Freshwater Mussels (Bivalvia: Unionida) of Vietnam:
Diversity, Distribution, and Conservation Status
Authors: Do, Van Tu, Tuan, Le Quang, and Bogan, Arthur E.
Source: Freshwater Mollusk Biology and Conservation, 21(1) : 1-18
Published By: Freshwater Mollusk Conservation Society
URL: https://doi.org/10.31931/fmbc.v21i1.2018.1-18
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Freshwater Mollusk Biology and Conservation 21:1–18, 2018
Ó Freshwater Mollusk Conservation Society 2018
REGULAR ARTICLE
FRESHWATER MUSSELS (BIVALVIA: UNIONIDA) OF
VIETNAM: DIVERSITY, DISTRIBUTION, AND
CONSERVATION STATUS
Van Tu Do1, Le Quang Tuan1, and Arthur E. Bogan2*
1
Institute of Ecology and Biological Resources (IEBR), Vietnam Academy of Science and
Technology, 18 Hoang Quoc Viet, Nghia Do, Cau Giay, Ha Noi, Vietnam, dovantu.iebr@gmail.com;
penguin0343@gmail.com
2
North Carolina Museum of Natural Sciences, 11 West Jones Street, Raleigh, NC 27601 USA
ABSTRACT
Vietnam has the second highest diversity of freshwater mussels (Unionida) in Asia after China. The
purpose of this paper is to compile an up-to-date list of the modern unionid fauna of Vietnam and its
current conservation status. Unfortunately, there has been relatively little research on this fauna in
Vietnam. Fifty-nine species of Unionida have been recorded from Vietnam based on literature, museum
records, and our fieldwork. Fifty were assessed in the International Union for Conservation of Nature
(IUCN) Red List 2016 in the IUCN categories of Critically Endangered (four species, 6.8%),
Endangered (seven species, 12%), Vulnerable (one species, 1.7%), Near Threatened (two species, 3.4%),
Least Concern (23 species, 39%), Data Deficient (11 species, 18.6%), and Not Evaluated (11 species,
18.6%). Considering the impacts of pollution, timbering, agriculture, and damming of rivers, research
on the diversity and conservation status of freshwater mussels is very urgently needed to propose
specific conservation measures for these species in Vietnam. If all taxa listed as Data Deficient are found
to be threatened, with around 42% of species threatened, this fauna would be one of the most
threatened freshwater molluscan faunas in Asia.
KEY WORDS: Unionidae, Margaritiferidae, IUCN Red List, extinct, endangered, citizen science
INTRODUCTION
Freshwater bivalves are widely distributed in the freshwaters
of the world and are considered one of the most imperiled
animal groups (Bogan 1993, 2008; Lydeard et al. 2004; Strayer
and Dudgeon 2010; Graf 2013; Haag and Williams 2014). Two
major diversity hotspots of unionid bivalves are the southeastern
United States (Neves et al. 1997; Bogan 2008; Haag 2012) and
east and southeastern Asia (Bolotov et al. 2017; Zieritz et al.
2017). Our understanding of the distribution of unionids in Asia
is limited and the coverage of information is uneven and mostly
historical (Dudgeon et al. 2006; Zieritz et al. 2017).
Overviews of the freshwater bivalve fauna across Asia,
including Vietnam, were provided by Haas (1910a, 1910b,
1910c–1920, 1923). Compendia covering the Unionida of the
*Corresponding Author: arthur.bogan@naturalsciences.org
world began with the Synopsis by Lea (1836, 1838, 1852,
1870) and were followed and expanded by Simpson (1900,
1914) and Modell (1942, 1949, 1964). Haas (1969a) wrote the
last comprehensive survey of the Unionida, and followed it
with a treatise (Haas 1969b) that covered both modern and
fossil taxa to the generic level. Higher classification of the
Unionida was overinflated by Starobogatov (1970). Brandt
(1974) summarized the Thai and Southeast Asian unionids,
but overlooked Haas (1969a, 1969b). Ða. ng et al. (1980)
followed the classification proposed by Starobogatov (1970).
The inflated taxonomy of the Unionidae erected by Starobogatov’s classification was reduced by Graf (2007). A
preliminary list of freshwater bivalve taxa from Vietnam from
literature and museum data was included in the summary of
the East and Southeast Asia freshwater bivalve fauna by
Zieritz et al. (2017). Analyzing molecular data, the Unionidae
has been confirmed as a monophyletic clade (Hoeh et al. 1998,
1
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DO ET AL.
2
Table 1. Unionid taxa listed and/or figured by in the four volumes by Thach (2005, 2007, 2012, 2016b).
Species
Anodonta harlandi Baird & Adams, 18671
Cristaria bialata (Lea, 1829)2
Cristaria discoidea3
Cristaria truncata
Hyriopsis cumingii4
Hyriopsis schlegeli5
Hyriopsis sp.
Lamprotula leai
Lanceolaria bilirata
Lanceolaria bogani
Lanceolaria fruhstorferi
Lanceolaria grayi
Lanceolaria grayana (Lea, 1834)
Lanceolaria laevis
Lanceolaria yueyingae He & Zhuang, 2013
Nodularia douglasiae
Oxynaia jourdyi
Oxynaia micheloti
Physunio inornatus
Physunio micropterus
Physunio modelli
Pilsbryoconcha exilis
Pilsbryoconcha lemeslei
Pseudodon cambodjensis
Pseudodon contradens tumidula (Lea, 1856)6
Pseudodon mouhoti [sic]7
Pseudodon vondembuschianus chaperi (Morgan, 1885)
Sinanodonta elliptica
Sinanodonta hunganhi
Sinanodonta woodiana
Trapezoideus exolescens
Uniandra contradens tumidula (Lea, 1856)8
2005
2007
2012
2016b
Fig.
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1
Anodonta harlandi is a junior synonym of Sinanodonta woodiana.
2
Cristaria bialata is a junior synonym of C. plicata.
3
Cristaria discoidea is a junior synonym of Pletholophus tenuis.
4
Hyriopsis cumingii moved to Sinohyriopsis cumingii (see Lopes-Lima et al. 2017).
5
Hyriopsis schlegeli is misidentification of Sinohyriopsis cumingii.
6
Misplaced species, belongs in Contradens.
7
Incorrect spelling of Pseudodon mouhotii
8
Correct genus is Contradens.
2001, 2002; Roe and Hoeh 2003; Graf and Cummings 2006;
Breton et al. 2007, 2010; Doucet-Beaupré et al. 2010; Whelan
et al. 2011; Pfeiffer and Graf 2015). The most recent
phylogenetic classification within the modern Unionidae is
that of Lopes-Lima et al. (2017) and Bolotov et al. (2017).
Early work on the Vietnamese molluscan fauna was
typically performed by European malacologists describing
freshwater bivalve species. These included the papers of
Morelet (1865, 1866), Mabille (1887), Morlet (1886a, 1886b,
1891), Dautzenberg (1900), Bavay and Dautzenberg (1901),
Martens (1902), Rochebrune (1904a, 1904b), Rolle (1904),
Dautzenberg and Fischer 1906a, 1906b, 1908), and Haas
(1910a, 1910b, 1910c-1920, 1913, 1923). Isaac Lea, living in
the United States, also described some unionids from
Southeast Asia (see Scudder 1885). The only comprehensive
treatment of the freshwater invertebrates, including freshwater
bivalves for northern Vietnam, is that by Ða. ng et al. (1980).
Thach, a retired physician and avocational shell collector
has published four volumes providing information and
pictures of marine, terrestrial, and freshwater mollusks of
Vietnam, including some Unionidae (Thach 2005, 2007, 2012,
2016b) (Table 1). A total of 21 species of freshwater mussels
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FRESHWATER MUSSELS OF VIETNAM
from Vietnam were reported and figured in the 4 volumes by
Thach. He also described two new species, Lanceolaria
bogani Thach, 2016 (Thach 2016a) and Sinanodonta hunganhi Thach, 2016 (Thach 2016b). Thach divided the freshwater
bivalves of Vietnam into Amblemidae and Unionidae, but did
not comment on the Margaritiferidae. His report and figure of
Pilsbryoconcha lemeslei is the first record of this species from
Vietnam (Thach 2007).
We recently have been surveying the freshwater mussel
fauna of Vietnam (Bogan and Do 2011, 2013a, 2013b, 2014a,
2014b, 2016). The objective of the present paper is to develop
an up-to-date list of the unionid fauna reported for Vietnam
and their distribution (Table 2; see also Supplemental Species
Range Maps). The modern unionid fauna of Vietnam is
divided between two families, Margaritiferidae and Unionidae.
Margaritiferidae is represented by a single species. The
Unionidae fauna of Vietnam is represented by 28 genera and
58 species (Table 2).
METHODS
Study Area
Vietnam lies on the eastern side of the Indochina Peninsula
encompassing 331,210 km2 (Fig. 1). The northern part of the
country is mountainous and contains the Red River basin,
which drains to the east, emptying into the Gulf of Tonkin.
The rivers draining to the west are tributaries of the Mekong
River basin, draining into the Gulf of Thailand. Southern
Vietnam is home to the Annamite Range, the Central or
Western Highlands and the extensive Mekong River delta and
its numerous distributaries. The major rivers of Vietnam are
shown on Figure 1.
Historical literature was used to document species
described from Vietnam and was compared with the list
published by Ða. ng et al. (1980). Taxonomy of these species
was challenging because Ða. ng et al. (1980) based their
taxonomy on Starobogatov (1970), but see Graf’s (2007)
revision of Starobogatov’s taxonomy. Taxonomy used in this
paper is based on available literature, as presented by LopesLima et al. (2017) and is the same as used by Zieritz et al.
(2017). Additional information was collected from photographs of specimens from various museum collections
available on the Internet, which were checked and identifications verified (Graf and Cummings 2017). Dates of publication
were verified and resulted in minor changes (Bogan and Do
2011; Bogan 2015).
Our fieldwork was carried out at the following times: July
2010 in northern Vietnam; November 2012 across 12 northern
provinces of Vietnam; March 2014 beginning in Hanoi,
including the Central Highlands and the Mekong Delta area of
southern Vietnam; and October–November 2016, concentrating on 7 northern provinces (Bogan and Do 2013a, 2013b,
2014a, 2014b, 2016). During each field trip, markets were
visited in the early morning and people selling freshwater
bivalves and gastropods were asked about where the animals
3
had been collected (Fig. 2). Other residents and fishermen
were asked about local freshwater mussels. We searched
streams and lakes crossed during our trips by wading and
feeling for mussels or using hand dredges. Collection methods
varied by location; some shell harvesters used handheld wire
baskets on a rope to collect mussels and gastropods in northern
Vietnam (Fig. 3). Long bamboo-handled rectangular wire
frame nets were used to collect mussels from stream and lake
bottoms in southern Vietnam (Fig. 4). One shell collector used
a surface-supplied-air diving apparatus to dive and collect
freshwater mussels.
The Vietnam Red Book was first developed in 1992, and it
was a collaboration of the Institute of Ecology and Biological
Resources (IEBR) and the International Union for Conservation of Nature (IUCN) with financial support from Sweden.
The criteria used in the book were built on the IUCN Red List
standards. The 1992 list was later revised and contained 13
species of freshwater mussels (Red Data Book of Vietnam
2000). This was expanded by the Institute of Science and
Technology of Vietnam. A country-wide assessment of the
conservation status of 416 species of animals in Vietnam
resulted in the publication of the Vietnam Red Data Book
(2007). This assessment included 11 species of freshwater
mussels; 1 species was added and 3 species were dropped from
the earlier lists: Cristaria herculea, Lamprotula liedtkei, and
Pilsbryoconcha suilla and was conducted using the IUCN
conservation assessment protocol in place at the time. The
Vietnam Red Data Book (2007) lists six Rare; four
Vulnerable, and a single indeterminate mussel species (Table
2).
Conservation assessments of all species listed here were
based upon the application of the criteria laid out in the IUCN
Red List Categories and Criteria (2012) and explained in the
IUCN Assessment Process (IUCN 2017) The categories of
threatened status include Extinct, Extinct in the Wild,
Critically Endangered, Endangered, Vulnerable, Near Threatened, Least Concern, Data Deficient, and Not Evaluated (see
Table 2). Each category is based on five criteria: (1) reduction
in population size, (2) geographic range, (3) population size
estimated to number at listed level for each threatened status,
(4) population size estimated to number less than a given level
for the threatened status, and (5) quantitative analysis showing
probability of extinction in the wild. The guidelines for the
application of these criteria are carefully explained by the
IUCN Standards and Petitions Subcommittee (2016). The
standards and an explanation of required data is presented in
tabular form based on the IUCN data entry format for
assessing the conservation status of each species (IUCN 2013).
The threatened species assessed by the IUCN from the IndoBurma region were initially listed in Köhler et al. (2012). The
conservation status of all the species listed in Table 2 and
supporting information can be found on the IUCN Red List
website (IUCN 2016).
This work has been registered with ZooBank and a copy
has been archived at Zenodo.org.
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4
DO ET AL.
Figure 1. Map of the major rivers of Vietnam.
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FRESHWATER MUSSELS OF VIETNAM
RESULTS
Margaritiferidae Henderson, 1929
Recent molecular studies of the Margaritiferidae have
considered it consists of a single genus, Margaritifera
Schumacher, 1816 (Bolotov et al. 2016b; Araujo et al.
2017). Bolotov et al. (2016b) recognized three clades, which
are treated as subgenera. Margaritifera laosensis is the only
species belonging to the subgenus Margaritanopsis Haas,
1910. Historically, M. laosensis was known from Dien Bien
Province in northwest Vietnam and in the adjacent part of Lao
People’s Democratic Republic (PDR). During recent fieldwork
in West Central Vietnam (Bogan and Do 2014a), a local stated
to have found similar shells but we did not procure any.
Phuong (2011) provided an IUCN conservation assessment.
The ecology and conservation of M. laosensis was detailed by
Bolotov et al. (2014), who listed three known viable
populations in the Lao PDR.
Unionidae Rafinesque, 1820
Aculamprotula nodulosa was reported from Cao Bang area
of northern Vietnam by Ða. ng et al. (1980). Graf and
Cummings (2007) continued to place this species in Lamprotula. However, Pfeiffer and Graf (2013) confirmed the split
of Lamprotula and Aculamprotula. Unio nodulosa was placed
in the genus Aculamprotula by He and Zhuang (2013) and
Graf and Cummings (2017). Live specimens have been
recently collected in northern Vietnam (Bogan and Do 2016).
Photographs presented by Graf and Cummings (2017) and
a photograph provided by S. Schneider (University of
Cambridge, Cambridge, UK, personal communication) are
identified as A. nodulosa, but these shells are much rounder
than the type figure of A. nodulosa.
Per Haas (1969a) Chamberlainia hainesiana [þ Simpsonia
demangei Rochebrune, 1904 is listed as occurring in Thailand,
Cambodia, and the Tonkin region of Vietnam. Simpsonia
demangei is listed from ‘‘Rivière Claire entre Vietri et TuyenQuas, (Tonkin)’’ (Rochebrune, 1904). Simpsonia demangei
was listed in the synonymy of C. hainesiana by Brandt (1974)
and Graf and Cummings (2017). Such a disjunct distribution
may suggest that the animal from Tonkin may be a separate
and distinct species from C. hainesiana. New live specimens
are needed to test the placement of this species described from
Vietnam using molecular methods.
Uniandra Haas, 1912 was used as a senior synonym of
Contradens Haas, 1913 by Brandt (1974). Contradens Haas,
1911 has priority over Uniandra Haas, 1912, based on the
dates of publication of the sections of Haas (1910c–1920)
(Bogan 2015).
Contradens contradens was listed by Brandt (1974) from
Thailand, Lao PDR, Cambodia and southern Vietnam [þ
Uniandra contradens tumidula (Lea, 1856)]. However, it was
not reported by Ða. ng et al. (1980). This species has been
collected from southern Vietnam (Bogan and Do 2014). Graf
and Cummings (2017) figured lots identified as Unio
5
semidecoratus Morlet, 1889, Tonkin Muséum national
d’Histoire naturelle, Paris (MNHN MP) 3861; Unio dautzenbergi Morlet, 1889, Tonkin; and Physunio crossei Deshayes,
1879, from Cochinchine MNHN MP 3774. All specimens
were identified as Contradens contradens by Graf and
Cummings (2017).
Considering that the type locality for Contradens semmelincki (Martens, 1891) is Borneo and the type locality for
Contradens fultoni is Manson, Tonkin (Mau Son, Lang Son),
C. fultoni is here considered a valid separate species. Graf and
Cummings (2017) treat C. fultoni as a junior synonym of C.
semmelincki.
Cristaria plicata is viewed as a valid, wide-ranging species
(Lopes-Lima et al. 2017). Ða. ng et al. (1980) recognized four
species of Cristaria in Vietnam: Cristaria bellua Morelet,
1866; Cristaria bialata Lea, 1829; Cristaria herculea
Middendorff, 1847; and Cristaria truncata Ða. ng, 1980.
Klishko et al. (2014, 2016) determined that C. herculea and
Cristaria tuberculata Schumacher, 1817 are synonyms of C.
plicata. Cristaria bellua, a misspelling of Anodonta bellus,
and Symphynota bialata were listed by Brandt (1974) and He
and Zhuang (2013) as synonyms of C. plicata.
Cristaria truncata is recognized as a valid species by Graf
and Cummings (2017) but is poorly known and its relationship
to C. plicata is unknown.
Cuneopsis demangei was described from Tonkin and
recognized by Haas (1969a), Ða. ng et al. (1980), and Graf and
Cummings (2017). This species is restricted to northern
Vietnam but has not been collected in the last several decades.
Cuneopsis pisiculus (Heude, 1874) was illustrated by Graf
and Cummings (2017) as University of Michigan Museum of
Zoology (UMMZ) 110095, purportedly from Tonkin, Vietnam. The accompanying original label listed it from Ningpo
and not from Tonkin. This is considered a spurious record and
not part of the fauna of Vietnam.
Diaurora aurorea was listed by Ða. ng et al. (1980), but has
not been collected recently. Early specimens identified as
Diaurora are housed in the Vietnam National University,
Hanoi University of Science, Museum of Biology, Hanoi, but
were not available for examination.
Ensidens sagittarius was reported by Brandt (1974) as a
synonym of Ensidens ingallsianus, but Graf and Cummings
(2017) treated E. ingallsianus as a separate species and
illustrated specimens from Cochinchine. However, the specimens listed as E. sagittarius from Cochinchine all appear to
be specimens of E. ingallsianus.
Gibbosula Simpson, 1900 was described containing only
Mya crassa Wood, 1815. Haas (1969a, 1969b) included
Gibbosula as a junior synonym of Lamprotula Simpson, 1900.
He and Zhuang (2013) and Graf and Cummings (2017)
recognized Gibbosula as a valid genus. Pfeiffer and Graf
(2013) recognized two clades in Lamprotula but did not
address the status of Gibbosula because the type species of
Lamprotula is poorly understood. Gibbosula is retained here
recognizing its uncertain status and placement.
Gibbosula crassa was listed as occurring in northern
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Species
IUCN
Red List
Vietnam Red
Data Books
CR
R
Chamberlainia hainesiana (Lea, 1856)
Contradens contradens (Lea, 1838)
Contradens fultoni Haas, 1939 *
Cristaria plicata (Leach, 1815)
Cristaria truncata Ða.ng, 1980
DD
LC
DD
DD
EN
V
Cuneopsis demangei Haas, 1929
Diaurora aurorea (Heude, 1883)
Ensidens ingallsianus (Lea, 1852)
Gibbosula crassa (Wood, 1815)
CR
DD
LC
CR
Harmandia somboriensis (Rochebrune, 1882)
Hyriopsis gracilis Haas, 1910
Hyriopsis delaportei (Crosse & Fischer, 1876)
Lamprotula bazini (Heude, 1877)
Lamprotula blaisei (Dautzenberg & Fischer, 1905)*
Lamprotula contritus (Heude, 1881)
Lamprotula leai (Gray in Griffith & Pidgeon, 1833)
Lamprotula quadrangulosa (Heude, 1881)
Lamprotula salaputium (Martens, 1902)*
Lanceolaria bilirata (Martens, 1902)*
Lanceolaria bogani Thach, 2016*
Lanceolaria fruhstorferi (Bavay & Dautzenberg, 1901)*
Lanceolaria gladiola (Heude, 1877)
Lanceolaria grayi (Gray in Griffith & Pidgeon, 1833)
Lanceolaria laevis (Martens, 1902)*
Margaritifera laosensis (Lea, 1863)
Nodularia dorri (Wattebled, 1886)*
Nodularia douglasiae (Gray, 1833)
Nodularia nuxpersica (Dunker, 1848)
Oxynaia diespiter (Mabille, 1887)*
Oxynaia gladiator (Ancey, 1881)*
Oxynaia jourdyi (Morlet, 1886)*
Oxynaia micheloti (Morlet, 1886)*
Physunio cambodiensis (Lea, 1856)
DD
LC
LC
DD
VU
EN
LC
LC
DD
CR
NE
NE
LC
DD
NE
EN
LC
LC
NE
EN
DD
NT
EN
LC
R
V
K-inderminate
R
R
V
R
Distribution Area (Province) in Vietnam, Comments
Northeast (Bang Giang River in Cao Bang, Ky Cung River in Lang Son, Thuong
River in Bac Giang, Tuyen Quang (IUCN listed as Lamprotula nodulosa)
Northeast (Phu Tho, Lo River in Tuyen Quang)
Mekong River delta (An Giang, Long An) IUCN listed as Uniandra contradens)
Northeast (Mau Son in Lang Son) (IUCN Listed as Uniandra semmelincki)
Northeast, Northwest, Red River delta
Northeast (Cao Bang, Bac Giang), Red River delta (Hai Duong, Ha Nam, Ninh
Binh, Nam Dinh)
Northeast (Da River in Phu Tho)
Northeast (?)
Central Highlands, South Central Coast, Southeast, Mekong River delta
Northeast (Bang Giang River in Cao Bang, Ky Cung River in Lang Son) (IUCN
listed as Lamprotula crassa)
Southeast, Mekong River delta
Southeast, Mekong River delta
Mekong River delta (An Giang)
Northeast (Ha Noi)
Northeast (Cao Bang, Lang Son, Bac Giang)
Northeast (Cao Bang, Bac Kan, Thai Thuyen)
Northeast
Northeast (Bang Giang River) (IUCN treated as synonym of L. leai not listed)
Northeast (Tuyen Quang)
Northeast (Cao Bang, Lang Son)
North Central Coast (Thua Thien Hue)
Northeast (Cao Bang, Lang Son) (IUCN lists as synonym of Lanceolaria grayana)
Northeast, Northwest, Red River delta
Northeast, Northwest, Red River delta
Northeast (Cong River in Thai Nguyen)
Northwest (Dien Bien Phu)
Northeast, Northwest, Red River delta
Northeast, Northwest, Red River delta, North Central Coast
Northeast (IUCN listed as Unio douglasiae)
Northeast (Bac Kan, Thai Nguyen)
Northeast
Northeast (Ha Noi, Vinh Phu, Phu Tho, Hoa Binh)
Northeast (Cao Bang, Phu Tho, Vinh Phuc)
Central Highlands (Serepok River in Dak Lak), Mekong River delta (An Giang)
DO ET AL.
Aculamprotula nodulosa (Wood, 1815)*
6
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Table 2. Complete list of freshwater mussel species found in Vietnam: 59 species, 28 genera, 2 families. The species conservation statuses are based on the International Union for Conservation of Nature (IUCN)
Red Listing (2016) and the Red Data Book of Vietnam (1992, 2000) and Vietnam Red Data Book (2007).
Species
IUCN
Red List
Physunio micropterus (Morelet, 1866)
LC
Physunio modelli Brandt, 1974
Physunio superbus (Lea, 1843)
Pilsbryoconcha compressa (Martens, 1860)
Pilsbryoconcha exilis (Lea, 1839)
Pilsbryoconcha lemeslei (Morelet, 1875)
LC
LC
LC
LC
LC
Pletholophus sp.*
Pletholophus tenuis (Gray in Griffith & Pidgeon, 1833)
Protunio messageri (Bavay & Dautzenberg, 1901)*
Pseudobaphia sp.
Pseudodon cambodjensis (Petit, 1865)
Pseudodon inoscularis (Gould, 1844)
Pseudodon mouhotii (Lea, 1863)
Pseudodon resupinatus Martens, 1902*
Pseudodon vondembuschianus (Lea, 1840)
Ptychorhynchus pfisteri (Heude, 1874)
Scabies crispata (Gould, 1843)
Sinanodonta hunganhi Thach, 2016*
Sinanodonta jourdyi (Morlet, 1866)*
Sinanodonta lucida (Heude, 1878)
Sinohyriopsis cumingii (Lea, 1852)
NE
LC
EN
NE
DD
LC
LC
EN
LC
NT
LC
NE
NE
NE
LC
Solenaia oleivora (Heude, 1877)
NE
Trapezoideus exolescens (Gould, 1843)
Trapezoideus misellus (Morelet, 1865)
Unionetta fabagina (Deshayes in Deshayes & Julien, 1874)
LC
DD
LC
Vietnam Red
Data Books
R
V
Distribution Area (Province) in Vietnam, Comments
Central Highlands (Dak Bla River tributary of Se San River in Kon Tum), Southeast
(Dong Nai), Mekong River delta
South Central Coast (Binh Thuan)
Mekong River delta (Can Tho)
Southeast, Mekong River delta (IUCN listed as synonym of P. exilis)
South Central Coast (Ninh Thuan), Southeast, Mekong River delta (Can Tho)
Central Highlands (Dak Bla River tributary of Se San River in Kon Tum, Khanh
Hoa)
Northeast, Red River delta
Throughout the country (IUCN listed as Cristaria tenuis)
Northeast (Bang Giang River in Cao Bang)
Northeast (Bang Giang River in Cao Bang)
Southeast, Mekong River delta
Southeast (Dong Nai), Mekong River delta
Southeast, Mekong River delta
Northeast (Vinh Phuc, Lang Son)
Southeast, Mekong River delta
Northeast (Vinh Phuc, Phu Tho), Northwest (Hoa Binh)
Northwest (Hoa Binh), Mekong River delta (An Giang)
North, Nghe An Province
Northeast, Northwest, Red River delta
Northeast, Northwest, Red River delta
Northeast (Bac Giang, Ha Noi, Ha Nam, Nam Dinh, Ninh Binh) (IUCN listed as
Cristaria cumingii)
Northeast (Phan River in Vinh Phuc, Day River in Ha Noi and Ha Nam, Thuong
River in Bac Giang)
Southeast, Mekong River delta
Northeast (Bang Giang River in Cao Bang, Vinh Phuc)
Central Highlands (Sa Thay River in Kon Tum)
FRESHWATER MUSSELS OF VIETNAM
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Table 2, continued.
CR ¼ Critically Endangered; DD ¼ Data Deficient; EN ¼ Endangered; IUCN ¼ International Union for Conservation of Nature; K ¼ Indeterminate; LC ¼ Least Concern; NE ¼ Not Evaluated; NT ¼ Near Threatened; R ¼ Rare; VU ¼
Vulnerable; V ¼ Vulnerable.
*Endemic to Vietnam.
7
8
DO ET AL.
Figure 2. Pan with several unionid species for sale in a market, Ha Noi,
Vietnam, Photograph by Arthur Bogan. November 18, 2014.
Vietnam by Ða. ng et al. (1980). Haas (1969a), Ða. ng et al.
(1980), and He and Zhuang (2013) all placed Lamprotula
mansuyi (Dautzenberg and Fischer, 1908), described from
northern Vietnam in the synonymy of Lamprotula crassa.
Live specimens of this species were recently collected from
the Bang River, Cao Bang Province, Vietnam, for anatomical
and molecular analyses (Bogan and Do 2016).
Harmandia somboriensis has been reported from the
Mekong and Mun rivers, but not mentioned as occurring in
Vietnam (Brandt 1974). Pfeiffer (personal communication)
found in it in shallow water around boulders. There are type
specimens collected from ‘‘Cochinchine’’ preserved in the
MNHN, Paris. This species collected from ‘‘Cochinchine’’ are
assumed to be part of the Vietnamese fauna.
The genus Hyriopsis Conrad, 1853 has some taxonomic
issues. Limnoscapha Lindholm, 1932 was proposed for a
group of bivalve species occurring from the Miocene to
Pliocene in what was, in 1978, the southern Soviet Union;
these species became extinct at the end of the Pliocene
(Gozhik 1978). This generic name was placed as a subgenus of
the modern Hyriopsis and has been used as a modern subgenus
(Modell 1950; Brandt 1974). Haas (1969b) and Graf and
Cummings (2006) listed Limnoscapha as a synonym of
Hyriopsis. Limnoscapha represents, in our opinion, an extinct
fossil group not related to the modern Hyriopsis species of
Asia.
Hyriopsis bialatus was listed from Malaysia, Thailand,
Cambodia, southern Vietnam, and Tonkin (Brandt 1974) and
recently confirmed from the Mekong Delta in Vietnam by
Bogan and Do (2014b). However, Hyriopsis bialatus has been
shown to be three separate species based on mitochondrial
DNA sequence data with H. bialatus being described from the
‘‘in Songi flumine Malaccae’’ (Sungi River, Malacca,
Malaysia) (Zieritz et al. 2016, 2017). Sungi is the Malay
Figure 3. Small clam rake used by local fishermen to collect freshwater
mussels from the local rivers. Photograph by Van Tu Do. Ca Lo River, Soc
Son District, Ha Noi, Vietnam. Photograph by Van Tu Do. November 23,
2012.
word for river, so the locality is unclear and Malaccae referred
to the southern side of peninsular Malaysia. An available name
for the species occurring in the Mekong River basin is
Hyriopsis gracilis Haas, 1910 (Haas 1910b). We are using H.
Figure 4. Long-handled rake with basket from southern Vietnam. Photograph
by Van Tu Do. April 4, 2014.
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FRESHWATER MUSSELS OF VIETNAM
gracilis for what has been historically listed as H. bialatus
only for the Mekong River basin populations.
Hyriopsis cumingii has been recognized by numerous
authors including Haas (1969a, 1969b), Brandt (1974), and He
and Zhuang (2013). Starobogatov (1970) described Sinohyriopsis with the type species Unio cumingii Lea, 1852 and
simultaneously described Nipponihyria Starobogatov, 1970
type species Hyriopsis schlegeli Martens, 1861. Ða. ng et al.
(1980) used the combination Sinohyriopsis cumingii. No use
of the name Nipponihyria has been found in the literature since
it was described. Sinohyriopsis is here given priority over
Nipponihyria based on usage since both names were published
in the same original work. Hyriopsis cumingii and H. schlegeli
have been placed in Sinohyriopsis based on recent phylogenetic work that separated them from the type species of
Hyriopsis, Hyriopsis bialatus, (Froufe et al. 2015; Lopes-Lima
et al. 2017). Sinohyriopsis cumingii is used here.
Hyriopsis delaportei was reported for the first time in An
Giang Province, Vietnam (Bogan and Do 2014b).
Lamprotula Simpson, 1900 was first divided by Wu (1998)
into Lamprotula and Aculamprotula based on differences in
shell shape, anatomy, and glochidial morphology. This
distinction was confirmed with molecular sequence data by
Zhou et al. (2007). Pfeiffer and Graf (2013) confirm this
division but raise questions about the other generic names
available for parts of this polyphyletic group.
Lamprotula bazini was reported from northern Vietnam by
Ða. ng et al. (1980) and recognized as valid by He and Zhuang
(2013) and Graf and Cummings (2017). This species has not
been collected in several decades.
Lamprotula blaisei was listed from northern Vietnam by
Ða. ng et al. (1980) and recognized as a valid by He and
Zhuang (2013) and Graf and Cummings (2017). This species
has not been collected in several decades.
Lamprotula contritus was reported from northern Vietnam
by Ða. ng et al. (1980). He and Zhuang (2013) and Graf and
Cummings (2017) consider this a synonym of Lamprotula
caveata (Heude, 1877). This species has not been collected
recently. Animals of this species from Vietnam do not appear
to be the same as L. caveata, as they lack the characteristic
depressions in the shell. We treat it here as a valid species
pending further genetic analysis.
Lamprotula leai was reported from northern Vietnam by
Ða. ng et al. (1980) and recognized as a valid species by He and
Zhuang (2013) and Graf and Cummings (2017). This species
is the most common species of this genus in northern Vietnam.
Lamprotula liedtkei (Rolle, 1904) was listed from northern
Vietnam by Ða. ng et al. (1980) and recognized as a junior
synonym of Aculamprotula nodulosa by He and Zhuang
(2013) and Graf and Cummings (2017). However, the
specimen of Quadrula liedtkei sp., with a Rolle manuscript
name figured by Graf and Cummings (2017) is from Tonkin
(United States National Museum, Smithsonian Institution
[USNM] 187462) and is much rounder than the figure of the
type of Unio liedtkei Rolle, 1904 and pictured by He and
Zhuang (2013). This USNM specimen resembles the shell
9
shape of Lamprotula similaris (Simpson, 1900) from China
(He and Zhuang 2013).
Lamprotula ponderosa (Dautzenberg and Fischer, 1905)
was recognized as a variety separate from Lamprotula leai.
However, it was listed as a junior synonym of Lamprotula leai
by Haas (1969a). He and Zhuang (2013) and Graf and
Cummings (2017) listed Unio (Quadrula) leai var. ponderosa
Dautzenberg and Fischer, 1905 as a primary junior homonym
of Unio ponderosa Rossmässler, 1842. It is also a junior
homonym to Unio ponderosa Hanley, 1842 and Unio
ponderosa Gray, 1825. The shell illustrated by Ða. ng et al.
(1980) may be a separate species or variation of L. leai.
Lamprotula quadrangulosa was described from China and
placed by Simpson (1914) as a junior synonym of L. caveata
(Heude, 1877) also described from China. Ða. ng et al. (1980)
viewed L. quadrangulosa as a valid species, separate from
both L. leai and L. caveata. Haas (1969a), He and Zhuang
(2013), and Graf and Cummings (2017) listed L. quadrangulosa as a synonym of L. caveata. Lamprotula quadrangulosa
from Vietnam does not appear to be the same as L. caveata,
lacking the characteristic depressions in the shell characteristic
of L. caveata. We treat L. quadrangulosa as a valid species
pending further genetic analysis.
Lamprotula salaputium described from Thuyen-Quan,
Annam, was not figured by Martens. Martens (1902) noted
he had a single specimen, making the specimen a holotype by
monotypy. Graf and Cummings (2017) do not provide a
picture of this species, but recognize it as a valid species. The
mollusk collections in Berlin and Frankfurt museums have
been queried and neither has this specimen. This species is an
unknown species described from Vietnam.
Ða. ng et al. (1980) reported four species of Lanceolaria
from northern Vietnam, Lanceolaria bilirata, Lanceolaria
fruhstorferi, Lanceolaria grayii and Lanceolaria laevis. Graf
and Cummings (2017) recognized three valid species from
Vietnam: L. bilirata, Lanceolaria gladiola, and L. grayii. This
genus needs a careful revision.
Lanceolaria bilirata was described from Tonkin, Vietnam,
and is apparently restricted to northern Vietnam. Haas (1969a)
placed this taxon as a subspecies of Lanceolaria oxyrhyncha
(Martens, 1894) and Kondo (2008) listed it as a synonym of
Lanceolaria grayana (Lea, 1834). The authors recognize this
species.
Lanceolaria bogani was recently described from Thura
Thien-Hue Province, Vietnam, and placed in the Unioninae by
Thach (2016a).
Lanceolaria fruhstorferi was recognized as a valid species
by Haas (1910c) but listed by He and Zhuang (2013) and Graf
and Cummings (2017) as a junior synonym of L. grayii. This
species was recognized as a separate valid species from
Vietnam by Thach (2016a).
Lanceolaria gladiola was described from China but Haas
(1911) and Graf and Cummings (2017) report it from Vietnam
and figure specimens that appear to be Lanceolaria grayii.
This is a doubtful record for Vietnam.
Lanceolaria grayii is the senior synonym for Unio
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DO ET AL.
grayanus Lea, 1834 described from China (Bogan and Do
2011). Graf and Cummings (2017) include L. fruhstorferi, L.
laevis and Lanceolaria gracillimus Rolle, 1904 as synonyms
of L. grayii. All three were described from Tonkin. This group
needs revision.
Lanceolaria laevis was originally described from Tonkin.
It has been recognized and figured by Ða. ng et al. (1980) and
Thach (2005, 2007). It was listed by He and Zhuang (2013)
and Graf and Cummings (2017) as a junior synonym of L.
grayii.
Nodularia Conrad, 1853 was erected with Unio douglasiae
Griffith & Pidgeon, 1833 as the type species. Simpson (1900,
1914) and Starobogatov (1970) used Nodularia as a valid
genus. However, Nodularia was considered a junior synonym
of Unio Retzius, 1788 (Haas 1969a, 1969b). Ða. ng et al.
(1980) included two taxa from northern Vietnam in Nodularia.
Graf and Cummings (2007) recognized the genus and included
four species, while Graf and Cummings (2017) included only
three species. Nodularidia Cockerell, 1901, was an unnecessary replacement name for Nodularia when it was considered
preoccupied. Nodularidia was used by He and Zhuang (2013)
for a single species in China. Nodularia and Unio represent
separate clades and may belong in separate tribes (Lopes-Lima
et al. 2017). We recognize three taxa in Nodularia from
Vietnam.
Nodularia dorri was reported by Ða. ng et al. (1980). Graf
and Cummings (2017) also used this species and noted its
distribution from the Gulf of Tonkin to the Mekong. It appears
to be restricted to northern and central Vietnam.
Nodularia douglasiae crassidens Haas, 1910 (Haas 1910a)
appeared in Simpson (1914) as a Nodularia (Nodularia)
douglasiae var. crassidens and was recognized by Ða. ng et al.
(1980). Haas (1969a) placed this taxon as a synonym of Unio
douglasiae douglasiae but listed it as occurring in China and
questionably in Cambodia. Graf and Cummings (2017)
synonymize this subspecies under Nodularia douglasiae.
Nodularia douglasiae is used here but it is recognized that
Nodularia needs extensive revision.
Nodularia nuxpersica is a new addition to the fauna of
Vietnam based on recent collections from markets in northern
Vietnam. He and Zhuang (2013) figured and included Unio
nuxpersica in the synonymy of Scabies crispata (Gould, 1843)
and Scabies chinensis Liu et al., 1991 (He and Zhuang 2013).
The Vietnam samples were shown to belong in Nodularia
based on DNA sequence data (Lopes-Lima et al. 2017).
Oxynaia was described by Haas, but the date of publication
has been confused. Haas (1969a) recorded it as 1911 but Haas
(1969b) used 1913. Starobogatov (1970) used Oxynaia Haas,
1912. Graf and Cummings (2017) noted the generic name was
associated with a figure in 1911 as a nomen nudum and only
described in 1913 (Haas 1913) and subsequently redescribed
in Haas (1913b). Based on the dates of publication for Haas
(1910c–1920) reported by Bogan (2015), plates 14, 15, 16 in
Haas (1911) are the first appearance of the generic name
Oxynaia associated with four described species published in
1911. Thus, the generic name Oxynaia was available from the
date of publication of the plates. This is considered an
indication for a genus named before 1931 under the
International Commission on Zoological Nomenclature (ICZN
1999) Code Article 12.11.2.2. The generic name Oxynaia
takes the date of publication as Haas, 1911 (see Bogan 2015).
The type species of Oxynaia was designated as Unio jourdyi
Morlet, 1886 by Haas (1913).
Oxynaia diespiter was described from Tonkin, and known
from the single type specimen and conchologically is very
close to Oxynaia jourdyi. Ða. ng et al. (1980) reported this
species from northern Vietnam. Graf and Cummings (2017)
illustrated a single specimen from Tonkin, MNHN_
MO_2998, which may be the unmarked holotype.
Oxynaia gladiator was described from Tonkin and
recognized as valid by Haas (1969a) and Graf and Cummings
(2017). Haas (1969a) listed this species from Tonkin and
Annam, Vietnam. However, it looks very much like Oxynaia
micheloti (Morlet, 1886). Adding this to the fauna of Vietnam
would bring the total Oxynaia species to four.
Oxynaia jourdyi is listed by Haas (1969a) as the type
species for the genus; it was described from Bac Hat etang de
la riviere Claire. Ða. ng et al. (1980) also listed this species.
Oxynaia micheloti was described from ‘‘Tonkin.’’ Haas
(1969a) mentions this species is only known from the original
description. Ða. ng et al. (1980) recognized the species. Based
on the figures provided by Graf and Cummings (2017), O.
micheloti does not have a very long shell and has a taller shell
than many of the lots figured as that species. Some of the other
figured lots appear to represent Nodularia species.
Physunio cambodiensis is listed from Cambodia, Lao PDR,
and Thailand by Haas (1969a), Brandt (1974), and Graf and
Cummings (2017). Davidson et al. (2006) collected this
species from a site on the Serepok River in Dak Lak Province,
and the Mekong River, An Giang and Long An Provinces,
Vietnam. These are the first records for this species in
Vietnam.
Physunio micropterus was reported by Brandt (1974) and
Graf and Cummings (2017) from Cambodia and Thailand.
Two specimens were reported from ‘‘Cochin China’’ or
southern Vietnam (Graf and Cummings 2017). Davidson et
al. (2006) collected this species from a site on the Dak Bla
River tributary of the Se San River, Kon Tum Province,
Vietnam. This species has been reported from Luy River, Binh
Thuan province.
Physunio modelli was described from north central Thailand and reported from Thailand and Lao PDR (Brandt 1974).
It has been reported from the Ham Tan district, Binh Thuan
Province by Thach (2007:190, plate 61, fig. 1057).
Physunio superbus was described from New Holland in
error and Brandt (1974) listed the distribution as possibly
including southern Vietnam. Ða. ng et al. (1980) did not report
this species. Graf and Cummings (2017) figure specimens
from Cochinchine or the Mekong Delta area of southern
Vietnam. There are specimens in the IEBR collections, Hanoi,
with the only locality data recorded as Vietnam.
Pilsbryoconcha compressa was recognized as a subspecies
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FRESHWATER MUSSELS OF VIETNAM
of Pilsbryoconcha exilis by Brandt (1974). Graf and Cummings (2017) listed P. compressa as a separate species. This
species was reported from An Giang Province, Vietnam
(Bogan and Do 2014b).
Pilsbryoconcha exilis was reported from southern Vietnam
(Brandt 1974). Graf and Cummings (2017) figured specimens
from Vietnam.
Pilsbryoconcha lemeslei was not mentioned by Ða. ng et al.
(1980). Haas (1969a) listed it only from Thailand and
Cambodia and Brandt (1974) did not mention this species
from Vietnam. Thach (2007) mentions this species from
Khanh Hoa, Vietnam, and figured a specimen that resembles
specimens figured by Graf and Cummings (2017).
Pilsbryoconcha suilla von Martens, 1902 was not
originally figured and was placed in the genus Pilsbryoconcha
by Simpson (1914). Simpson remarked that Martens thought
this species reminded him of Pilsbryoconcha. Ða. ng et al.
(1980) used the combination Pilsbryoconcha suilla but did not
figure this species. Anodonta suilla has been used as a junior
synonym of Sinanodonta woodiana by Haas (1969a) and Graf
and Cummings (2017). If it has a shell shape close to
Sinanodonta jourdyi Morlet, it would become a synonym of S.
jourdyi of Vietnam (Lopes-Lima, personal communication).
Cristaria discoidea (Lea, 1834) has been placed in
Cristaria (Pletholophus) by Simpson (1900, 1914) and Haas
(1969a). Unio tenuis Gray in Griffith and Pidgeon, has been
dated as 1834 and listed as a junior synonym of Unio
discoidea Lea, 1834 by Lea (1836, 1838, 1852, 1870),
Simpson (1900, 1914) and Haas (1969a). Petit and Coan
(2008) determined the date of publication of the figure of Unio
tenuis Gray in Griffith and Pidgeon as 1833, and noted Unio
tenuis has priority over the later name Unio discoidea Lea,
1834. This case does not meet the requirements of the Code
23.9 (ICZN 1999) for usage and the older name has date
priority and must prevail. The correct name for Cristaria
discoidea is Cristaria tenuis (Gray in Griffith and Pidgeon,
1833). Cristaria tenuis was used by He and Zhuang (2013)
and Graf and Cummings (2017). Ða. ng et al. (1980) elevated
Pletholophus to generic level and included three species:
Pletholophus swinhoei (Adams, 1866); Pletholophus inangulatus (Haas, 1910a), and Pletholophus discoideus (Lea, 1834).
All of them are considered synonyms under Cristaria tenuis
(He and Zhuang 2013; Graf and Cummings 2017). Placement
of C. tenuis in Pletholophus and separate from Cristaria was
confirmed by Lopes-Lima et al. (2017).
DNA sequence analyses of Pletholophus samples from
Vietnam revealed a second distinct species, here assumed to
represent a new species, Pletholophus sp. (Lopes-Lima,
personal communication).
Protunio Haas, 1912 was described by Haas but various
dates have been listed for its publication. Haas (1969a, 1969b),
Starobogatov (1970), and Graf and Cummings (2017) all listed
1913 (Haas 1913) as the date of publication of Protunio. Graf
and Cummings (2017) cited Haas 1912 (plate 32) for Protunio
but used 1913 as the date for the generic description (Haas
1913). Based on the dates of publication for Haas (1910c–
11
1920) reported by Bogan (2015), plate 32 is the first
appearance of the generic name Protunio associated with a
described species published in 1912 (Bogan 2015). Thus, the
generic name Protunio was available from the date of
publication of the plate and since Protunio was published in
association with a single species, Unio messageri Bavay and
Dautzenberg, 1901 is the type species by monotypy. This is
considered an indication for a genus named before 1931 under
ICZN Code Article 12.112.2. The generic name Protunio
takes the date of publication as Haas, 1912 (see Bogan 2015).
Protunio was subsequently redescribed by Haas with the date
on the signature of 1914 and the date on the cover of the
Lieferung containing the four signatures of 1919 (Bogan
2015).
Protunio messageri was recognized by Haas (1969a,
1969b) and Graf and Cummings (2017) as restricted to
northern Vietnam. Ða. ng et al. (1980) reported this species
from around Cao Bang and Lang Son. However, the specimen
` (in press) is
illustrated by Ða. ng et al. (1980) and Ða. ng and Hô
not Protunio messageri when compared with the original
figures (Heude 1877) and figures provided by Graf and
Cummings (2017). The specimen figured by Ða. ng et al.
` (in press) represents a different
(1980) and Ða. ng and Hô
species. The figured shell shape is very similar to Pseudobaphia biesiana (Heude, 1877) figured by Haas (1910c–1920),
He and Zhuang (2013), and Graf and Cummings (2017) but
represents an undescribed species, Pseudobaphia sp. Pseudobaphia sp. is known from only three lots of specimens, one lot
in IEBR, Hanoi, one lot in the North Carolina Museum of
Natural Sciences, Raleigh, and a large lot in the Vietnam
National University, Hanoi University of Science, Museum of
Biology, Hanoi. This species has not been collected since
1971.
Pseudodon inoscularis is the type species of Pseudodon
and has been reported from throughout Southeast Asia. Brandt
(1974) recognized one species as a Rassenkreis or a ring
species with a variety of subspecies. This group is in dire need
of a taxonomic revision. Graf and Cummings (2017) claimed
the range of P. inoscularis from South Vietnam but did not
illustrate any specimens.
Pseudodon callifer (Martens, 1860) was listed by Brandt
(1974) as subspecies of P. inoscularis callifer and reported
from Thailand, Cambodia and southern Vietnam.
Pseudodon cambodjensis was not reported by Brandt
(1974) or Ða. ng et al. (1980) from Vietnam, but Graf and
Cummings (2017) list a specimen (Museum of Comparative
Zoology, Harvard [MCZ] 37431) from Cochinchine and
mapped it in southern Vietnam.
Pseudodon ellipticus Conrad, 1865 was listed by Graf and
Cummings (2017) from Thailand, Cambodia, and southern
Vietnam but they did not figure any specimens. It was
described from Cambodia. Brandt (1974) used it as a
subspecies of Pseudodon vondembuschianus.
Pseudodon inoscularis was recognized by Graf and
Cummings (2017), who listed it from southern Vietnam. Haas
(1969a) listed this species from Tenasserim, Myanmar [þ
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DO ET AL.
Burma], only. Brandt (1974) treated the species as a ring
species ranging from Myanmar, Thailand, Malaysia, Lao
PDR, Cambodia, and southern Vietnam. This wide-ranging
species appears to grade from one form to another across its
range and should be carefully examined with molecular
techniques.
Pseudodon mouhotii was recognized and listed from
Vietnam by Brandt (1974) and Graf and Cummings (2017),
but was not discussed by Ða. ng et al. (1980). Pseudodon exilis
(Morelet, 1866) was considered by Brandt (1974) a junior
synonym of Pseudodon mouhotii.
Pseudodon resupinatus was described from Than Moi,
Tonkin, but was not originally figured. It is recognized as a
valid species by Simpson (1914), Haas (1920), Ða. ng et al.
(1980), and Graf and Cummings (2017). This species is
endemic to northern Vietnam.
Pseudodon vondembuschianus, as used by Brandt (1974),
contained three subspecies including Pseudodon vondembuschianus ellipticus Conrad, 1865, reported from southern
Vietnam. Ða. ng et al. (1980) and Graf and Cummings (2017)
do not list this species from Vietnam. It is found in southern
Vietnam.
Ptychorhynchus pfisteri was reported by Ða. ng et al. (1980)
from northern Vietnam. The species was not listed from
Vietnam by Haas (1969a, 1969b), He and Zhuang (2013), or
Graf and Cummings (2017). Recently specimens have been
collected by the authors in Hanoi.
Scabies crispata was reported by Ða. ng et al. (1980) and
Brandt (1974) from Vietnam. Graf and Cummings (2017)
report this species from Cochinchine and Tonkin, Vietnam,
but some of the specimens figured appear to be specimens of
Nodularia. Recently, a field survey conducted by L. A.
Prozorova and N. X. Quang (personal communication)
recorded this species from Bung Binh Thien, An Phu, An
Giang (southern Vietnam).
Sinanodonta hunganhi was described from around Vinh
City, Nghê An Province, in northern Vietnam and reported it
living ‘‘along rivers’’ [sic] (Thach 2016b). Based on the
published figure, this species appears to fit within the shell
variation of S. jourdyi.
Sinanodonta jourdyi and Sinanodonta elliptica (Heude,
1878) were identified from northern Vietnam (Ða. ng et al.
1980). Specimens collected from Vietnam were identified as
both species and were examined genetically. These specimens
represent a single species, distinct from Sinanodonta woodiana of China (Lopes-Lima, personal communication). The
name available for the Vietnamese species is S. jourdyi.
Because S. elliptica was described from China, it is likely a
synonym of S. woodiana and was not found in Vietnam
(Lopes-Lima, personal communication).
Sinanodonta lucida was reported from North Vietnam
(Ða. ng et al. 1980). Graf and Cummings (2017) treated S.
lucida as a junior synonym of S. woodiana. Bolotov et al.
(2016a) reported it as a separate, valid species but from China.
Sinanodonta woodiana has been assumed to be a wideranging and plastic species, with 103 synonyms listed for this
species (Graf and Cummings 2017). Haas (1969a) treated all
three taxa reported from Vietnam as synonyms of Anodonta
(Anodonta) woodiana (Lea, 1834). Ða. ng et al. (1980)
recognized three species in the genus Sinanodonta in northern
Vietnam, S. jourdyi (Morlet, 1886), S. elliptica (Heude, 1878),
and S. lucida (Heude, 1878). He and Zhuang (2013) and Graf
and Cummings (2017) listed all three taxa as synonyms of S.
woodiana. Preliminary DNA sequence analyses have separated S. jourdyi from S. woodiana occurring in China. Because
the type locality for S. elliptica is China, it is not recognized in
Vietnam and is considered part of the greater S. woodiana
complex (Lopez-Lima, personal communication). The complexity of S. woodiana has been documented for Asia and the
species invading Europe (Bolotov et al. 2016a). Bolotov et al.
(2016a) documented seven separate lineages within what has
been named Sinanodonta woodiana. These analyses also
separate Sinanodonta lucida as a separate linage within the S.
woodiana complex. This taxonomic puzzle will require further
analyses to resolve this group.
Solenaia oleivorus has been collected from the Phan River
(Vinh Phuc Province), Day River (Ha Noi and Ha Nam
provinces), and Thuong River (Bac Giang province) in
Vietnam. This species was synonymized by He and Zhuang
(2013) under Solenaia iridinea (Heude, 1874) in China, but
determined to be a separate and valid species by Ouyang et al.
(2011). The authors recognize it as a valid species.
Trapezoideus was recognized as containing a single
species with three subspecies by Brandt (1974). He remarked
Trapezoideus might occur in southern Vietnam. Ða. ng et al.
(1980) recognized Trapezoideus misellus. Graf and Cummings
(2017) placed T. misellus as a junior synonym of Trapezoideus
excolescens (Gould, 1843). Preliminary genetic analyses have
suggested Trapezoideus is not a monotypic genus, but may
harbor several cryptic species (Lopes-Lima, personal communication). Now, we have chosen to recognize the wide-spread
species T. exolescens in southern Vietnam and T. misellus
from northern Vietnam.
Unionetta fabagina was listed by Haas (1969a) from
Cambodia and Lao PDR. Brandt (1974) reported Unionetta
fabagina from the Mekong River, but not from Vietnam. A
shell of U. fabagina has been collected from Sa Thay River,
Kon Tun and Gia La provinces (Central Highlands), Vietnam.
DISCUSSION
Distribution of the Fauna
The unionid fauna of Vietnam can be divided along the
major river basins draining the country (Fig. 1). The Red River
basin is the major basin in the north, passing through Hanoi.
There are representatives of the fauna in the far north of
Vietnam that have relationships with the fauna of the Zhu
River basin and the Yangtze River basin farther north in
southern China (e.g., Lamprotula species). The western border
areas of Vietnam, including Dien Bien Province and rivers of
the Central Highland and southern Vietnam, are drained by
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FRESHWATER MUSSELS OF VIETNAM
tributaries to the Mekong River basin in the north. The
Mekong River fauna described by Brandt (1974) for Thailand,
Lao PDR, and Cambodia extends into southern Vietnam in the
distributaries in the Mekong Delta. The unionid fauna in the
eastern rivers of the Central Highland is still poorly known.
Seventeen species are considered endemic to Vietnam and
comprise 28.8% of the total species found in Vietnam (Table
2). Forty species were found in the north while 18 species
were reported from the south and only one species
(Sinanodonta jourdyi) is widely distributed throughout the
country (Table 2; see also Supplemental Individual Species
Maps).
Our recent surveys demonstrated that the northeastern area
of Vietnam possess the highest diversity of freshwater mussels
in Vietnam. Se San and Serepok, tributaries of the Mekong
River, can be ranked as a second hotspot of diversity of this
group. It is necessary to conduct more surveys in the northwest
of Vietnam in Dien Bien Province and the Central Highlands
where data are quite scarce. We have only spent part of a
single field trip in southern Vietnam and need to spend more
time in the Mekong Delta area of Vietnam.
Conservation Assessment
The IUCN hosted a training session in Phenom Penh,
Cambodia, and a workshop in Vientiane, Lao PDR, to assess
the conservation status of the freshwater fauna of the IndoBurma area, including Vietnam (Allen et al. 2012). Freshwater
bivalves and gastropods were a part of this program (Köhler et
al. 2012). Our current conservation assessment began with the
information from IUCN Red List (2016) and is supplemented
with our recent field data and museum records. The IUCN
conservation status of the freshwater bivalves of Vietnam is
summarized in Table 2.
Based on the 59 species recorded in Vietnam (Table 2),
those taxa assessed in the IUCN Red List (2016) included 4
species (6.8%) assessed as Critically Endangered, 7 species
(12%) assessed as Endangered, 1 species (1.7%) assessed as
Vulnerable, 2 species (3.4) assessed as Near Threatened, 23
species (39%) assessed as Least Concern, 11 species (18.6%)
assessed as Data Deficient, and 11 species (18.6%) that were
not evaluated.
Based on our survey results beginning in 2010 and using
the IUCN Red List categories and criteria (IUCN 2012), we
herein recommend that the conservation status of following
four species should be changed and we will submit a revised
species assessment to the IUCN:
(1) Solenaia oleivora should be assessed as Vulnerable. It was
found only in the Phan, Day, and Thuong rivers; population
reduction was estimated about 50% over 10 yr; extent of
occurrence was estimated as smaller than 20,000 km2. This
species is assessed only on the Vietnamese portion of its range.
(2) Lamprotula quadrangulosa should be assessed as
Vulnerable. It was found only in the Bang River; population
reduction was estimated about 50% over 10 yr; extent of
13
occurrence was estimated as smaller than 20,000 km2. This
species is assessed only on the Vietnamese portion of its range.
(3) Trapezoideus misellus should be assessed as Vulnerable. It
was found only in the Bang River; population reduction was
estimated about 50% over 10 yr; extent of occurrence was
estimated as smaller than 20,000 km2. This species assessed
only on data from northern Vietnam.
(4) Contradens fultoni should be assessed at least as
Endangered. It was found in only the Mau Son River, Lang
Son Province, based on an early record from 1930; extent of
occurrence was estimated as smaller than 5,000 km2. This
species is endemic to Vietnam.
We are particularly concerned about the status and
continued survival of four species in the Bang River basin
and tributaries of Li Chiang in northeastern Vietnam and
China: Aculamprotula nodulosa, Lamprotula contritus, Lamprotula quadrangulosa, and Gibbosula crassa. These species
have been very scarce in the past several decades. Lamprotula
blaisei, Contradens fultoni, Pseudobaphia sp., and Cuneopsis
demangei have not been seen or collected since the early
1970s. No shells or live specimens of these last four taxa were
collected during our fieldwork, strongly suggesting that these
taxa are extirpated from their former range in Vietnam. The
first three species are endemic to Vietnam and they may
already be extinct.
Impacts
Freshwater mussels are long-lived animals (10 to 200 yr)
and have a unique life cycle with a parasitic larval stage on the
gills or fins of fish, so negative impacts on the species may not
be immediately apparent (Bogan 1993, 2006; Vaughn and
Taylor 1999; Vaughn 2010; Haag 2012). Mussels provide a
variety of ecosystem services that directly and indirectly
impact the local human populations including biofiltration,
food source for animals and humans, and resource materials
(e.g., inlay materials) (Vaughn 2017). Negative impacts on
freshwater mussels are numerous, including channel modification and habitat destruction from dredging; sedimentation;
clear-cutting of watersheds; monoculture cropping; loss of
riparian buffers along streams; pollution in many forms
including fertilizers, pesticides, industrial effluents, and
domestic sewage; mining; urbanization and damming of
rivers; commercial exploitation; introduced species; expansion
of nonnative parasite hosts; and the loss of native host fish
(e.g., Dudgeon et al. 2006; Gillis 2012; Haag 2012; Vaughn
2012; Sousa et al. 2014; Zieritz et al. 2016). There is no
research on the impacts of current agricultural and forestry
practices on the unionid fauna of Vietnam; however, research
in Malaysia (Zieritz et al. 2017) has indicated that such
activities have had a negative effect on the distribution of
freshwater mussels.
Many impacts on the freshwater environment were visible
during our surveys including deforestation, road construction
(Fig. 5), in-stream sand and gravel mining (Fig. 6), open-pit
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14
DO ET AL.
Figure 5. Road construction and disposal of debris over the edge of the road
and the impact on the local river; the river is the same color as the earth being
dumped. Lang Son Province, Vietnam. Photograph by Arthur Bogan.
November 11, 2012.
mining runoff (Fig. 7), harvesting for food (Fig. 2), domestic
pollution, and construction of various dams and hydroelectric
projects (Fig. 8). Streams have been locally modified by
restriction of channels, diversion of water for rice fields, and
terracing of flood plains for rice production. Most of the large
trees have been cut from the mountains, affecting the rain
runoff patterns and thus affecting water temperature and clarity
and increasing runoff (see Naiman and Dudgeon 2011; Zieritz
et al. 2016). These modifications are impacting freshwater
mussels and gastropod populations. A few species are doing
well in disturbed habitats and show up in the markets as food
items, including Sinanodonta spp., Cristaria plicata, Pletholophus tenuis, and Nodularia spp. (V.T. Do, personal
observations).
Current and planned dams in the Mekong River basin and
their impacts have been summarized by Winemiller et al.
(2016). Dams and reservoirs not only impact the mussel and
Figure 6. In-stream gravel mining, Dien Bien Province, Vietnam. Photograph
by Arthur Bogan. November 19, 2012.
Figure 7. Open-pit mining, Cao Bang Province, Vietnam. Photograph by
Arthur Bogan. November 13, 2012.
fish fauna in the footprint of the reservoir but also below the
reservoir. Downstream of the dam (Fig. 8) water flow patterns,
water temperature, and chemistry are changed and can
dramatically impact mussels, their biology, and native host
fish (Vaughn and Taylor 1999).
Mussels are used a food item and are intensively collected
in some areas of Vietnam. A local mussel harvester
commented that currently he is only able to collect about
100 kg per day where in the past he was collecting 500 to 800
kg per day to sell in local markets. This decline in harvest may
be due to overharvesting or mussel population declines due to
a combination of overharvesting, disturbance, and pollution.
Consider the number of people across Vietnam collecting
mussels each day and this impact on a yearly basis is
staggering. Decline in the mussel fauna of Vietnam can be
considered on a local scale as well as considering the national
impact on the freshwater bivalve fauna. These impacts will
also impact those species that range beyond Vietnam’s
borders.
Figure 8. Dam in central Vietnam with the riverbed downstream dewatered.
Photograph by Arthur Bogan. November 18, 2014.
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FRESHWATER MUSSELS OF VIETNAM
15
Conservation Recommendations
ACKNOWLEDGMENTS
In addition to established national parks and conservation
areas, beginning in 2008 and extending to 2020, Vietnam will
establish 45 areas for protection of inland water bodies. Some
parts of the Red and Da rivers (in northern Vietnam) will be
included in protected areas. However, conservation activities
for freshwater bivalves have never been mentioned. About
60% of the Vietnamese freshwater bivalve fauna is currently
imperiled. Some freshwater bivalve species already appear to
have disappeared from northern Vietnam and the future is
bleak for endangered freshwater bivalve species.
New outreach materials need to be developed for
distribution to local people, aquaculture agencies, primary
and high schools, and universities which illustrate the
Vietnamese freshwater bivalve fauna and their unique life
cycle. Aquaculture programs can be encouraged to develop
captive propagation programs that will identify the native host
fish for species to assist in conservation. Public activities and
education on the importance and role of freshwater bivalves
have not been mentioned. Some of freshwater bivalve species
already seem to have disappeared from northern Vietnam and
there is reason to be pessimistic about the future of endangered
freshwater bivalve species.
Knowledge of the taxonomy of marine mollusks is actively
expanding due to activities of the amateur shell collectors who
are regularly contributing to the description of new species
(see Bouchet et al. 2016). Developing a public program not
only to provide information on freshwater mollusks of
Vietnam, but also a website to provide free information or
assist with identifications, is needed. This would stimulate
more public involvement in examining the local freshwater
fauna. Shell clubs are common in the United States and
Europe and there are already shell dealers and people
interested in shells in Vietnam (see Thach 2005, 2007, 2012,
2016b). These people would be a great source of information
and local knowledge about these animals and should be
encouraged to participate. A template might be the Cornell
FeederWatch project.
We want to express our gratitude for the support of the
Institute of Ecology and Biological Resources (IEBR),
Vietnam Academy of Science and Technology (VAST), and
North Carolina Museum of Natural Sciences for their support
of this collaborative research. We also acknowledge the help
of our assistants, Mr. Quoc Toan Phan, Nguyen Tong Cuong,
Nguyen Dinh Tao, and our drivers, Mr. Nguyen Ngoc Quyen,
who was our driver for three trips, and Mr. Dang Thanh Ha,
who was our driver for the 2016 trip. We also appreciate the
Nagao Natural Environment Foundation support for Do Van
Tu’s research project ‘‘The study of Unionoida biodiversity
and conservation status in Bang and Ky Cung River basins in
Northeast Vietnam.’’ Ms. Jamie M. Smith and Mrs. Heather B.
Leslie have been instrumental in databasing specimens
deposited in the North Carolina Museum of Natural Sciences
Mollusk Collection from Vietnam. Mrs. Cynthia M. Bogan
and Ms. Jamie M. Smith have reviewed the manuscript for
content and English composition. We thank our anonymous
reviewers for their very helpful and thorough comments and
suggestions; they have been very helpful.
Future Research Needs
Our understanding of the phylogenetic relationships of the
freshwater mussels of Vietnam is only beginning to be
explored (e.g., Lopes-Lima et al. 2017). Using results of our
expanding molecular analyses, we have recognized a smallsized species, Nodularia nuxpersica, new to Vietnam, and one
undescribed new species, Pletholophus sp. By comparing
photographs and museum collections, we have recognized a
misidentified species as new, Pseudobaphia sp. If the results
of the work in Malaysia are any indication, some of the
currently recognized Vietnamese unionid species may be
overturned as some species are being recognized as a species
complex (e.g., Hyriopsis bialatus) (Zieritz et al. 2016). This
effort will require continued collaboration with colleagues
throughout Asia.
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