Зоологический журнал, 2024, № 4

Российская академия наук
ЗООЛОГИЧЕСКИЙ 
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том 103       № 4       2024       Апрель
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© Редколлегия “Зоологического журнала”
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СОДЕРЖАНИЕ
Том 103, Номер 4, 2024
The diversity of commercial marine bivalves in the Nha Phu Lagoon,  
Khanh Hoa Province, south-central Vietnam
N. T. Lan, P. Yu. Dgebuadze, P. T. Huan	
3
A new species of the wolf spider genus Mongolicosa (Araneae, Lycosidae) from Mongolia
S. L. Esyunin, A. L. Ustinova, B. Tsagaanbileg	
22
Первая находка Canthocamptus (Canthocamptus) microstaphylinus Wolf 1905  
(Harpacticoida, Copepoda) в прибрежно-соровой зоне озера Байкал
Т. М. Алексеева, Е. Б. Фефилова, Н. Г. Шевелева, О. А. Тимошкин	
26
Восточно-азиатский вид Thermocyclops taihokuensis (Harada 1931)  
(Crustacea, Copepoda, Cyclopoida) в Таганрогском заливе Азовского моря
В. И. Лазарева, Р. З. Сабитова	
36
Генетические линии Parisotoma notabilis sensu lato (Hexapoda, Collembola)  
и их использование в биологическом мониторинге
А. В. Стрючкова, Н. А. Кузнецова	
44
Долгосрочная память трипсов Thrips tabaci (Thysanoptera, Thripidae)
М. А. Федорова, С. Э. Фарисенков, А. А. Полилов	
53
Паразитоиды (Hymenoptera, Eulophidae, Braconidae) как фактор смертности  
липовой моли-пестрянки (Phyllonorycter issikii, Lepidoptera, Gracillariidae) в Предуралье
И. В. Ермолаев, З. А. Ефремова, С. А. Белокобыльский, В. В. Курбатов, Е. Н. Егоренкова	
58
Первый опыт мечения GPS-GSM трансмиттерами ушастой совы (Asio otus, Strigidae, Aves) 
в европейской части России
А. В. Шариков, С. В. Волков, А. С. Педенко, Т. С. Массальская, А. В. Макаров	
66
Через крах к возрождению: нестационарная динамика сообщества грызунов  
в новый цикл опустынивания в Калмыкии
Е. Н. Суркова, Л. Е. Савинецкая, А. В. Чабовский	
70
Наследование признаков родительских видов гибридами большого,  
желтого и крапчатого сусликов  (Spermophilus, Sciuridae, Rodentia)
С. В. Титов, О. В. Чернышова, М. Д. Симаков, Н. А. Картавов, А. А. Кузьмин	
80
Методика зоологических исследований
Использование съемки с беспилотных летательных аппаратов в исследованиях  
поведения млекопитающих
Е. А. Березина, А. Н. Гилёв, К. А. Каренина	
100

Contents
Volume 103, No 4, 2024
The diversity of commercial marine bivalves in the Nha Phu Lagoon,  
Khanh Hoa Province, south-central Vietnam
N. T. Lan, P. Yu. Dgebuadze, P. T. Huan	
3
A new species of the wolf spider genus Mongolicosa (Araneae, Lycosidae) from Mongolia
S. L. Esyunin, A. L. Ustinova, B. Tsagaanbileg	
22
First record of Canthocamptus (Canthocamptus) microstaphylinus Wolf 1905  
(Harpacticoida, Copepoda) from a shallow coastal area of Lake Baikal
T. M. Alekseeva, E. B. Fefilova, N. G. Sheveleva, O. A. Timoshkin	
26
The East Asian species, Thermocyclops taihokuensis (Harada 1931)  
(Crustacea, Copepoda, Cyclopoida) in the Taganrog Bay of Azov Sea
V. I. Lazareva, R. Z. Sabitova	
36
Genetic lineages of Parisotoma notabilis sensu lato (Hexapoda, Collembola) and their use  
in biological monitoring
A. V. Striuchkova, N. A. Kuznetsova	
44
Long-term memory in Thrips tabaci (Thysanoptera, Thripidae)
M. A. Fedorova, S. E. Farisenkov, A. A. Polilov	
53
Parasitoids (Hymenoptera, Eulophidae, Braconidae) as a mortality factor for the lime leaf miner  
(Phyllonorycter issikii, Lepidoptera, Gracillariidae) in the Cis-Urals
I. V. Ermolaev, Z. A. Yefremova, S. A. Belokobylskij, V. V. Kurbatov, E. N. Yegorenkova	
58
First observations of the movements of the Long-eared owl (Asio otus, Strigidae, Aves)  
using GPS-GSM telemetry in the European part of Russia
A. V. Sharikov, S. V. Volkov, A. S. Pedenko, T. S. Massalskaya, A. V. Makarov	
66
Through a collapse to restoration: the non-stationary dynamics of the rodent community  
in a new cycle of desertification in Kaфlmykia
E. N. Surkova, L. E. Savinetskaya, A. V. Tchabovsky	
70
Inheritance of signs of parent species by hybrids of Russet, Yellow and Speckled ground squirrels  
(Spermophilus, Sciuridae, Rodentia)
S. V. Titov, O. V. Chernyshova, M. D. Simakov, N. A. Kartavov, A. A. Kuzmin	
80
Methods of Zoological Investigations
The use of drones for studying the behaviour of mammals
E. A. Berezina, A. N. Giljov, K. A. Karenina	
100

ЗООЛОГИЧЕСКИЙ ЖУРНАЛ,  2024, том 103, № 4,  с.  3–21
УДК 594.124; 594.121; 594.133; 594.113
THE DIVERSITY OF COMMERCIAL MARINE BIVALVES  
IN THE NHA PHU LAGOON, KHANH HOA PROVINCE,  
SOUTH-CENTRAL VIETNAM
© 2024   N. T. Lana,*, P. Yu. Dgebuadzeb, P. T. Huana
aCoastal Branch of the Joint Vietnam-Russia Tropical Science and Technology Research Center,  
Nha Trang, Khanh Hoa, 57000 Vietnam
bA. N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, 119071 Russia
*e-mail: nguyenlanst1805@gmail.com
Received October 13, 2023

Accepted April 19, 2024
The study was conducted as part of the Joint Vietnam–Russia Tropical Science and Technology Research 
 project on indicator organisms in the marine environment of south-central Vietnam. 
The research focused on coastal areas with significant anthropogenic impacts, revealing that biological 

and subject to both human and natural factors. The material from the soft-bottom zone was collected 
using a grab rake with a mesh bag at the end. Scuba diving equipment was used in areas with a hard substrate and mudflats. In addition, during sample collection, several mollusc specimens were acquired from 
fishermen who cought marine organisms in that lagoon. Comparing the results with previous studies on 
bivalve resources in the area, the Nha Phu Lagoon was found to harbour a wide range of economically 

the family Veneridae having the highest number of species (eight). The most abundant catches were from 
the families Ostreidae and Mytilidae, primarily consisting of 
 (Thunberg 1793) and Perna 

 (Linnaeus 1758), both cultured in the lagoon. Apart from their economic value, these two species 
also play crucial roles in the ecosystem by filtering water and retaining organic debris in the environment.
Keywords: sea molluscs, economic value, ecosystem roles
DOI: 10.31857/S0044513424040016, EDN: 
  
There are approximately 10,000 known living species 
Currently, the fisheries and aquaculture industry is 

-
creasing global demand for food. According to statistics 
from the Food and Agriculture Organization (FAO), 
the total production of fisheries and aquaculture in 
2020 reached 214 million tons (FAO, 2020), including 
178 million tons of animals and 36 million tons of seaweed. The global production of marine bivalves exceeds 
15 million tons per year, with aquaculture accounting 
for 89% and wild catches 11% (Wijsman et al., 2019). 
Asian countries dominate the production of bivalves, 
contributing to more than 85% of the global production, and this is expected to further increase to ensure 
food security (Steeves et al., 2018).
However, under economic pressures, fisheries in 
general and for bivalves in particular have significantly 
increased. The consequences of overfishing and other 
forms of exploitation (Hiscock, 2014; Costello et al., 
2021), destruction of mangroves, loss of coral reefs and 
seaweed beds, changes in wetland, water, estuarine, 
of bivalve molluscs worldwide; they are the second largest taxonomic class within the Mollusca (Bieler et al., 

including those found in estuarine and brackish water 
environments (Huber, 2015). They are widely distribu- 
ted, highly adaptable to various habitats and have diverse life habits, such as burrowing in sand and mud, 
clinging to shells and rocks, burrowing into wood and 
rock corals, crawling through seaweed thickets (Turgeon 
et al., 2009) and even clinging to mangrove leaves in 
mangrove forests. Due to their diverse species composition, extensive distribution and consumption of organic 
matter, they play a crucial role in marine ecosystems 
and contribute to ecosystem metabolism (Weitzman et 
al., 2019; Smyth et al., 2017; Gosling, 2003; National 
Research Council, 2000). In addition to their ecological 

that helps ensure global food security (Martins et al., 

3

LAN et al.
Nha Phu Lagoon is the main agricultural area in 
sandbar and mudflat landscapes (Lotze et al., 2010; 
Fredston-Hermann et al., 2013), pollution and marine debris (Pitacco et al., 2018) are causing biodiversity loss, including bivalves (Halpern et al., 2015; Boyce 
 
et al., 2020). 
Vietnam is one of the world’s leading seafood exKhanh Hoa Province, providing seafood for the pro- 
vince’s residents, especially serving tourists in Nha 
Trang City. The main forms of fishing in this area are 
fishing by motorized or non-motorized boats, scuba 
diving equipment was used in areas with hard substrate 
and manual collection on tidal flats. Around the lagoon, there are shrimp ponds, fish and molluscs raised 
in cages on the lagoon water surface. Considering the 
increasing demand for seafood consumption and export alongside economic and life development, there is 
a need to promote resource research and evaluation activities. This study focuses on investigating the composition of economically valuable bivalve molluscs in Nha 
Phu Lagoon (Khanh Hoa) to assess their resources and 
provide a scientific basis for management, monitoring 
and exploitation.
MATERIAL AND METHODS
porters and plays a crucial role in the global seafood 
supply. According to the Vietnam Seafood Exporters 
and Producers Association (Ministry of agriculture 
and rural development, 2021), marine molluscs are the 
second-largest biological resource after marine fish, 
with an export value of 141,000 million USD in 2021. 
Favourable natural conditions contribute to the development of exploitation and aquaculture in Vietnam. 
Firstly, Vietnam is located in an area of global marine 
biodiversity. Studies on species richness of bivalves in 
Southeast Asia have identified approximately 1200– 
1500 species (Adrianov, Lutaenko, 2016), including 
594 species in the Gulf of Thailand, 368 species in 
the Gulf of Tonkin, 299 species in Hong Kong and 
 
Research area
Nha Phu Lagoon (Fig. 1) is situated to the north 
815 species in Vietnam (Hylleberg, Kilburn, 2003). Do 
and Le (Do, Le, 2015) reported that Vietnam’s marine 
waters contain approximately 2200–2500 species of 
molluscs, with more than 1000 species belonging to the 
class Bivalvia.
Secondly, Vietnam’s long curving coastline forms 
of Nha Trang City, in Khanh Hoa Province. Spanning 
an approximate area of 4,500 hectares (Strehlow, Peter, 
2004), this lagoon is protected by the Hon Ho Mountain range and is accompanied by small islands such as 
Hon Hoai, Hon Sam, Hon Thi and Hon Khi Island. It 
is recognized as one of the prominent sites for development and aquaculture in Khanh Hoa Province. Shrimp 
ponds can be found surrounding the lagoon, while cages 
on the water surface are primarily used for farming fish 
and molluscs.
Samples collection
The material was collected six times between 2019 
semi-enclosed bays stretching from north to south, including Ha Long Bay (Quang Ninh), Tam Giang Lagoon (Thua Tien Hue), Thi Nai Lagoon (Binh Dinh), 
Vung Ro Bay and Xuan Dai Bay (Phu Yen), Van Phong 
Bay, Nha Phu Lagoon, Thuy Trieu Lagoon (Khanh 
Hoa) and Nai Lagoon (Ninh Thuan), which are key 
areas for aquaculture and exploitation. However, unplanned aquaculture, a lack of control over space and 
density and direct dumping have led to environmental degradation in these areas. Pham, Nguyen (2006), 
Pham et al. (2010), Nguyen et al. (2009), Nguyen et al. 
(2010), Nguyen et al (2016) and Vo et al. (2018) have 
shown that localized pollution occurs in most marine 
areas along the coast.
and 2022. A grab rake with a mesh bag at the end 
was used to sample from the soft-bottom zone (depth 
3–20  m). The sample was then washed through a 
sieve with a mesh diameter of 1 cm. Scuba diving 
Fig. 1. The study region. Nha Phu Lagoon.
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THE DIVERSITY OF COMMERCIAL MARINE BIVALVES IN THE NHA PHU LAGOON 
5
	
THE DIVERSITY OF COMMERCIAL MARINE BIVALVES IN THE NHA PHU LAGOON 
5
Identification of bivalves was carried out using a 
equipment was used in areas with hard substrate and 
mudflats (depth 6–25 m). Three 1 m2 frames were 
randomly placed underwater at each sample collection site. All detected bivalve specimens were collected from the frame area and transported to the laboratory for further identification. In addition, during the 
sample collection, several mollusc specimens were 
acquired from fishermen who catch marine organisms in this lagoon. Information on the location and 
type of the bottom substrate at the fishing spot was 
 
obtained.
comparative-morphological method based on morphological criteria of external and internal body structure 
following the works of Abbott and Dance (1982), Carpenter and Niem (1998), Hylleberg and Kilburn (2003), 
Nguyen N. T. (2005), Do and Le (2015), Adrianov and 
Lutaenko (2016). In this study, the classifications and 
accepted names of Bivalvia taxa are used as given in 
the WoRMS database (WoRMS, 2023). Higher taxa 
and species in frame of families are presented in alpha- 
betical order.
To obtain information on the biological resources 
RESULTS
A total of 27 commercial marine bivalve species were 
of the lagoon, our group conducted a survey in small 
markets around Nha Phu Lagoon (e.g., Vinh Luong, 
Ninh Ich, Ninh Hoa markets) to find molluscs that 
are actively caught by fishermen and determine their 
 
fishing season.
identified in Nha Phu lagoon, Khanh Hoa Province, 
Vietnam (Table 1).
Table 1. List of species, habitat and some commercial information
Habitat, commercial information
№
Species
Soft-bottom 
Hard substrate 
Acquired from 
Actually 
zone
and mudflats
fishermen
sold
1
Solen vagina Linnaeus 1758
+
+
+
2
Vasticardium pectiniforme (Born 1780)
+
+
+
3
Asaphis violascens (Forsskål 1775)
+
+
+
+
4
Lutraria rhynchaena Jonas 1844
+
+
5
Anomalodiscus squamosus (Linnaeus 1758)
+
+
6
Gafrarium pectinatum (Linnaeus 1758)
+
+
+
+
7
Marcia hiantina (Lamarck 1818)
+
+
8
Meretrix lyrata (G. B. Sowerby II 1851)
+
+
+
9
Meretrix meretrix (Linnaeus 1758)
+
+
+
10
Paphia euglypta (R. A. Philippi 1847)
+
+
+
11
Paratapes undulatus (Born 1778)
+
+
+
12
Periglypta exclathrata (Sacco 1900)
+
+
+
13
Anadara antiquata (Linnaeus 1758)
+
+
+
+
14
Tegillarca granosa (Linnaeus 1758)
+
+
+
15
Trisidos semitorta (Lamarck 1819)
+
+
+
16
Perna viridis (Linnaeus 1758)
+
+
+
17
Isognomon isognomum (Linnaeus 1758)
+
+
18
Pinctada imbricata Röding 1798
+
+
19
Pinctada margaritifera (Linnaeus 1758)
+
+
+
20
Magallana gigas (Thunberg 1793)
+
+
+
21
Atrina pectinata (Linnaeus 1767)
+
+
+
22
Atrina vexillum (Born 1778)
+
+
+
23
Amusium pleuronectes (Linnaeus 1758)
+
+
+
+
24
Annachlamys striatula (Linnaeus 1758)
+
+
+
+
25
Decatopecten plica (Linnaeus 1758)
+
+
+
26
Mimachlamys sanguinea (Linnaeus 1758)
+
+
+
+
27
Spondylus squamosus Schreibers 1793
+
+
+
ЗООЛОГИЧЕСКИЙ ЖУРНАЛ
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LAN et al.
Systematic account
D i s t r i b u t i o n  i n  V i e t n a m .  Along the coast 
Infraclass Heteroconchia J. E. Gray 1854 
Order Adapedonta Cossmann & Peyrot 1909 
Family Solenidae Lamarck 1809 
of the Gulf of Tonkin, along the coast of Nha Trang 
(Do, Le, 2015), in the Khanh Hoa, Can Gio, Ha Tinh, 
North Vietnam, etc. areas. Habitat: sandy bottom at 
1–50 m (Nguyen, 2005).
Genus Solen Linnaeus 1758
In Nha Phu Lagoon – occurs on sandy, sandy-silt 
bottoms.
Order Cardiida A. Férussac 1822 
Family Cardiidae Lamarck 1809 
1. Solen vagina Linnaeus 1758
(Fig. 2A)
Solen vagina Linnaeus 1758 (Linnaeus, 1758)
Solen vagina: Abbott and Dance, 1982: p. 338; CarGenus Vasticardium Iredale 1927
penter and Niem, 1998: p. 289, p. 285 fig. 6; Hylleberg 
and Kilburn, 2003: p. 194; plate 6 fig. 4; Nguyen, 2005: 
p. 280, plate 86 fig. 2.
D e s c r i p t i o n .  Shell medium sized, reaching 
2. Vasticardium pectiniforme (Born 1780)
(Fig. 2C)
Cardium pectiniforme Born 1780 (Born, 1780)
Vasticardium pectiniforme: Abbott and Dance, 1982: 
8–14 cm in length. Sculpture consists or numerous fine 
concentric lines and growth pauses. Colour externally 
purplish (under the greenish periostracum) and internally whitish.
p. 328; Carpenter and Niem, 1998: p. 256, p. 252 
fig. 13; Hylleberg and Kilburn, 2003: p. 185, plate 4 
fig. 15 a, b, c; Adrianov and Lutaenko, 2016: plate 13 
 
Season of capture fisheries – from May to November.
figs C, D.
A
B
C
D
Fig. 2. A – Solen vagina (93 mm), B – Asaphis violascens (49 mm), C – Vasticardium pectiniforme (32 mm),  D – Lutraria 
rhynchaena (64 mm).
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THE DIVERSITY OF COMMERCIAL MARINE BIVALVES IN THE NHA PHU LAGOON 
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D e s c r i p t i o n .  Shell medium sized, reaching 
Lutraria rhynchaena: Abbott and Dance, 1982: 
p. 336; Hylleberg and Kilburn. 2003: p. 187; Nguyen, 
2005: p. 279, plate 86 fig. 1; Adrianov and Lutaenko, 
2016: plate 14 figs G, H.
D e s c r i p t i o n .  Shells medium-sized (4–8 cm 
4–6 cm in height. Shell moderately thick, oblong-ovate 
and higher than long, slightly inequilateral with posterior part tending to be slightly expanded. Thirty promi- 
nent, rounded and rugose radial ribs at each valve. 
Outside of shell whitish, periostracum straw colour to 
 
olive-brown, sometimes with a scattering of black spots. 
Interior completely white, or more or less stained yellow.
in length), elongated oval.  The shell is fairly thick. 
Outer sculpture fine concentric lines corresponding to 
periods of growth. The basic colour is white and the 
 
Season of capture fisheries – from May to Sep- 
tember.
periostracum is brown, but the latter is usually abraded. 
The interior surface is glossy white. 
D i s t r i b u t i o n  i n  V i e t n a m .  Usually found 
Season of capture fisheries – exploited year-round, 
in coastal waters (Do, Le, 2015).
concentrated from May to November.
In Nha Phu Lagoon – found on soft bottoms near 
D i s t r i b u t i o n  i n  V i e t n a m .  Occurs in tidal 
the estuary.
areas in the Khanh Hoa Province. Habitat: sandy bottom at 1–15 m (Nguyen, 2005; Dang et al., 2009).
In Nha Phu Lagoon – occurs in the coastal alluvial 
FamilyPsammobiidae J. Fleming 1828 
zone (between low and high tide).
Genus Asaphis Modeer 1793
Family Veneridae Rafinesque 1815 
Genus Anomalodiscus Dall 1902
3. Asaphis violascens (Forsskål 1775)
(Fig. 2B) 
Venus violascens Forsskål 1775 (Niebuhr, 1775)
Asaphis violascens: Abbott and Dance, 1982: p. 347; 
5. Anomalodiscus squamosus (Linnaeus 1758)
(Fig. 3A) 
Venus squamosa Linnaeus 1758 (Linnaeus, 1758)
Anomalodiscus squamosus: Carpenter and Niem, 
Carpenter and Niem, 1998: p. 307, p. 305 fig. 1; Okutani, 2000: p. 987, plate 491 fig. 15; Hylleberg and Kilburn, 2003: p. 204; Nguyen, 2005: p. 285, plate 87, 
figs  8, 10; Adrianov and Lutaenko, 2016: plate 17 
 
figs A, B.  
D e s c r i p t i o n .  Shell length commonly to 4– 
1998: p. 328, p. 324 fig. 24; Okutani, 2000: p. 1003, 
plate 499 fig. 7; Hylleberg and Kilburn, 2003: p. 209; 
Do and Le, 2015: p. 162 fig. 113; Adrianov and Lutaenko, 2016: plate 18 figs E, F.
D e s c r i p t i o n .  Shell ranges 2–4 cm in length. 
Shell small, solid, inflated and strongly inequilateral 
in shape, subtrigonal in outline. Anterior side of shell 
rounded, posterior side produced and tapering to posterior end. Outer sculpture of valves strong, of dense radial riblets and low concentric cords, giving the surface 
a finely granulated and latticed aspect. Outside of shell 
dull yellowish white, often flushed with slightly darker 
 
greyish to purplish brown on lunule and escutcheon 
 
6 cm. Shell solid, rather large, inflated, slightly gaping 
posteriorly, elongate-ovate in outline but quite variable 
in shape and sculpture. Outer sculpture of numerous, 
strong, rounded and often forked radial ribs, developed 
throughout the valves and crossed by concentric lines 
that do not interrupt the ribs but make them slightly 
scaly or nodulose, at least on posterior slope. Outside 
of shell dull coloured, usually creamy white with a pale 
greyish brown periostracum. Interior slightly glossy, 
whitish, often with yellow or orange hue centrally and a 
dark purplish blotch posteriorly. 
areas. Interior whitish. 
Season of capture fisheries – from May to Sep- 
Season of capture fisheries – from May to No- 
tember.
vember.
D i s t r i b u t i o n  i n  V i e t n a m .  Commonly 
D i s t r i b u t i o n  i n  V i e t n a m .  Found only in 
found in coastal waters (Do, Le, 2015), in the Khanh 
Hoa, Southwest Vietnam, North Vietnam, etc. areas. 
Habitat: coarse sand or gravels near rocks at 1–15 m 
(Nguyen, 2005).
the southern Vietnam Sea (Dang et al., 2009) in the 
Khanh Hoa, Hoi An, Southwest Vietnam, North Vietnam area along the coast and around islands (Do, Le, 
2015; Nguyen, 2005).
In Nha Phu Lagoon – occurs in the muddy zone 
In Nha Phu Lagoon – found in sand, near mangrove forests.
(place between low and high tide) and hard bottom 
zone.
Genus Gafrarium Röding 1798
Order Venerida Gray 1854 
Family Mactridae Lamarck 1809 
Genus Lutraria Lamarck 1799
6. Gafrarium pectinatum (Linnaeus 1758)
(Fig. 3C)
Venus pectinata Linnaeus 1758 (Linnaeus, 1758)
Gafrarium pectinatum: Carpenter and Niem, 1998: 
4. Lutraria rhynchaena Jonas 1844
(Fig. 2D)
Lutraria rhynchaena Jonas 1844 (Jonas, 1844)
p. 332, p. 322 fig. 9; Okutani, 2000: p.1007, plate 501 fig. 24; 
 
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LAN et al.
Hoa, Ca Na, Hue, Southwest Vietnam, Quang Ninh, 
Hai Phong. Habitat: sand at 5–25 m (Nguyen, 2005). 
Hylleberg and Kilburn, 2003: p. 213; Nguyen, 2005: 
p. 294, plate 90 fig. 24; Do and Le, 2015: p. 114–115, 
fig. 77; Adrianov and Lutaenko, 2016: plate 20 figs C, D. 
In Nha Phu Lagoon – widespread, occurring at 
D e s c r i p t i o n .  Shell length 2–4 cm. Shell thick 
most sampling sites.
Genus Marcia H. Adams & A. Adams 1857
7. Marcia hiantina (Lamarck 1818)
(Fig. 3B) 
Venus hiantina Lamarck 1818 (Lamarck, 1818)
Marcia hiantina: Carpenter and Niem, 1998: p. 334; 
Hylleberg and Kilburn, 2003: p. 215; Nguyen, 2005: 
p. 299, plate 89 fig. 15; Adrianov and Lutaenko, 2016: 
plate 18, figs A, B.
and solid, with a variable, relatively compressed, elongated shape, elliptical-ovate in outline. Outer sculpture 
rather fine but well-marked, with many small concentric ridges and slightly nodulous, diverging radial riblets rapidly weakening medially and anteriorly. Main 
sculpture of median area of valves of concentric ridges. Outside coloration of shell variable, off-white to 
buff-coloured and generally with fawn or brown blotches or spots throughout. Interior porcelaneous white, often with yellowish hue inside the pallial line, and with 
purple-brown blotching on lunular margin, posteroventral part of hinge and posterior margin of valves. 
D e s c r i p t i o n .  Maximum shell length 5 cm. 
Season of capture fisheries – from April to No- 
vember.
D i s t r i b u t i o n  i n  V i e t n a m .  Found along 
Shell solid, with a variable, moderately inflated, inequilateral shape, roughly rounded-ovate in outline. 
Outer surface of valves covered with numerous, somewhat irregular concentric grooves and cords, the latter 
the shore to a depth of 20m (Do, Le, 2015) in Khanh 
A
B
C
D
Fig. 3.  A – Anomalodiscus squamosus (29 mm), B – Marcia hiantina (37 mm), C – Gafrarium pectinatum (29 mm),  
D – Periglypta exclathrata (58 mm).
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THE DIVERSITY OF COMMERCIAL MARINE BIVALVES IN THE NHA PHU LAGOON 
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p. 168-169, fig. 118; Adrianov and Lutaenko, 2016: 
plate 20 figs A, B.
D e s c r i p t i o n .  Shell length 3–5 cm. Shell thick, 
sometimes flatter and less apparent, but always present, near posterior half of ventral margin. outside of 
shell variable in colour and pattern, whitish to grey, 
fawn-coloured or brown, frequently with various patterns of white, fawn or purplish radial bands, zigzags or 
trigonal spots or irregular blotches. Interior dirty white, 
often dark coloured on hinge area.
Season of capture fisheries – from March to November.
moderately inflated with a variable, inequilateral shape, 
subtrigonal in outline. Outer surface of shell with incised concentric grooves fading out toward anterodorsal 
and posterodorsal margins, and sometimes also at ventral margin of larger specimens. Outside of shell light 
fawn to brown under the translucent, glossy, yellowish 
periostracum. Interior porcelaneous white.
D i s t r i b u t i o n  i n  V i e t n a m .  Found in 
Season of capture fisheries – exploited year-round.
D i s t r i b u t i o n  i n  V i e t n a m .  Found along 
Khanh Hoa, Long Hai, Ca Na, Northern Vietnam, etc. 
Habitat: sand at 1–15 m (Nguyen, 2005).
In Nha Phu Lagoon – found alive buried in sand in 
coastal areas.
Genus Meretrix Lamarck 1799
the coastal region of South Vietnam and can now be 
found in Central and North Vietnam (Do, Le, 2015). 
According to Nguyen, 2005: Ben Tre, Soc Trang, Bac 
Lieu, Go Cong, Tien Giang, Tra Vinh, Ca Mau, Rach 
Gia, Khanh Hoa. Habitat: sandy mud at 1–15 m.
In Nha Phu Lagoon – occurs on the sandy bottom 
along the shore.
8. Meretrix lyrata (G. B. Sowerby II 1851)
(Fig. 4B) 
Cytheraea lyrata G. B. Sowerby II 1851 (Sowerby II, 
1851)
Meretrix lyrata: Carpenter and Niem, 1998: p. 336, 
9. Meretrix meretrix (Linnaeus 1758)
(Fig. 4A) 
Venus meretrix Linnaeus 1758 (Linnaeus, 1758)
Meretrix meretrix: Carpenter and Niem, 1998: p. 37, 
p. 323 fig. 18; Hylleberg and Kilburn, 2003: p. 216; 
Nguyen, 2005: p. 295, plate 89 fig. 6; Do and Le, 2015: 
p. 323 fig. 17; Hylleberg and Kilburn, 2003: p. 216; 
A
B
C
D
Fig. 4.  A – Meretrix meretrix (47 mm), B – Meretrix lyrata (32 mm), C – Paratapes undulatus (73 mm), D – Paphia euglypta 
(56 mm).
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