed. The collection is stored at Jeju National
University and the Zoological Museum, Science and Educational
Museum of the Far Eastern Federal University (ZMFU). Identifica-
tion was made using large regional literature studies and by
consulting the voucher collections in the aforementioned in-
stitutions. Taxonomic and biogeographic comments are provided
for some species when necessary. The regional setting of the
Gangwon coast, with relation to faunal analysis, was described by
Lutaenko et al (2002).
Results
In total, our study revealed 70 species belonging to 27 families
(Table 1;Figures 2e24). In a previous rapid study in 1997 (Lutaenko
et al 2002), we collected from three localities in Gangwon Province,
Sachondan, Taejin Village, and Gallam Village, with only 24 species
of bivalves obtained; all of those were found in the 2016 study.
Therefore, a more extensive collecting effort (13 sites in 2016 vs. 3in
1997) brought three times more species. A total of 143 species of
marine bivalves is known from all of Gangwon Province (Lutaenko
and Noseworthy 2012, 2014). In our list (Table 1), 17 species are
recorded for the first time for Gangwon-do: Anadara talmiensis
(Kalishevich, 1976), A. broughtonii (Schrenck, 1867), Striarca
symmetrica (Reeve, 1844), Mytilus trossulus Gould, 1850, Modiolus
nipponicus (Oyama, 1950), Spondylus cruentus Lischke, 1868, Clino-
cardium likharevi (Kafanov in Scarlato, 1981), Cardita leana Dunker,
1860, Conchocele scarlatoi Ivanova et Moskaletz, 1984, Kellia
japonica Pilsbry, 1895, Trapezium liratum (Reeve, 1843), Nuttallia
obscurata (Reeve, 1857), Gari chinensis Deshayes, 1855, Macoma
incongrua (Martens, 1865), Meretrix lusoria (Röding, 1798), Irus
ishibashianus Kuroda et Habe, 1952, and Myadoropsis transmontana
(Yokoyama, 1922). Among them, two species are new records for
Korea: G. chinensis and Cl. likharevi. Another five species have been
known in the Korean mollusk literature under different names and
thus are reidentifications: Acila vigilia Schenck, 1936,A. talmiensis,
C. scarlatoi,Potamocorbula amurensis (Schrenck, 1861), and Petricola
habei Huber, 2010. Although the previous names for these species
Figure 1. Map of the area studied along the coast of Gangwon Province, Korea. Names of sampling localities: 1, Mangsan Beach; 2, Gamchu Beach; 3, Anin Beach; 4, Yeomjeon
Beach; 5, Sacheon Beach; 6, Yeongeok Beach; 7, Chuam Beach; 8, Samcheok-si, Gwangjin-gil, Saecheonnyeondo-ro; 9, Maenbang Beach; 10, Deoksan Port; 11, Geumjin Port; 12,
Geumjin Beach; 13, Okgye Beach.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 15

Table 1. List of bivalve mollusk species collected in Gangwon Province.
Species Collecting sites Biogeographic characteristics
12345678910111213
Fam. Nuculidae Gray, 1824
1. Acila insignis (Gould, 1861) þs/b
2. A. vigilia Schenck, 1936 þlb
Fam. Arcidae Lamarck, 1809
3. Arca boucardi Jousseaume, 1894 þþþþ þþ þ þ þ s/b
4. Anadara talmiensis Kalishevich, 1976 þs
5. A. broughtonii (Schrenck, 1867)þs
6. Striarca symmetrica (Reeve, 1844) þþt/s
Fam. Parallelodontidae Dall, 1898
7. Porterius dalli (Smith, 1885) þ þþþþþþ þ þ þ s
Fam. Glycymerididae Dall, 1908
8. Glycymeris yessoensis (G. B. Sowerby III, 1889) þþþþþþþ þ þ þ lb
9. G. albolineata (Lischke, 1872)þþ þ s
10. G. sp. þþ
Fam. Mytilidae Rafinesque, 1815
11. Mytilus galloprovincialis Lamarck, 1819 þþ þþþþþþþ þ þ þ s/b
12. M. trossulus Gould, 1850 þ þþ þþþþ þ wdb þcb
13. M. coruscus Gould, 1861 þþþþþþþ þ s
14. Septifer virgatus (Wiegmann, 1837) þþþþþþþþþ þ þ þ t/s
15. S. keenae Nomura, 1936 þþ þ þ þ þ s
16. Modiolus kurilensis Bernard, 1983 þþþs/b
17. M. nipponicus (Oyama, 1950) þþ þ þ þ t/s
Fam. Ostreidae Rafinesque, 1815
18. Crassostrea gigas (Thunberg, 1793) þ þþþþþþþ þ þ s/b
19. C. nippona (Seki, 1934) þþþþþþþþ s
Fam. Anomiidae Rafinesque, 1815
20. Anomia chinensis Philippi, 1849 þþþþþ þ þþ t/s
Fam. Pectinidae Wilkes, 1810
21. Chlamys farreri (Jones et Preston, 1904) þþþ þþ s
22. C. swiftii (Bernardi, 1858) þþlb
23. Scaeochlamys sp. þþ þ þ þ
24. Mizuhopecten yessoensis (Jay, 1857) þþ þþ þ þ þ lb
Fam. Spondylidae Gray, 1826
25. Spondylus cruentus Lischke, 1868 þþs
Fam. Lucinidae Fleming, 1828
26. Pillucina pisidium (Dunker, 1860) þt/s
Fam. Thyasiridae Dall, 1900
27. Conchocele scarlatoi Ivanova et Moskaletz, 1984 þwdb þcb
Fam. Carditidae Férussac, 1828
28. Cardita leana Dunker, 1860 þþ t/s
Fam. Trapezidae Lamy, 1920
29. Trapezium liratum (Reeve, 1843) þt/s
Fam. Cardiidae Lamarck, 1809
30. Clinocardium likharevi (Kafanov in Scarlato, 1981)þlb
31. Fulvia mutica (Reeve, 1844) þþs
Fam. Lasaeidae Gray, 1842
32. Kellia japonica Pilsbry, 1895 þs/b
Fam. Mactridae Lamarck, 1809
33. Mactra chinensis Philippi, 1846 þ þþþþ þ þ þ þ s/b
34. M. quadrangularis Deshayes in Reeve, 1854 þt/s
35. Spisula sachalinensis (Schrenck, 1861)þþþ þ þ lb
Fam. Tellinidae Blainville, 1814
36. Cadella lubrica (Gould, 1861) þþþs/b
37. Megangulus venulosus (Schrenck, 1861)þþþþ lb
38. M. zyonoensis (Hatai et Nisiyama, 1939) þlb
39. Macoma incongrua (Martens, 1865) þþþ þ s/b
40. M. calcarea (Gmelin, 1791) þb/a
41. M. irus (Hanley, 1844) þþþ s
42. M. sectior (Oyama, 1950) þþ s
43. Angulus sp. þþ
Fam. Psammobiidae Fleming, 1828
44. Nuttallia obscurata (Reeve, 1857) þþþ þ þ þ s
45. Gari chinensis (Deshayes, 1855) þt/s
Fam. Ungulinidae Gray, 1854
46. Felaniella usta (Gould, 1861) þþs/b
47. Diplodonta semiasperoides Nomura, 1932 þþ þlb
Fam. Veneridae Rafinesque, 1815
48. Dosinia japonica (Reeve, 1850) þþ þt/s
49. Callista brevisiphonata (Carpenter, 1864) þ þþlb
50. Saxidomus purpurata (G.B. Sowerby II, 1852) þs
51. Ruditapes philippinarum (Adams et Reeve, 1850) þþþþ þ þ þ þ s/b
52. Mercenaria stimpsoni (Gould, 1861) þ þþþþ þ þ þ s
53. Protothaca jedoensis (Lischke, 1874) þþþþþþþ þþs
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4416

are not necessarily synonyms, they may represent species existing
in other areas of Korea. At present, the Gangwon-do marine bivalve
molluscan fauna contains 159 species. A latitudinal species richness
gradient, reflecting changes in the number of species from north to
south along the eastern Korean coast (Lutaenko and Noseworthy
2014), can now be presented as follows: Gangwon, 159 species /
Gyeongbuk, 131 species /Gyeongnam, 183 species. The Gyeong-
buk fauna is obviously underestimated, and a greater collecting
effort should be made to reveal the species composition of bivalves
of that region of Korea. The taxonomy of some collected species
from Gangwon is complicated and requires some comment.
Taxonomic and distributional comments
Family Nuculidae Gray, 1824
Acila (Acila) vigilia Schenck, 1936
(Figure 2B, C)
Specimens examined. 1ex, Geumjin Port, 11 v 2016 KA Lutaenko &
RG Noseworthy), ZMFU no. 45790/Bv-7402.
Remarks.Acila (Acila)divaricata vigilia Schenck, 1936 was listed
as a subspecies of Acila (Acila)divaricata (Hinds, 1843) for Gangwon
and Gyeongsangbuk provinces of Korea by Lee and Min (2002), Min
et al (2004), and Lee (2014; as “A. divaricata vigila”). A. vigilia was
believed to be a synonym of A. divaricata (Kafanov 1991; Lutaenko
and Noseworthy 2012). The latter species was thought to be
recorded in Russian waters of the East Sea (Sea of Japan) only
around Moneron Island, southwestern Sakhalin (Scarlato 1981).
Japanese workers (Higo et al 1999; Okutani 2000) split A. divaricata
into four subspecies: A. divaricata divaricata (with a synonym,
Nucula mirabilis Adams et Reeve, 1850, also synonymized by
Scarlato (1981) (Boso Peninsula to central Japan Sea to Kyushu),
A. divaricata balabacensis Schenck, 1936 (Aichi Prefecture to South
China Sea and the Philippines), A. divaricata vigilia (Tohoku to
Hokkaido), and A. divaricata archibenthalis Okutani, 1964 (Sagami
Bay to Shikoku). Meanwhile, Chinese malacologists (Qi 2004; Xu
1984, 1999) separated A. divaricata and A. mirabilis and showed
that A. divaricata is distributed only from the East China to the
South China seas, whereas A. mirabilis is a member of the Yellow
Sea Cold Water Mass community. Later, this opinion was supported
by Huber (2010a):A. divaricata, originally described from China, is
smaller, generally less than 20 mm, with weaker ribbing, less
rostrate posteriorly, with a very fine regular ribbing on the inner
ventral margin, and is distributed in the Philippines and Chinese
waters and not found in Japan or in Russia. Recently, Zhang et al
(2014) clearly confirmed this geographical pattern and morpho-
logical separation by a genetic-molecular study: according to them,
A. mirabilis is commonly found in the Yellow Sea, the northern East
Sea (Sea of Japan), and the Russian Far Eastern seas. The distribution
of A. divaricata and A. mirabilis in the Yellow Sea and East China Sea
is separated by the Changjiang (Yangtze) River Estuary. Recent
shells from Sagami Bay depicted by Slodkewitsch (1967, pl. 4, Fig-
ures 2e4; as “Acila divaricata submirabilis”) also belong to
A. mirabilis. A. mirabilis is somewhat variable in convexity, thick-
ness, and rostration and has several synonyms: Nucula sculpta
Pease, 1860; Acila divaricata submirabilis Schenck, 1936;Acila
schencki Habe, 1958; and Acila archibenthalis Okutani, 1964 (Huber
2015). Omitted in literature, the recent Acila (Acila)divaricata bonini
Slodkewitsch, 1967 (type locality: Celebes (Sulawesi), Indonesia) is
probably a synonym of A. divaricata.
The large, biogeographically restricted A. vigilia (northern Hon-
shu, southern Hokkaido, Russia, and Sakhalin), originally described
as Acila (Acila)divaricata var. vigilia Schenck, 1936 (Schenck 1936,p.
101, pl. 17, Figures 1e6), with a strong black periostracum in fresh
specimens, appears distinct enough to be considered a valid species
(Huber 2010a; Zhang et al 2014). It is indeed somewhat closer to
A. divaricata than to A. mirabilis and by far the largest Acila known,
almost twice the size of A. mirabilis (Huber 2010a);however, A. vigilia
also differs from thetwo other species by its rather oblong shape, flat
umbo, and strong black periostracum (Zhang et al 2014). The holo-
type of A. divaricata vigilia is figured in color by Higo et al (2001) and
also at the website of the Smithsonian Institution (https://
collections.nmnh.si.edu/search/iz/). In this regard, our specimens
collected from fish nets in Geumjin Port are A. vigilia, as well as
Table 1 (continued )
Species Collecting sites Biogeographic characteristics
12345678910111213
54. P. euglypta (Sowerby, 1914) þþþ þs
55. P. adamsii (Reeve, 1863) þlb
56. Gomphina melanaegis Römer, 1860 þþþþþþþ þ þ þ þ s
57. G. multifaria (Kong, Matsukuma et Lutaenko, 2012) þþþþþ þs
58. Petricola habei Huber, 2010 þs
59. Irus ishibashianus Kuroda et Habe, 1952 þþs/b
60. Meretrix lusoria (Röding, 1798) þþþs
Fam. Corbulidae Lamarck, 1818
61. Anisocorbula venusta (Gould, 1861) þþs/b
62. Potamocorbula amurensis (Schrenck, 1861)þþs/b
Fam. Hiatellidae Gray, 1824
63. Hiatella orientalis (Yokoyama, 1920) þþ s/b
64. Panopea japonica A. Adams, 1850 þþ s/b
Fam. Solenidae Lamarck, 1809
65. Solen krusensternii Schrenck, 1867 þs/b
Fam. Pharidae H. et A. Adams, 1856
66. Siliqua pulchella (Dunker, 1852) þs
Fam. Lyonsiidae P. Fischer, 1887
67. Entodesma navicula (Adams et Reeve, 1850) þwdb þcb
Fam. Myochamidae Carpenter, 1861
68. Myadoropsis transmontana (Yokoyama, 1922) þþs
Fam. Pandoridae Rafinesque, 1815
69. Pandora pulchella Yokoyama, 1926 þs/b
70. P. wardiana Adams, 1860 þs/b
b/a, boreal-arctic species (including widely distributed boreal-arctic); lb, lowboreal species; s/b, subtropical-boreal (mostly subtropical-lowboreal) species; s, subtropical
species; t/s, tropical-subtropical species; wdb þcb, widely distributed boreal þcircumboreal species.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 17

specimens depicted from the northern East Sea (Sea of Japan) by
Scarlato (1981, photos 4e6; as “A. divaricata”). Min et al (2004, p. 366,
Figure 1163), as “A. divaricata vigila”, and Lee (2014, p. 129, Figure 5),
as “A. divaricata divaricata”, illustrated a typical A. mirabilis, whereas
Min’s“A. divaricata”(op. cit., p. 366, Figure 1162) and Lee’s(2013,p.
130, Figure 6) “A. divaricata vigila”are A. vigilia.“A. divaricata”,
collected in the southeastern bathyal area of the East Sea (Sea of
Japan) (Honshu) (Okutani and Saito 2014,Figure 2A) and from
Tatarsky Strait, northern East Sea (Sea of Japan) (Slodkewitsch 1967,
pl. 3, Figures 4, 6, non Figure 6), is obviously A. vigilia, indicating the
wide range of this species throughout this sea from south to north.
Lutaenko and Noseworthy’s“A. divaricata”(2012, pl. 1, Figures AeD)
from the northern East Sea (Sea of Japan), depth of 150 m, is A.
mirabilis. Thus, the Russian and Korean regions of the East Sea (Sea of
Japan) are inhabited by two species of Acila (Acila): A. vigilia and
A. mirabilis.
Figure 2. A, Acila insignis (Gould, 1861): Gangwon-do, site 12, shell length 8.0 mm, ZMFU no. 45789/Bv-7401; B and C, Acila vigilia Schenck, 1936: Gangwon-do, site 12, shell length
8.0 mm, ZMFU no. 45789/Bv-7401; site 11, right (B) and left (C) valves, shell length 26.3 mm, ZMFU no. 45790/Bv-7402; D, Anadara broughtonii (Schrenck,1867): Gangwon-do, site 1,
shell length 71.2 mm, ZMFU no. 46267/Bv-7548; E, Anadara talmiensis (Kalishevich, 1976): Gangwon-do, site 12, shell length 39.1 mm, ZMFU no. 46583/Bv-7570; F, Striarca
symmetrica (Reeve, 1844): Gangwon-do, site 12, shell length 12.1 mm, ZMFU no. 46588/Bv-7575; G and H, Arca boucardi Jousseaume,1894: Gangwon-do, site 11, right (G) and left (H)
valves, shell length 40.1 mm, ZMFU no. 46583/Bv-7570.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4418

Family Arcidae Lamarck, 1809
Anadara (Scapharca)talmiensis Kalishevich, 1976
(Figure 2E)
Specimens examined. 1ex, Geumjin Beach, 11 v 2016 (KA
Lutaenko and RG Noseworthy), ZMFU no. 46583/Bv-7570.
Remarks. A specimen from Geumjin Beach belongs to the species
complex “Anadara (Scapharca)inaequivalvis Bruguière (1789)”as it
has been identified for a long time in Korea, China, and Japan
(e.g. Habe 1965; Xu and Zhang 2008; Lee 2013). The taxonomy of this
anadarine species complex is very complicated, especially in the
tropicalIndo-Pacific whereit includes a number of speciesand names
to be resolved. Lutaenko (2006) showed that true A. inaequivalvis
from Southern India, its type locality, is clearly different from
northeastAsian “A. inaequivalvis aucct.”. Following Huber (2010a),we
believe that Japanese-Korean-Chinese “A. inaequivalvis”, including
subfossil Holocene species from the northwestern East Sea (Sea of
Japan) (Lutaenko 1988,1993), is a distinct species, and it differs from
Anadara (Scapharca)kagoshimensis (Tokunaga, 1906), although they
were often confused in the literature. Huber (2010a) incorrectly used
the name Anadara sativa (Bernard, Cai et Morton, 1993) for the
aforementioned species, which has been proposed as a nom. nov. pro
Arca subcrenata Lischke,1869 non Michelotti, 1861, a junior synonym
of A. kagoshimensis. It appears that the correct name for the northeast
Asian “A. inaequivalvis aucct.”is the little known, subfossil Anadara
talmiensis Kalishevich, 1976 (Kalishevich 1976) described from the
Holocene deposits of Talmi Lagoon, East Sea (Sea of Japan), near the
Russian-Korean border, whereas the allied Anadara masudai Noda,
1966 (Noda 1966) (Pleistocene, Tokyo Formation, Honshu) is a syn-
onym of A. kagoshimensis. The type locality of A. talmiensis is close to
Figure 3. A and B, Porterius dalli (Smith, 1885): Gangwon-do, site 11, left (A) and right (B) valves, shell length 29.2 mm, ZMFU no. 45804/Bv-7407; C, Glycymeris sp.: Gangwon-do,
site 9, shell length 15.5 mm, ZMFU no. 46582/Bv-7569; D, Glycymeris cf. yessoensis (G. B. Sowerby III, 1889): Gangwon-do, site 9, shell length 12.1 mm, ZMFU no. 46582/Bv-7569.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 19

Gangwon,and it is obvious that this species spread in the Holocene to
the northwestern East Sea (Sea of Japan) (Peter the Great Bay) due to
climatic warming periods and existed there between ca. 7000 and
500 years ago, now being locally extinct(Lutaenko 1993).This species
was extremely abundant in the Holocene semienclosed bays and
inner open bays in the Peter the Great Bay area, often found in the
Neolithic to Medieval shell-middens and natural outcrops (Lutaenko
1988; Lutaenko and Artemieva 2017; Rakov and Lutaenko 1997;
Sayenko et al 2015), but it appears to be rare at present along the
EastSea(SeaofJapan)coastofKorea(Lutaenko et al 2003; Lutaenko
2014)asitdoesnotfind suitable conditions in open, usually sand-
dominated coastal environments. The shell collected at Geumjin
appears subfossil and may be washed up on the beach from the
Holocene deposits.
Family Mytilidae Rafinesque, 1815
Mytilus (Mytilus) trossulus Gould, 1850
(Figure 7C, D)
Specimens examined. 22ex, Mangsan, 6 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 45925/Bv-7439; Yeomjeon, 7 v 2016 (KA
Lutaenko & RG Noseworthy), ZMFU no. 45951/Bv-7465; Yeongeok, 8
v 2016 (KA Lutaenko & RG Noseworthy), ZMFU no. 45937/Bv-7451;
Sacheon, 8 v 2016 (KA Lutaenko & RG Noseworthy), ZMFU no. 45950/
Bv-7464; Maenbaeng, 9 v 2016 (KA Lutaenko & RG Noseworthy),
ZMFU no. 45935/Bv-7439; Chuam, 9 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 45949/Bv-7463; Deoksan Port, 9 v 2016 (KA
Lutaenko & RG Noseworthy), ZMFU no. 45786/Bv-7398; Geumjin, 11
v 2016 (KA Lutaenko & RG Noseworthy), ZMFU no. 45936/Bv-7450.
Figure 4. A, Glycymeris cf. yessoensis (G. B. Sowerby III, 1889): Gangwon-do, shell length 38.8 mm, ZMFU no. 45855/Bv-7569; B, Glycymeris albolineata (Lischke, 1872): Gangwon-do,
site 1, shell length 57.2 mm, ZMFU no. 46579/Bv-7566; C, Glycymeris cf. yessoensis (G. B. Sowerby III, 1889): Gangwon-do, site 1, shell length 39.3 mm, ZMFU no. 46580/Bv-7567.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4420

Remarks. This species was found in eight localities along the
coast of Gangwon Province (Table 1), indicating its wide distribu-
tion, at least, in northern South Korea. First Je et al (1990) and then
Lutaenko and Noseworthy (2012) and Lee (2013) believed that all
previous records of “Mytilus edulis L., 1758”belong to the Medi-
terranean mussel Mytilus galloprovincialis Lamarck, 1819, an inva-
sive species in the East Sea (Sea of Japan) (see for review Lutaenko
and Kolpakov 2016). However, in the first color atlas of Korean
mollusks, true M. trossulus is figured under the name “M. edulis”
(Yoo 1976, pl. 23, Figures 9, 10); Kartavtsev et al (2005) genetically
detected hybrids between M. trossulus and M. galloprovincialis in
the Busan area. Therefore, we definitively record M. trossulus from
Gangwon-do and reinstate this species in the Korean fauna,
although it is also known from North Korea (Lutaenko and
Pretsiniek 2014). It appears that M. trossulus also has a wider dis-
tribution in Japan (Hokkaido and southwards) (Iwasaki et al 2017).
Family Pectinidae Wilkes, 1810
Scaeochlamys sp.
(Figure 10C, D)
Specimens examined. 5ex, Yeongeok, 8 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 46265/Bv-7546; Sacheon (KA Lutaenko &
RG Noseworthy), 8 v 2016, ZMFU no. 46264/Bv-7545; Samcheok-si,
Figure 5. A, Glycymeris yessoensis (G. B. Sowerby III, 1889): Gangwon-do, site 12, shell length 32.4 mm, ZMFU no. 46581/Bv-7568; B and C, Modiolus kurilensis Bernard, 1983:
Gangwon-do, site 11, right (B) and left (C) valves, shell length 4 4.7 mm, ZMFU no. 45814/Bv-7416; D and E, Modiolus kurilensis Bernard, 1983: Gangwon-do, site 11, right (D) and left
(E) valves, hell length 26.1 mm, ZMFU no. 45814/Bv-7416.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 21

9 v 2016 (KA Lutaenko & RG Noseworthy), ZMFU no. 46404/Bv-
7550; Geumjin, 11 v 2016 (KA Lutaenko & RG Noseworthy), ZMFU
no. 46266/Bv-7547.
Remarks. Our specimens can be provisionally identified as
Scaeochlamys cf. squamea Dijkstra and Maestrati, 2009. However,
they are atypical in having a finer radial sculpture than type spec-
imens (Dijkstra and Maestrati 2009; Chang et al 2011). S. squamea is
distributed in southern Japan, East China and South China seas, and
the Philippines to Australia with a bathymetric range of 5e50 m
(Djikstra 2013). Specimens depicted as “Chlamys irregularis
(Sowerby II, 1842)”(Min et al 2004, Figure 1306; Lutaenko and
Noseworthy 2012, pl. 20, Figures E, F) and “Chlamys lemniscata
(Reeve, 1853)”(Min et al 2004, Figure 1314; Lutaenko and
Noseworthy 2012, pl. 21, Figures A, B) known from Gangwon are
instead true Ch. squamea (H. Dijikstra, pers. comm., February 3 and
9, 2015).
Family Lucinidae Fleming, 1828
Pillucina pisidium (Dunker, 1860)
(Figure 12H)
Specimens examined. 1ex, Maenbaeng, 9 v 2016 (KA Lutaenko &
RG Noseworthy), ZMFU no. 48589/Bv-8068.
Remarks. For a long time, this species has been considered to be
subtropical, ranging from the East China Sea, Ryukyu Islands, and
Kyushu to Hokkaido and the East Sea (Sea of Japan) [Korea to Russia
(Peter the Great Bay)] (Lee and Min 2002; Higo et al 1999; Scarlato
1981). Glover and Taylor (2001) found that Japanese and Australian
specimens are very similar and considered them as conspecific and
defined the species range as Indo-West Pacific, extending from
Japan and China to East Africa and Madagascar, including Kenya,
Tanzania, Mozambique, Reunion, Mauritius, Seychelles, Maldives,
Andaman Islands, Thailand, Australia, Philippines, and New
Figure 6. A, Modiolus nipponicus (Oyama, 1950): Gangwon-do, site 2, shell length 37.4 mm, ZMFU no. 45782/Bv-7394; B and C, Mytilus galloprovincialis Lamarck, 1819: Gangwon-do,
site 10, right (B) and left (C) valves, shell length 107.4 mm, ZMFU no. 45769/Bv-7387.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4422

Caledonia. However, it appears they examined only a few speci-
mens from Japan and China. Northeast Asian specimens have a
weaker and less distinct sculpture and finer crenulation of the inner
margin. A molecular study is needed to resolve taxonomy.
Family Thyasiridae Dall, 1900
Conchocele scarlatoi Ivanova et Moskaletz, 1984
(Figure 12A, B)
Specimens examined. 1ex, Deoksan Port, 9 v 2016 (KA Lutaenko &
RG Noseworthy), ZMFU no. 46263/Bv-7544.
Remarks. A specimen collected from Gangwon is very similar to
Conchocele scarlatoi Ivanova et Moskaletz, 1984 (Figure 25B) found
from Peter the Great Bay, northwestern East Sea (Sea of Japan)
(Ivanova and Moskaletz 1984); it was previously reported from
Korea as Conchocele disjuncta (Gabb, 1866) (Kwon et al 2001;
Min et al 2004). However, Russian authors did not compare
C. scarlatoi to any fossil species from the northern Pacific, although
the diversity of the Cenozoic thyasirids is very high (Gladenkov et al
1984; Hickman 2015; Hryniewicz et al 2017; Kafanov et al 2001,
2002; Krishtofovich 1964; Moore 1988), and there are many fossil
and some recent forms which are very similar to C. scarlatoi
Figure 7. A and B, Mytilus cf. coruscus Gould, 1861: Gangwon-do, site 11, left (A) and right (B) valves, shell length 54.5 mm, ZMFU no. 45813/Bv-7415 (not typical specimen, possible
hybrid with Mytilus galloprovincialis Lamarck,1819); C and D, Mytilus trossulus Gould, 1850: Gangwon-do, site 10, lef t (C) and right (D) valves, shell length 68.1 mm, ZMFU no. 45786/
Bv-7398; E, Mytilus cf. coruscus Gould, 1861: Gangwon-do, site 11, left (E) and right (F) valves, shell length 34.5 mm, ZMFU no. 46584/Bv-7571.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 23

(Figure 25). Yabe and Nomura (1925, pl. 23, Figures 8, 10) figured
Thyasira bisecta Conrad, 1849 from the Neogene of Sakhalin and
Japan; this species is very closely related to C. scarlatoi.Conchocele
disjuncta var. alta (Krishtofovich, 1936) from the Upper Miocene
(Sakhalin, Kamchatka, and Sado Island, western Japan) and Con-
chocele disjuncta var. wajampolkana (Krishtofovich, 1936) from the
Pliocene of Kamchatka (Gladenkov et al 1984; Ilyina 1954;
Krishtofovich 1964; Slodkewitsch 1938) are other species close to
C. scarlatoi (Figure 25C). The American Cenozoic Conchocele bath-
yaulax Hickman, 2015 (upper Eocene to lower Oligocene) closely
resembles C. scarlatoi: This species is relatively higher and more
quadrate in outline than C. bisecta and has a more evenly and more
broadly rounded ventral margin and a straighter anterior margin
(Hickman 2015).
Some authors have synonymized C. scarlatoi with C. bisecta
(Coan et al 2000; Lutaenko and Noseworthy 2012), whereas others
separate them (Kafanov 1991; Kamenev et al 2001). As Oliver and
Frey (2014) mentioned, although there has been considerable
debate on the identity and number of living species of Conchocele,
including the relationship between C. bisecta and C. disjuncta
(Bernard 1972; Coan et al 2000; Habe 1958, 1977; Kamenev et al
2001), the current view is that only a single species, C. bisecta,is
Figure 8. A, Septifer virgatus (Wiegmann, 1837): Gangwon-do, site 3, shell length 31.7 mm, ZMFU no. 45812/Bv-7414; B, Septifer virgatus (Wiegmann, 1837): Gangwon-do, site 3,
shell length 37.8 mm, ZMFU no. 45812/Bv-7414; C and D, Septifer keenae Nomura, 1936: Gangwon-do, site 11, right (C) and left (D) valves, shell length 19.0 mm, ZMFU no. 45856/Bv-
7434; E, Crassostrea gigas (Thunberg, 1793): Gangwon-do, site 10, shell height 95.1 mm, ZMFU no. 45776/Bv-7388; F, Anomia chinensis Philippi, 1849: Gangwon-do, site 5, shell
length 38.5 mm, ZMFU no. 45787/Bv-7399.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4424

extant. C. bisecta was previously distinguished from C. disjuncta by
the concave outline of the anterior surface and the more promi-
nent umbones (Bernard 1972). Hryniewicz et al (2017) pointed out
that both C. bisecta and C. disjuncta are based on fossil rather than
extant material, and each fossil species was erected based on the
material of different ages (Miocene and Pliocene, respectively), and
thus caution should be taken when applying these names to extant
material. However, there is a name for a North PacificConchocele
which has been applied to a living species, Thyasira disjuncta var.
ochotica Krishtofovich, 1936; the holotype was designated based
on recent material from the United States National Museum.
“C. ochotica”might be used for extant North Pacific species
(Hryniewicz et al 2017). Krishtofovich (1964) later selected
another “lectotype”for T. disjuncta ochotica from fossil material,
but this action is not valid. Kharlamenko et al (2016) believe that
C. scarlatoi differs from C. bisecta in having a more pointed umbo
and an apical angle of less than 90

, which are characters that do
not change with age; these authors also provide a good image of
the holotype of C. scarlatoi. A molecular study is needed to resolve
this issue. However, if C. scarlatoi is proven to be a separate species,
C. alta (Krishtofovich, 1936) and C. wajampolkana (Krishtofovich,
1936) should be regarded as prior available names for this recent
species.
Family Trapezidae Lamy, 1920
Trapezium (Neotrapezium) liratum (Reeve, 1843)
(Figure 12F, G)
Specimens examined. 1ex, Geumjin, 11 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 45819/Bv-7421.
Figure 9. A, Crassostrea nippona (Seki, 1934): Gangwon-do, site 10, shell height 45.9 mm, ZMFU no. 45777/Bv-7389; B, Crassostrea nippona (Seki, 1934): Gangwon-do, site 10, shell
height 59.6 mm, ZMFU no. 45777/Bv-7389; C, Spondylus cruentus Lischke, 1868: Gangwon-do, site 8, shell length 40.1 mm, ZMFU no. 45818/Bv-7420; D, Chlamys swiftii (Bernardi,
1858): Gangwon-do, site 11, shell height 43.0 mm, ZMFU no. 45805/Bv-7408.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 25

Remarks. This species has been reported from the west and
southwest coasts of Korea as far as Busan (Min et al 2004). Based on
available faunal and ecological studies, it has previously never been
recorded in Gangwon and Gyeongsangbuk provinces (Lutaenko
and Noseworthy 2012; Lutaenko et al 2002, 2003; Park et al 2011
and bibliography therein; Lutaenko 2014). It appears that this
tropicalesubtropical species has a disjunct distribution in the East
Sea (Sea of Japan), now being known from Busan, Gangwon, where
it is rare, and Peter the Great Bay in the Russian region, where it
occurs only in semienclosed bays. This species is associated with
oyster settlement and lives in protected, low-energy environments
(Kolpakov and Kolpakov 2013; Kondo et al 2009). It is now en-
dangered due to low population density, limited spatial distribu-
tion, and relic origin in both Peter the Great Bay and lagoons in
northern Honshu, Japan (Kondo et al 2009; Lutaenko 2004).
Family Cardiidae Lamarck, 1809
Clinocardium (Ciliatocardium) likharevi (Kafanov in Scarlato,
1981)
(Figure 13AeD)
Specimens examined. 12ex, Deoksan Port, 9 v 2016 (KA Lutaenko
& RG Noseworthy), ZMFU no. 45781/Bv-7393.
Remarks. An allied species, Clinocardium (Ciliatocardium) cil-
iatum (Fabricius, 1790), a well-known, widely distributed boreal-
arctic species, has been reported from the eastern coast of Korea
(Gangwon and Gyeongbuk) (Lutaenko and Noseworthy 2012; Min
et al 2004). This represents the southernmost record of this spe-
cies along the western coast of the East Sea (Sea of Japan). C.
likharevi, described from Tatarsky Strait, western Sakhalin, in the
northern East Sea (Sea of Japan) (Scarlato 1981), has not been
Figure 10. A and B, Chlamys farreri (Jones et Preston, 1904): Gangwon-do, site 11, shell height 62.3 mm, ZMFU no. 45807/Bv-7410; C, Scaeochlamys cf. squamea Dijkstra and
Maestrati, 2009: Gangwon-do, site 5, shell height 18.9 mm, ZMFU no. 46264/Bv-7545; D, Scaeochlamys cf. squamea Dijkstra and Maestrati, 20 09: Gangwon-do, site 5, shell
height 11.7 mm, ZMFU no. 46264/Bv-7545.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4426

recognized in the Japanese fauna and was provisionally synony-
mized with C. ciliatum by Coan et al (2000), although at present,
World Register of Marine Species (WoRMS) regards it as a separate
species (Poorten 2014). C. likharevi differs from C. ciliatum by its
equilateral, nonoblique, and unbent shell, by the anterior concavity
of its dorsal margin, and by its relatively shorter nymph (on
average, no more than 1/5 of the shell length) (Kafanov 2001). The
significant morphological variability of C. ciliatum was subdivided
into five subspecies by Kafanov (2001), and this prevented Japanese
and Korean workers from identifying C. likharevi. We previously
recognized and figured this species for the middle Primorye area
(Lutaenko 1999), Amursky Bay (Lutaenko 2002), Ussuriysky Bay
(Lutaenko 2005), and Peter the Great Bay (Lutaenko and Volvenko
2017) in the East Sea (Sea of Japan), which is in accord with
Kafanov’s (2001) data. It is also known from the Okhotsk Sea coast
of Sakhalin (Aniva and Terpenya bays), South Kurile Islands, and
Nagasaki, Kyushu, Japan (l.c.). Our finding of C. likharevi at Deoksan
Port, Gangwon Province, is the first record for Korea. In Russia, it
has been collected at a depth of 33e117 m on mud and sand
(Kafanov 2001).
Family Psammobiidae Fleming, 1828
Gari (Psammotaena) chinensis (Deshayes, 1855)
(Figure 16E, F)
Figure 11. AeE, variability of Mizuhopecten yessoensis (Jay, 1857): Gangwon-do, site 11, shell height 55.8 mm (A), 30.00 mm (B), 44.6 mm (C), 30.3 mm (D), 34.9 mm (E), ZMFU no.
45806/Bv-7409.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 27

Specimens examined. 1ex, Yeomjeon, 7 v 2016 (KA Lutaenko &RG
Noseworthy), ZMFU no. 45779/Bv-7391.
Remarks. This species has been identified in Japanese literature
as “Psammotaea virescens (Deshayes, 1855)”(Higo et al 1999;
Okutani 2000). It is believed to be distributed in Japan from
Tokyo and Wakasa bays southward to the Indian Ocean, living in
the muddy bottom of embayments (Okutani 2000). This species has
not been found along the continental coast of the East Sea (Sea of
Japan) (Lutaenko and Noseworthy 2012) but has been recorded
from other localities in Korea (Kwon et al 2001; Min et al 2004).
According to Huber (2010a),Gari virescens (Deshayes, 1855) is close
to Gari chinensis (Deshayes, 1855), but these are two distinct spe-
cies: G. virescens is smaller and more narrowly elongate, with a
strongly rounded, broader, and shorter pallial sinus. Figured syn-
types of both species are found in the study by Higo et al (2001).
However, “G. virescens”of Japanese authors represents instead
G. chinensis; true G. virescens is found in southern China, the
Philippines, and westward, but not in Japan or eastern China
(Huber 2010a). Synonyms of G. chinensis are Soletellina moesta
Lischke, 1872, from Tokyo Bay (type locality) (Lischke 1872), and
Figure 12. A and B, Conchocele scarlatoi Ivanova et Moskaletz, 1984: Gangwon-do, site 10, left (A) and right (B) valves, shell length 47.6 mm, ZMFU no. 46263/Bv-7544; C, Mya-
doropsis transmontana (Yokoyama,1922): Gangwon-do, site 1, shell length 8.0 mm, ZMFU no. 45811/Bv-7413; D and E, Cardita leana Dunker, 1860: Gangwon-do, site 5, right (D) and
left (E) valves, shell length 16.6 mm, ZMFU no. 45791/Bv-7403; F and G, Trapezium liratum (Reeve, 1843): Gangwon-do, site 12, right (F) and lef t (G) valves, shell length 18.0 mm,
ZMFU no. 45819/Bv-7421; H, Pillucina pisidium (Dunker, 1860): Gangwon-do, site 9, shell length 8.2 mm, ZMFU no. 48589/Bv-8068; I and J, Kellia japonica Pilsbry, 1895: Gangwon-
do, site 11, left (I) and right (J) valves, shell length 10.7 mm, ZMFU no. 46589/Bv-7576.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4428

Sanguinolaria (Psammotaea)castanea Scarlato, 1965, from China
[unnecessary nom. nov. pro Capsa (Capsella) chinensis Deshayes,
1855, non Solenotellina chinensis Mörch, 1853 (Scarlato 1965)(¼
Hiatula chinensis (Mörch, 1853)]. Our material is obviously
G. chinensis and represents the first record for Korea. “Psammotaea
virescens”of Korean workers (Kwon et al 2001; Min et al 2004)is
Gari (Psammotaena) elongata (Lamarck 1818). For taxonomy, syn-
onymy, and illustrations, see Willan (1993); Huber (2010a).
Family Veneridae Rafinesque, 1815
Petricola (Petricola) habei Huber, 2010
(Figure 19C)
Specimens examined. 2ex, Geumjin, 11 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 45850/Bv-7427.
Remarks. This species has been known in Korean and Japanese
literature as Pseudoirus mirabilis (Deshayes, 1853) (Higo et al 1999;
Kwon et al 2001; Min et al 2004; Okutani 2000) in Petricolidae,
now a subfamily of Veneridae. This species is distributed from
southern Hokkaido and southward to Kyushu, Korea, and the Yel-
low Sea, living from the intertidal zone to a depth of 20 m (Higo
et al 1999). Chinese workers (Xu and Zhang 2008; Zhang et al
2016) do not list this species for China. Coan (1996, 1997) synony-
mized Pseudoirus Habe, 1951 with Petricola Lamarck, 1801, as their
sculpture is similar. Moreover, Coan (1996, 1997) showed that
Petricola mirabilis Deshayes, 1853, described from Monterey, Cali-
fornia [type is illustrated by Coan (1997, Figure 9)] is a synonym of
Petricola (Petricola) carditoides (Conrad, 1837), an American species.
Japanese and Korean authors, following Lischke (1871), used
“mirabilis”for an Asian Petricola until Huber (2010b) redescribed it
Figure 13. A, Clinocardium likharevi (Kafanov in Scarlato, 1981): Gangwon-do, site 10, shell length 33.4 mm, ZMFU no. 45781/Bv-7393; B, Clinocardium likharevi (Kafanov in Scarlato,
1981): Gangwon-do, site 10, shell length 27.8 mm, ZMFU no. 45781/Bv-7393; C and D, Clinocardium likharevi (Kafanov in Scarlato, 1981): Gangwon-do, site 10, left (C) and right (D)
valves, shell length 23.8 mm, ZMFU no. 45781/Bv-7393.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 29

as Petricola (Petricola) habei Huber, 2010; type locality Nobi,
Kanagawa Prefecture, Honshu, Japan.
Meretrix lusoria (Röding, 1798)
(Figure 21A, B)
Specimens examined. 2ex, Maenbaeng, 9 v 2016 (KA Lutaenko &
RG Noseworthy),ZMFU no. 48592/Bv-8071; Geumjin,11 v 2016 (KA
Lutaenko & RG Noseworthy), ZMFU no. 46587/Bv-7574.
Remarks. This species has been reported from Busan on the
southeastern coast of Korea (Lutaenko and Noseworthy 2012) but is
also distributed from southern Hokkaido and southward in Japan
(Higo et al 1999). It is quite abundant in the tidal flats of western
Korea and the Yellow Sea. The genetically and morphologically
close Meretrix petechialis Lamarck, 1818 (Zhang et al 2012) can form
hybrids with M. lusoria on the southwestern coast of Korea, at
Ganjin Bay (Torii et al 2010), and is found in Gyeongnam (Min et al
2004) but not distributed further north. As is the case with
T. liratum (Trapezidae), M. lusoria has a disjunct distribution in the
East Sea (Sea of Japan), being known from Busan and Gangwon
Province, and it has also been found in mid-Holocene deposits of
Peter the Great Bay, Russia (Evseev 1981; Rakov and Lutaenko 1997;
Sharova and Rakov 2011). However, we do not rule out the possi-
bility that our Gangwon shells may be washed out of the Holocene
lagoonal/estuarine deposits containing relic shells such as
A. talmiensis,T. liratum, and Mactra quadrangularis Deshayes in
Figure 14. A, Potamocorbula amurensis (Schrenck, 1861): Gangwon-do, site 12, shell length 9.5 mm, ZMFU no. 45792/Bv-7404; B, Anisocorbula venusta (Gould, 1861): Gangwon-do,
site 9, shell length 7.1 mm, ZMFU no. 46586/Bv-7573; C, Megangulus zyonoensis (Hatai et Nisiyama, 1939): Gangwon-do, site 12, shell length 94.7 mm, ZMFU no. 45784/Bv-7396; D,
Megangulus venulosus (Schrenck, 1861): Gangwon-do, site 5, shell length 62.5 mm, ZMFU no. 45821/Bv-7423; E, Cadella lubrica (Gould, 1861): Gangwon-do, site 12, shell length
13.1 mm, ZMFU no. 45820/Bv-7422; F, Nitidotellina sp.: Gangwon-do, site 4, shell length 20.9 mm, ZMFU no. 46590/Bv-7577.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4430

Reeve, 1854. They cannot be discarded as restaurant shells as their
size is too small, 19e34 mm in length, and we have never seen such
small specimens of M. lusoria in Korean markets.
Gomphina (Macridiscus) multifaria (Kong, Matsukuma et
Lutaenko, 2012)
(Figure 20E, F)
Specimens examined. 34ex, Yeomjeon, 7 v 2016 (KA Lutaenko &
RG Noseworthy), ZMFU no. 43122/Bv-7430; Sacheon, 8 v 2016 (KA
Lutaenko & RG Noseworthy), ZMFU no. 46037/Bv-7518; Yeongeok,
8 v 2016 (KA Lutaenko & RG Noseworthy), ZMFU no. 46035/Bv-
7516; Maenbaeng, 9 v 2016 (KA Lutaenko & RG Noseworthy), ZMFU
no. 46038/Bv-7517; Geumjin, 11 v 2016 (KA Lutaenko & RG Nose-
worthy), ZMFU no. 46038/Bv-7519.
Remarks. This species was long known under the names “Gom-
phina aequilatera (G.B. Sowerby I, 1825)”or “Gomphina veneriformis
(Lamarck, 1818)”, and these two names are still in use in ecological
and related literature in Korea. The former name is a nomem
dubium or nomen nudum, and the latter one is based on a spec-
imen of Donacidae (Lutaenko 2001). In regional literature, this
species was often confused or synonymized with Gomphina
(Macridiscus) melanaegis Römer, 1860 (Figure 22AeD), although
Yoo ( 1976) and Lutaenko (2001) clearly separated them. After
molecular confirmation, it was redescribed as G. multifaria (Kong
et al 2012). We would like to stress that some specimens
collected from Gangwon have an intermediate shell shape between
that of G. multifaria and G. melanaegis, and we regard them as hy-
brids. Hybridization of these two species was first morphologically
detected in the eastern East Sea (Sea of Japan) (Tanaka 1979).
Figure 15. AeD, variability of Mactra chinensis Philippi, 1846: Gangwon-do, site 5, shell length 52.2 mm (A), 49.2 mm (B), 52.7 mm (C), 44.5 mm (D), ZMFU no. 45815/Bv-7417.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 31

Family Corbulidae Lamarck, 1818
Potamocorbula amurensis (Schrenck, 1861)
(Figure 14A)
Specimens examined. 2ex, Geumjin, 11 v 2016 (KA Lutaenko & RG
Noseworthy), ZMFU no. 45792/Bv-7404.
Remarks. From two to four species of Potamocorbula Habe, 1965
are known from northeastern Asia (Higo et al 1999; Zhuang and
Cai 1983). Korean species, including Gangwon specimens, were
believed to belong to Potamocorbula ustulata (Reeve, 1844) (Min
et al 2004; Park et al 2011). Owada et al (2013) showed that
Potamocorbula spp. distributed in the waters adjacent to Japan and
Korea are divided into two groups on the basis of morphological
and genetic characteristics; those groups are closely related
genetically, but reproductive isolation between them is possible,
and they can be treated as at least two distinct species. One spe-
cies lives in the Yellow Sea, at Ganghwa-do, and another one
around Korea and Japan (P. a mure nsi s ); however, specimens from
Mae-ho Lake in Gangwon Province and Hinuma Lake in Japan can
be morphologically identified as P. us tula t a (Owada et al 2013).
The status of the latter species is unclear: Corbula laevis Hinds,
1843 and Corbula ustulata Reeve, 1844 are regarded as earlier
names for this species complex (Coan 2002). P. amurensis is
extremely variable in shell shape and other parameters.
Figure 16. A, Spisula sachalinensis (Schrenck, 1861): Gangwon-do, site 5, shell length 64.8 mm, ZMFU no. 45816/Bv-7418; B and C, Nuttallia obscurata (Reeve, 1857): Gangwon-do,
site 9, outer (B) and inner (C) views of shell, shell length 35.9 mm, ZMFU no. 45785/Bv-7397; D, Mactra quadrangularis Deshayes in Reeve, 1854: Gangwon-do, site 1, shell length
33.4 mm, ZMFU no. 48590/Bv-8069; E and F, Gari chinensis (Deshayes, 1855): Gangwon-do, site 4, shell length 29.9 mm, ZMFU no. 45779/Bv-7391.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4432

Specimens from Gangwon and the northwestern East Sea (Sea of
Japan) (Lutaenko and Noseworthy 2012; Lutaenko and Volvenko
2017; Scarlato 1981) generally correspond to the original
description and illustrations of P. a m ure n sis (Schrenck, 1861), and
the type material is figured here for the first time (Figure 26).
P. amurensis is also listed for the Holocene deposits of Gangwon
lagoons (Nakanishi et al 2017; Yum et al 2004). Huber (2010a)
believes that there are few doubts that Corbula nimbosa Hanley,
1843 (now Potamocorbula) is the same as Corbula labiata Reeve,
184 4 a nd C. ustulata from Singapore. According to him, P. nim bos a
is similar to P. am urens i s but has a larger and thicker shell with a
more trigonal, deeper pallial impression. Tsuchida and Okamura
(1997) found that P. ustu lata from Shanghai, China, is different
from the Japanese P. am u ren s is and P. la evi s, a distinct species that
has recently immigrated from China to the Ariake Sea in Japan (see
also: Horikoshi and Okamoto 1994). Thus, it appears that
P. am u ren sis ,P. laevis,andP. n imb o sa (¼P. ustulata) are three
distinct species, but this conclusion requires further molecular
confirmation. Eastern Korean and Russian populations belong to
P. am u ren sis .
Biogeographic and ecological remarks
The bivalve molluscan fauna of Gangwon is biogeographically
extremely important as it contains a composition of “northern”and
“southern”species coexisting in the East Sea (Sea of Japan), a
unique “mini-ocean”with strong currents and contrasting ocean-
ographic conditions. This sea lies in two different geographical and
climatic zones: temperate in the north, with a subarctic regime in
winter, and subtropical in the south, thus making the fauna a
Figure 17. A, Macoma incongrua (Martens, 1865): Gangwon-do, site 3, shell length 29.0 mm, ZMFU no. 45822/Bv-7424; B, Macoma sectior (Oyama, 1950): Gangwon-do, site 6, shell
length 51.0 mm, ZMFU no. 46585/Bv-7572; C, Macoma irus (Hanley, 1844): Gangwon-do, site 3, shell length 46.2 mm, ZMFU no. 45849/Bv-7426; D, Macoma calcarea (Gmelin,1791):
Gangwon-do, site 10, shell length 36.1 mm, ZMFU no. 45848/Bv-7425.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 33

combination of species of various origins, boreal, boreal-arctic, and
tropical, which appears to have been recognized originally by
Schrenck (1867). The sea is divided by a frontal zone into two
distinct areas: the southeastern warm-water area washed by the
Tsushima Current with three branches, and the northwestern area
dominated by the cold Liman(ian), Primorskoye, and North Korean
Currents. The western part of the East Sea (Sea of Japan) is inhabited
by 371 species and subspecies of bivalve mollusks: 316 species are
known for South Korea, 50 species for North Korea, and in Russia,
163 species for southern Primorye (Peter the Great Bay) and 130
species for middle/northern Primorye, the area of the continental
coast of this sea from Cape Povorotny to Tatarsky Strait (Lutaenko
2017). The South Korean fauna is quite rich and is comparable
with that of the East China Sea, which has 337 species (Xu and
Zhang 2011). Species richness of the Gangwon bivalve fauna (159
species) is comparable to that of Peter the Great Bay (163 species) in
the Russian region, and it constitutes half of the fauna of the East
Sea (Sea of Japan) coast of South Korea (50.4%). In the course of our
study, we collected 70 species or 44% of the total Gangwon fauna;
17 species, 11% of this fauna, were recorded for the first time.
According to a preliminary biogeographic analysis (Lutaenko
and Noseworthy 2014), the Gangwon fauna has a rather high pro-
portion of species found only in this province, 57 or 40% of the total
provincial fauna, and are not known southward. However, in
contrast to Gyeongnam, the “endemics”are predominantly boreal
species: widely distributed boreal, circumboreal, and lowboreal,
with an admixture of some warm-water mollusks. Gangwon is very
rich in cold-water families; for example, the Nuculanidae and
Figure 18. A, Saxidomus purpurata (G.B. Sowerby II,1852): Gangwon-do, site 3, shell length 51.5 mm, ZMFUno. 45851/Bv-7428; B, Protothaca jedoensis (Lischke, 1874): Gangwon-do,
site 1, shell length 20.5 mm, ZMFU no. 45852/Bv-7429; C, Protothaca euglypta (Sowerby, 1914): Gangwon-do, site 3, shell length 30.4 mm, ZMFU no. 46039/Bv-7520; D, Mercenaria
stimpsoni (Gould, 1861): Gangwon-do, site 12, shell length 79.5 mm, ZMFU no. 45853/Bv-7431.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4434

Yoldiidae are well represented within Korean East Sea (Sea of
Japan) waters but mainly in Gangwon. Many mainly boreal genera
such as Robaia (Nuculanidae), Portlandia,Megayoldia (Yoldiidae),
Dacrydium (Mytilidae), Parvamussium (Propeamussiidae), Serripes
(Cardiidae), Mactromeris (Mactridae), Liocyma and Turtonia (Ven-
eridae), Bankia (Teredinidae), Panomya (Hiatellidae), and Thracia
(Thraciidae) are recorded only from Gangwon; two families,
Nuculanidae and Thraciidae, are found exclusively in Gangwon
(Lutaenko and Noseworthy 2012). Species belonging to these
genera are rather deep-water faunal elements inhabiting mainly
the lower subtidal zone in the northern East Sea (Sea of Japan),
although they may live in shallow water in high latitudes. Among
the species encountered in Gangwon Province for the first time, in
the course of this study, the majority are warm-water mollusks
(subtropical, tropicalesubtropical, and subtropicalelowboreal),
and only three are boreal species (lowboreal and circumboreal).
Thus, we extended the northern limit of distribution in Korean
waters for 14 bivalve species.
As mentioned previously, the regional distribution of some
warm-water bivalves in the East Sea (Sea of Japan) is obscure. These
include Anadara talmiensis (Kalishevich, 1976), Trapezium liratum
(Reeve, 1843), Mactra quadrangularis Deshayes in Reeve, 1854,
Macoma tokyoensis Makiyama, 1927, and Meretrix lusoria (Röding,
1798). They were not previously known from Gangwon (except for
M. quadrangularis)(Lutaenko and Noseworthy 2012; Min et al
2004) but presently live further north, except for the regionally
extinct A. talmiensis, in Peter the Great Bay on the Russian coast
(42

N). There they are found only in semienclosed bays with
Figure 19. A, Dosinia japonica (Reeve, 1850): Gangwon-do, site 12, shell length 9.3 mm, ZMFU no. 46056/Bv-7435; B, Irus ishibashianus Kuroda et Habe, 1952: Gangwon-do, site 9,
shell length 9.9 mm, ZMFU no. 48591/Bv-8070; C, Petricola habei Huber, 2010: Gangwon-do, site 12, shell length 14.4 mm, ZMFU no. 45850/Bv-7427; D, Callista brevisiphonata
(Carpenter, 1864): Gangwon-do, site 10, shell length 9.9 mm, ZMFU no. 46057/Bv-7536.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 35

intense summer warming, where they are considered to be relic
faunal elements left after the Holocene warm periods, especially
the climatic optimum (ca. 5000e6000 years ago) (Lutaenko 1991).
These species are characteristic of the fauna of the tidal flats of the
Yellow Sea and are found in the Korea Strait (South Sea) including
Holocene deposits around Busan (Oh 1994). Thus, a combination of
factors such as the considerable indentation of the coastline,
including “ria”type bays with shallow-water, semienclosed areas at
their upper ends, and penetration of the warm Tsushima waters to
the northwestern East Sea (Sea of Japan) which intensified the ef-
fect of local warming, resulting in the formation of a subtropical-
type molluscan fauna in some areas of the northern East Sea (Sea
of Japan) during the Middle Holocene. The same pattern of Holo-
cene shifts of distribution of bivalve mollusks is known for Hok-
kaido (Matsushima 1984). Some workers call these warm-water
faunas that existed in the Holocene as “thermally anomalous
molluscan assemblages”(TAMA) (DeVries and Wells 1990)or
interzonal faunas in modern seas (Golikov et al 1990).
However, recent paleogeographic and sedimentological studies
have revealed that warm-water mollusks existed in Gangwon in
the Holocene. In the Holocene deposits of the brackish-water
Hwanjinpo Lagoon, located very close to the area studied (38

27
0
Figure 20. A and B, Gomphina melanaegis Römer, 1860: Gangwon-do, site 1, outer (A) and inner (B) views of shell, shell length 31.6 mm, ZMFU no. 45780/Bv-7392; C and D,
Gomphina melanaegis Römer, 1860: Gangwon-do, site 1, outer (C) and inner (D) views of shell, shell length 45.2 mm, ZMFU no. 45780/Bv-7392; E and F, Gomphina multifaria (Kong,
Matsukuma et Lutaenko, 2012): Gangwon-do, site 4, left (E) and right (F) valves, length 31.6 mm, ZMFU no. 43122/Bv-7430.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4436

N, 128

28
0
E), a typical warm-water embayment assemblage of
bivalves was identified: Potamocorbula amurensis (Schrenck, 1861)
(mostly at the late stages of the lagoon development), M. tokyoensis,
Dosinia japonica (Reeve, 1850), Ruditapes philippinarum (Adams et
Reeve, 1850), and “Anadara subcrenata”[¼A. kagoshimensis
(Tokunaga, 1906)] (Yum et al 200 4). P. amurensis,Crassostrea gigas
(Thunberg, 1793), T. liratum, and Corbicula japonica Prime, 1864
were collected from the Late Holocene deposits of Mae-ho Lagoon
(37

57
0
0
00
N, 128

46
0
12
00
E) in Gangwon (Nakanishi et al 2017).
This indicates that limited embayment, semienclosed bay condi-
tions existed in the open, high-energy coastal environment of
Gangwon Province and further north during the Holocene, thus
enabling warm-water “Yellow Sea tidal flat species”populations to
exist here and serve as “seed populations”for the Holocene
molluscan migrations to the Russian coast in the northwestern East
Sea (Sea of Japan), where they found suitable conditions and
flourished in the Middle to Late Holocene (Lutaenko 1988;
Lutaenko and Artemieva 2017). These species (A. talmiensis,
M. quadrangularis,T. liratum, and M. lusoria) found during the
present study may represent either elements of the past Holocene
populations as empty shells washed ashore from eroded Holocene
deposits, or they may still live in some sheltered habitats in low
abundance. These species were not found further south in Yeongil
Bay (Lutaenko et al 2003) that has summer temperatures more
closely related to those critical for their successful reproduction.
Thus, despite cold water currents along the coast of Gangwon
(Kim and Kim 1983), tropicalesubtropical and subtropical species
still survive in this area and may also exist in some localities in
Figure 21. A, Meretrix lusoria (Röding, 1798): Gangwon-do, site 12, shell length 34.3 mm, ZMFU no. 46587/Bv-7574; B, Meretrix lusoria (Röding, 1798): Gangwon-do, site 9, shell
length 19.4 mm, ZMFU no. 48592/Bv-8071; C, Ruditapes philippinarum (Adams et Reeve, 1850): Gangwon-do, site 3, shell length 44.7 mm, ZMFU no. 46047/Bv-7526; D, Protothaca
adamsii (Reeve, 1863): Gangwon-do, site 10, shell length 14.7 mm, ZMFU no. 45854/Bv-7432.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 37

North Korea. An understanding of their precise distribution along
the East Sea (Sea of Japan) coast of Korea is important in relation to
historical biogeography, faunal origins, and expected faunal
changes due to global warming.
Some species collected in our study are very abundant in the
intertidal and upper subtidal zones of Gangwon and are of com-
mercial importance; their biology and ecology have been studied in
this area by previous workers, and these data supplement
biogeographic considerations. The Japanese scallop Mizuhopecten
yessoensis (Jay, 1857) grows quickly around Jumunjin within the
water column from 10e30 m, with temperatures ranging from 5 to
23

C and with the highest growth in cages at a water depth from 10
to 15 m; however, a water depth below 25 m was best to avoid mass
mortality during high water temperature periods (Park et al 2000).
This is in accord with data from Peter the Great Bay in the
northwestern East Sea (Sea of Japan); the largest specimens of
Japanese scallops were found in sites with good water exchange
and relatively stable temperatures, at a depth of approximately
20 m, although it lives there at a depth range of 4e80 m (Scarlato
1981). Spawning in scallop populations starts at a temperature
range of 7e9

C(Ivin and Kalashnikov 2005), similar to that in
Gangwon, 7e11

C(Uddin et al 2007). This species may demon-
strate the so-called biogeographic submergence: relatively cold-
water animals are found in deeper areas at the edge of their ranges.
Mass mortality of farmed M. yessoensis has been observed in
Gangwon in August due to high temperatures (Jo et al 2009). This
species was found only at a depth of 25e52 m in outer Yeongil Bay,
in Gyeongbuk (Lutaenko et al 2003), the southernmost limit of its
geographic range. The geoduck clam, Panopea japonica A. Adams,
1850 was found to inhabit relatively deep water around Geumjin at
Figure 22. A, Panopea japonica A. Adams, 1850: Gangwon-do, site 10, shell length 107.4 mm, ZMFU no. 45778/Bv-7390; B and C, Hiatella orientalis (Yokoyama, 1920): Gangwon-do,
site 11, left (B) and right (C) valves, shell length 18.6 mm, ZMFU no. 45808/Bv-7411.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4438

about 10e50 m, but mainly at a depth range of 20e35 m, in sand, at
a temperature range of 7e21

C(Kim et al 1991)or5e17

C(Lee et al
1998). This is why this species was not found in beach thanato-
coenoses but only in fishing nets at Deoksan and Geumjin ports.
In the intertidal zone, Septifer virgatus (Wiegmann, 1837) is very
abundant along the Gangwon coastline, especially among rocks
exposed to strong wave action (Kim et al 1983). Furthermore, dense
intertidal settlements of M. galloprovincialis were observed in the
1990s (Lutaenko et al 2002), and this mussel is also dominant on
artificial reefs (Kim et al 2008). A subtidal study around Gangneung
revealed the abundance of a small kelliellid, Alveinus ojianus
(Yokoyama, 1927) and Mactra chinensis Philippi, 1846 (Choi et al
2000); the latter species was frequent in beach thanatocoenoses.
The large bivalves Spisula sachalinensis (Schrenck, 1861),
M. chinensis,Megangulus venulosus (Schrenck, 1861), Mercenaria
stimpsoni (Gould, 1861), Siliqua pulchella (Dunker, 1852), and Solen
krusensternii Schrenck, 1867 (as “Solen corneus”) were abundant in
dredge samples at a depth of 9e12 m around Gangneung (An et al
2014).
In our beach collections, shells of Gomphina melanaegis Römer,
1860 were abundant at Mangsan (site 1), Yeomjeon (site 4), Chuam
(site 7), Maenbang (site 9), and Geumjin (site 12). Other common
Figure 23. A, Diplodonta semiasperoides Nomura, 1932: Gangwon-do, site 5, shell length 18.1 mm, ZMFU no. 48593/Bv-8072; B and C, Felaniella usta (Gould, 1861): Gangwon-do, site
12, shell length 13.8 mm, ZMFU no. 45857/Bv-7435; D, Pandora pulchella Yokoyama, 1926: Gangwon-do, site 10, shell length 30.1 mm, ZMFU no. 45802/Bv-7405; E and F, Pandora
wardiana Adams, 1860: Gangwon-do, site 10, shell length 28.5 mm, ZMFU no. 45803/Bv-7406.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 39

species were Glycymeris spp. (Mangsan and Chuam), Ruditapes
philippinarum (Adams et Reeve, 1850) [Anin (site 3) and Geumjin
(site 12)], Mytilus coruscus Gould, 1861, and Crassostrea gigas
(Thunberg, 1793) [near Samcheok (site 8)], and M. chinensis
(Mangsan). On biofouled buoys (Geumjin Port, site 11), live Arca
boucardi Jousseaume, 1894, Porterius dalli (Smith, 1885), and
M. galloprovincialis were dominant.
An euryhalineeestuarine assemblage observed near river
mouths and larger estuaries merits special attention. Living Nut-
tallia obscurata (Reeve, 1857) was very common intertidally in es-
tuaries of rivers flowing into the sea near Maenbang and Yeomjeon.
The latter locality was the only site where we collected live speci-
mens of Gari chinensis (Deshayes, 1855). Numerous shells of
Corbicula spp. were found throughout the entire studied area.
P. amurensis, found at Maenbang and Geumjin, is also an element of
this assemblage. N. obscurata has also been obtained further north
in Namdaechon estuary, Yangyang (Hong et al 2000;as“Nuttallia
olivacea”).
A cold-water and relatively cold-water (boreal) assemblage
(Modiolus kurilensis Bernard, 1983, Mytilus trossulus Gould, 1850,
Clinocardium likharevi (Kafanov in Scarlato, 1981), Megangulus
venulosus (Schrenck, 1861), Megangulus zyonoensis (Hatai et
Nisiyama, 1939), Macoma calcarea (Gmelin, 1791), Spisula sachali-
nensis (Schrenck, 1861), Protothaca adamsii (Reeve, 1863), Con-
chocele scarlatoi Ivanova et Moskaletz, 1984) is indicative of the
influence of cold currents on the shelf fauna of Gangwon, as well as
the location of this region in the northern area of South Korea.
Some species belonging to this group, e.g. M. calcarea and
P. adamsii, appeared at the edge of the South Korean shelf when the
sea level was much lower (120e130 meters), dated at around
Figure 24. A and B, Entodesma navicula (Adams et Reeve, 1850): Gangwon-do, site 11, shell length 49.5 mm, ZMFU no. 45810/Bv-7412; C and D, Siliqua pulchella (Dunker, 1852):
Gangwon-do, site 12, shell length 24.8 mm, ZMFU no. 45965/Bv-7475; E, Solen krusensterni Schrenck, 1867: Gangwon-do, site 10, shell length 89.6 mm, ZMFU no. 45783/Bv-7395.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e4440

Figure 25. Fossil and recent Conchocele from the northwestern Pacific: A, Conchocele disjuncta Gabb, 1866, Recent, Japan (after: Habe 1958, pl. 2, Figure 5); B, Conchocele scarlatoi
Ivanova et Moskaletz, 1984, recent northwestern East Sea (Sea of Japan), Possjet Bay, depth 30 m (after: Ivanova and Moskaletz 1984, Figure 1); C, Conchocele disjuncta var. alta
(Krishtofovich, 1936), southern Sakhalin, Maruyamskaya Formation, Upper Miocene (after: Krishtofovich 1964, pl. 53, Figure 5); D, Conchocele scarlatoi Ivanova et Moskaletz, 1984,
Recent, northwestern East Sea (Sea of Japan), Possjet Bay, subtidal (after: Lutaenko and Noseworthy 2012, pl. 25, Figure G).
Figure 26. Syntypes of Corbula amurensis Schrenck, 1861: Zoological Institute of the Russian Academy of Sciences (St. Petersburg), estuary of Amur River; A, shell length 11.9 mm; B,
shell length 13.9.
KA Lutaenko, RG Noseworthy / Journal of Asia-Pacific Biodiversity 12 (2019) 14e44 41

15000 years BP (Later Pleistocene) (Lee et al 2003). Many of them
live in Korea only in Gangwon and Gyeongbuk provinces.
Conflict of interest
The authors declare that there is no conflict of interest.
Acknowledgments
The authors would like to thank Joy Noseworthy for her help in
the logistics and management of the fieldwork on the coast of
Gangwon-do in May 2016. Irina E. Volvenko, Zoological Museum,
Far Eastern Federal University, Vladivostok, Russia, took all photo-
graphs and prepared plates. Henk H. Dijkstra, Naturalis Biodiversity
Center, Leiden, the Netherlands, kindly consulted us on Pectinidae
identification. Partial financial support for travel (K. Lutaenko) was
made possible through Professor Joong-Ki Park, Ewha Womans
University, Seoul, Republic of Korea, to whom the authors are very
grateful. This research did not receive any specific grant from
funding agencies in the public, commercial, or not-for-profit
sectors.
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