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Corresponding author: Peng Bun Ngor ( pengbun.ngor@gmail.com ) Academic editor: Graham Oliver
© 2020 Ting Hui Ng, Ekgachai Jeratthitikul, Chirasak Sutcharit, Samol Chhuoy, Kakada Pin, Arthit Pholyotha, Warut Siriwut, Ruttapon Srisonchai, Zeb S. Hogan, Peng Bun Ngor.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ng TH, Jeratthitikul E, Sutcharit C, Chhuoy S, Pin K, Pholyotha A, Siriwut W, Srisonchai R, Hogan ZS, Ngor PB (2020) Annotated checklist of freshwater molluscs from the largest freshwater lake in Southeast Asia. ZooKeys 958: 107-141. https://doi.org/10.3897/zookeys.958.53865
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The Tonle Sap Lake in Cambodia is a crucial freshwater biodiversity hotspot and supports one of the world’s largest inland fisheries. Within the Tonle Sap basin, freshwater molluscs provide vital ecosystem services and are among the fauna targetted for commercial harvesting. Despite their importance, freshwater molluscs of the Tonle Sap basin remain poorly studied. The historical literature was reviewed and at least 153 species of freshwater molluscs have been previously recorded from throughout Cambodia, including 33 from the Tonle Sap basin. Surveys of the Tonle Sap Lake and surrounding watershed were also conducted and found 31 species, 15 bivalves (five families) and 16 gastropods (eight families), in the Tonle Sap basin, including three new records for Cambodia (Scaphula minuta, Novaculina siamensis, Wattebledia siamensis), the presence of globally invasive Pomacea maculata and potential pest species like Limnoperna fortunei. This study represents the most comprehensive documentation of freshwater molluscs of the Tonle Sap basin, and voucher specimens deposited at the Inland Fisheries Research and Development Institute, Cambodia, represent the first known reference collection of freshwater molluscs in the country. In order to combat the combined anthropogenic pressures, including invasive species, climate change and dams along the Mekong River, a multi-pronged approach is urgently required to study the biodiversity, ecology, ecosystem functioning of freshwater molluscs and other aquatic fauna in the Tonle Sap basin.
alien species, bivalves, Cambodia, diversity, economic species, Lower Mekong basin, snails, Tonle Sap Lake
“Clams, cockles, snails, et cetera can be obtained just by cupping one’s hands into the Fresh Water Lake” (Zhou in
Being the largest natural lake in Southeast Asia, the Tonle Sap Lake is a crucial freshwater habitat for various animals, including fish and birds (
In addition, there has been interest in economically-important molluscs, including freshwater apple snails (Ampullariidae) and Asian clams (Cyrenidae) (
Furthermore, most of the recent studies did not include voucher specimens nor photographs of species that could be used by local stakeholders and research scientists. The lack of proper reference material may lead to incorrect identification of species, allowing non-native species to establish and spread unnoticed (e.g.,
It is therefore imperative to document the biodiversity of freshwater molluscs of Tonle Sap as a foundation for further evolutionary and ecological research, and to make available the necessary information to the government, local residents, and other stakeholders to allow for more effective management of threatened, economic- and medically-important freshwater molluscs in this unique freshwater habitat. In this study, we aimed to conduct 1) a review of historical literature on freshwater molluscs of Cambodia; and 2) surveys of Tonle Sap Lake and the surrounding watershed to provide an updated checklist of species diversity for the freshwater molluscs in the largest lake in Southeast Asia, which is also one of the world’s most productive lakes.
Historical data on freshwater mollusc records from Cambodia were gathered from relevant literature based on a search of the Global Biodiversity Information Facility (GBIF.org 2019), the IUCN Red List (
We conducted surveys in and around the Tonle Sap Lake and its major tributaries over two one-week periods in May and December 2019, respectively. Samples were taken from 44 locations from the study area, covering major landing sites (Chhnouk Tru in Kampong Chhnang Province, Kampong Loung in Pursat Province, Chong Khneas in Siem Reap Province and Boeung Chhmar Ramsar wetlands in Kampong Thom Province), major Tonle Sap tributaries (Pursat, Sangkae, Mongkol Borei, Serei Saophoan, Sreng, Chi Kraeng, Staung, Sen, and Chinit Rivers), rain-fed and flooded zones along national road No. 5 and 6 around the Tonle Sap Lake (Fig.
Sampling localities of freshwater molluscs collected from the Tonle Sap basin in 2019.
No. | Locality code and locality details | Coordinates |
---|---|---|
1 | C078-Serei Saophoan River, Serei Saophoan city, Banteay Meanchey Province | 13°35'08.9"N, 102°58'41.2"E |
2 | C073-Serei Saophoan River, Preah Ponlea, Serei Saophoan District, Banteay Meanchey Province | 13°34'33.1"N, 102°58'58.5"E |
3 | C017-Paoy Samraong, Preah Netr Preah, Preah Netr Preah District, Banteay Meanchey Province | 13°35'01.8"N, 103°09'24.8"E |
4 | C075-River in Chob Vari, Preah Netr Preah District, Banteay Meanchey Province | 13°37'24.0"N, 103°11'50.0"E |
5 | C080-Lotus pond in Phnum Lieb, Preah Netr Preah District, Banteay Meanchey Province | 13°36'31.8"N, 103°18'02.3"E |
6 | C079-River in Phnum Lieb, Preah Netr Preah District, Banteay Meanchey Province | 13°36'16.9"N, 103°19'02.0"E |
7 | C074-Sreng River in Kralanh City, Kralanh District, Siem Reap Province | 13°35'29.8"N, 103°24'15.8"E |
8 | C018-Tonle Sap Lake open area near Chong Khneas, Chong Khneas, Siem Reap Province | 12°30'20.1"N, 103°50'06.2"E |
9 | C085-Siem Reap River, Leang Dai, Angkor Thom District, Siem Reap Province | 13°29'24.3"N, 103°55'14.4"E |
10 | C081-Stream near Banteay Srei District, Siem Reap Province | 13°35'44.0"N, 103°57'43.3"E |
11 | C020-Tonle Sap Lake open area near Kampong Phluk, Prasat Bakong District, Siem Reap Province | 13°11'16.2"N, 103°57'25.4"E |
12 | C021-Tonle Sap Lake open area near Kampong Phluk, Prasat Bakong District, Siem Reap Province | 13°11'40.6"N, 103°57'50.5"E |
13 | C082-Chi Kraeng River in Kampong Kdei, Chi Kraeng District, Siem Reap Province | 13°07'55.3"N, 104°20'16.8"E |
14 | C023-Ponds in Thnol Keng, Kampong Kdei, Chi Kraeng District, Siem Reap Province | 13°05'25.4"N, 104°23'32.5"E |
15 | C024-Paddy fields near Trach, Kampong Chen Cheung, Stoung District, Kampong Thom Province | 12°58'43.7"N, 104°33'52.3"E |
16 | C009-Tonle Sap Lake open area in Kampong Thom Province | 12°42'42.5"N, 104°11'58.2"E |
17 | C008-Tonle Sap Lake open area near Pean Bang, Stoung District, Kampong Thom Province | 12°43'36.9"N, 104°14'11.6"E |
18 | C007-River flows from Boeng Tonle Chhma to Tonle Sap Lake, Stoung District, Kampong Thom Province | 12°44'15.4"N, 104°15'30.9"E |
19 | C004-Don Sdeung, Peam Bang, Staung District, Kampong Thom Province | 12°46'25.4"N, 104°17'02.9"E |
20 | C005-Don Sdeung, Boeung Chhmar Fish Sanctuary, Staung District, Kampong Thom Province | 12°47'25.1"N, 104°17'55.7"E |
21 | C006-Provincial Fisheries Office, Boeung Chhmar Fish Sanctuary, Staung District, Kampong Thom Province | 12°48'48.6"N, 104°18'11.6"E |
22 | C087-Sen River in Balang, Damrei Choan Khla, Stung Sen District, Kampong Thom Province | 12°41'48.5"N, 104°53'58.8"E |
23 | C025-Sen River in Kampong Samraung, Srayav, Stung Sen District, Kampong Thom Province | 12°40'58.3"N, 104°54'33.1"E |
24 | C090-Tang Krasang River in Santuk District, Kampong Thom Province | 12°34'04.4"N, 105°03'02.3"E |
25 | C092-Chinit River in Kampong Thma, Santuk District, Kampong Thom Province | 12°29'45.7"N, 105°07'18.4"E |
26 | C088-Canal from Makara Dam, Ballangk, Baray District, Kampong Thom Province | 12°29'13.8"N, 105°08'47.0"E |
27 | C068-Recreational area, Sangkae River in Traeng, Rotanak Mondol, Battambang Province | 12°49'51.4"N, 102°55'44.3"E |
28 | C071-Sangkae River in Chaeng Mean Chey, Banan District, Battambang Province | 12°52'03.5"N, 103°06'03.8"E |
29 | C070-Sangkae River in Chheu Teal, Banan District, Battambang Province | 12°58'54.7"N, 103°08'36.2"E |
30 | C014-Sangkae River in Wat Ta Meum, Oudambang Muoy, Sangkae District, Battambang Province | 13°04'04.7"N, 103°12'17.2"E |
31 | C067-River in Ou Ta Paong, Bakan District, Pursat Province | 12°39'34.8"N, 103°40'26.2"E |
32 | C013-Pursat River near Damnak Ampel Irrigation Dam, Damnak Ampel, Lolork Sor, Sampaov Meas District, Pursat Province | 12°29'19.3"N, 103°48'29.7"E |
33 | C012-Pursat River in Wat Loung, Lolork Sor, Sampaov Meas District, Pursat Province | 12°30'20.1"N, 103°50'07.8"E |
34 | C064-Pursat River in Sorya, Krong Pursat, Pursat Province | 12°31'00.3"N, 103°54'54.7"E |
35 | C066-Thliem Ma-Orm River in Boeng Kantuot, Krakor District, Pursat Province | 12°31'37.3"N, 104°03'13.8"E |
36 | C011-Tonle Sap Lake open area in Krakor District, Pursat Province | 12°35'42.1"N, 104°12'13.1"E |
37 | C010-Tonle Sap Lake open area in Krakor District, Pursat Province | 12°38'20.2"N, 104°12'12.1"E |
38 | C003-Kampong Chhnok Tru landing point, Chhnok Tru, Boribo District, Kampong Chhnang Province | 12°30'36.5"N, 104°27'18.2"E |
39 | C063-Tributary of Tonle Sap Lake in Phumi Phsar, Kampong Chhnang Province | 12°22'50.7"N, 104°29'0.2"E |
40 | C002-Tonle Sap River in Kampong Prasat, Saeb, Kampong Tralach District, Kampong Chhnang Province | 12°04'23.6"N, 104°46'23.8"E |
41 | C097-Tonle Sap River in Kaoh Thkov, Chol Kiri District, Kampong Chhnang Province | 12°03'31.7"N, 104°46'22.4"E |
42 | C001-Boeung Po, Sorvong, Tacheise, Kampong Tralach District, Kampong Chhnang Province | 12°01'28.3"N, 104°43'54.1"E |
43 | C094-Tonle Sap River in Samretthi Chey, Kampong Tralach district, Kampong Chhnang Province | 11°53'55.2"N, 104°46'01.6"E |
44 | C093-Tonle Sap River in Kaoh Chen, Popnhea Lueu district, Kandal Province | 11°49'01.6"N, 104°48'40.5"E |
All the specimens were identified to genus or species level based on shell characteristics by referring to the historical literature with original species descriptions,
Voucher specimens are deposited in the following institutions:
CIFI Inland Fisheries Research and Development Institute, Cambodia;
MUMNH Mahidol University Museum of Natural History, Thailand;
Almost 300 species of freshwater molluscs have previously been recorded from Cambodia to date, but only 153 are currently considered to be valid species and among these, 33 species were recorded from the Tonle Sap basin (Table
A third of the previously recorded species are medically-important Pomatiopsidae and Stenothyridae, which are intermediate hosts of zoonotic parasites like Schistosoma mekongi Voge, Bruckner & Bruce, 1978, and are distributed in the Mekong River (
At least 15 species of bivalves from five families, and 16 species of gastropods from eight families (Table
Summary of freshwater molluscs recorded in historical records of Cambodia, historical records that specify the Tonle Sap basin, and collected from the Tonle Sap basin in 2019. Higher classification follows
Higher classification | Order | Superfamily | Family | Number of species | ||
---|---|---|---|---|---|---|
Historical records of Cambodia | Historical records mentioning Tonle Sap basin | Present study of Tonle Sap basin | ||||
Bivalvia | ||||||
Pteriomorphia | Arcida | Arcoidea | Arcidae | 1 | – | 1 |
Mytilida | Mytiloidea | Mytilidae | 2 | 2 | 2 | |
Heterodonta | Adapedonta | Solenoidea | Pharidae | – | – | 1 |
Venerida | Cyrenoidea | Cyrenidae | 14 | 6 | > 1 | |
Palaeoheterodonta | Unionida | Unioidea | Unionidae | 40 | 13 | 10 |
Gastropoda | ||||||
Caenogastropoda | Architaenioglossa | Ampullarioidea | Ampullariidae | 7 | 4 | 4 |
Viviparoidea | Viviparidae | 19 | 3 | 4 | ||
Littorinimorpha | Truncatelloidea | Bithyniidae | 4 | 1 | 2 | |
Iravadiidae | 1 | – | – | |||
Pomatiopsidae | 46 | – | – | |||
Stenothyridae | 6 | – | – | |||
Neogastropoda | Buccinoidea | Nassariidae | 7 | 2 | 2 | |
(Cohort) Sorbeoconcha | Cerithioidea | Pachychilidae | 2 | 1 | 1 | |
Thiaridae | 2 | 1 | 1 | |||
Heterobranchia | ||||||
Hygrophila | Lymnaeoidea | Bulinidae | 1 | – | 1 | |
Lymnaeidae | 1 | – | 1 | |||
Total | 153 | 33 | 31 |
Superfamily Arcoidea Lamarck, 1809
Family Arcidae Lamarck, 1809
Scaphula minuta Ghosh, 1922: 1143–1144. Type locality: “Tale Sap or inland Sea of Singgora on the east coast of peninsular Siam”.
CIFI.MOL.044,
Pursat River in Pursat Province, and Sreng River in Siem Reap Province (locality no. 7 and 32). Found attached to rocks in slow-moving waters.
Scaphula minuta was first described from Songkhla Lake in southern Thailand (Ghosh 1929) and only recently reported to be distributed in southern Vietnam (
Superfamily Mytiloidea Rafinesque, 1815
Family Mytilidae Rafinesque, 1815
Volsella fortunei Dunker, 1856: 361, 362. Type locality: “Mare Chinense”.
Limnoperna siamensis:
Limnoperna fortunei:
Tonle Sap River at Kampong Chhnang Province, and Tonle Sap Lake at Kampong Chhnang and Siem Reap Provinces (locality no. 8, 12, 37, 38 and 40). Occur in colonies attached by byssus threads to hard surfaces like man-made structures (e.g., boats, jetties, wooden pillars of homes), on unionid bivalves, and occasionally the shells of large freshwater gastropods.
Limnoperna fortunei is native to East Asia, and although previously thought to be naturally distributed in the countries south of China (e.g.,
Dreissena harmandi Rochebrune, 1882: 102. Type locality: “Lac de Rhom-Penh, Mekong”.
Sinomytilus harmandi:
Similar to that of Limnoperna fortunei.
Sinomytilus harmandi appears to be limited in range to the Lower Mekong River basin, and it may have previously been mistaken for Limnoperna fortunei, resulting in a lack of historical records (
Superfamily Solenoidea Lamarck, 1809
Family Pharidae H. Adams & A. Adams, 1856
Novaculina siamensis Morlet, 1889: 198, pl. 9, fig. 4. Type locality: “Marais de Chantakam Siam”.
MUMNH.PHA.001, MUMNH.PHA.002
Tonle Sap River in Kampong Chhnang Province (locality no. 41 and 43); tough clay bottom substrate, in which it makes cylindrical holes.
The discovery of Novaculina siamensis in Tonle Sap River is a new record for Cambodia and fills in the distribution gap of the genus in Indochina. This species was first described from “Chantakam, Siam” [Prachantakham District, Prachinburi Province, Thailand], but the type series is thought to be lost (
Freshwater bivalves of the Tonle Sap basin, Cambodia (Pharidae and Unionidae) A Novaculina siamensis B Bineurus mouhotii C Contradens contradens D Ensidens ingallsianus E Hyriopsis bialata F Hyriopsis delaportei G Monodontina cambodjensis H Physunio micropterus I Pilsbryoconcha linguaeformis J Pilsbryoconcha lemeslei and K Scabies mandarinus. Scale bars: 10 mm, unless stated otherwise. Photographs by K Macharoenboon.
Superfamily Cyrenoidea Gray, 1840
Family Cyrenidae Gray, 1840
CIFI.MOL.029, CIFI.MOL.030, CIFI.MOL.031, CIFI.MOL.032, MUMNH.COR.0194, MUMNH.COR.0195, MUMNH.COR.0196, MUMNH.COR.0197, MUMNH.COR.0198, MUMNH.COR.0199, MUMNH.COR.0200, MUMNH.COR.0201, MUMNH.COR.0202, MUMNH.COR.0203, MUMNH.COR.0204, MUMNH.COR.0205, MUMNH.COR.0206, MUMNH.COR.0207, MUMNH.COR.0208, MUMNH.COR.0209, MUMNH.COR.0210, MUMNH.COR.0211, MUMNH.COR.0212,
Tonle Sap River and Lake, and most tributaries (locality no. 3, 6, 7, 8, 11, 12, 16, 18, 19, 20, 21, 22, 23, 24, 27, 32, 38, 39 and 40). Buried just below surface of soft, muddy substrate of the water bodies.
Approximately 17 different species of Corbicula have previously been recorded from Cambodia (Suppl. material
Corbicula species are the only bivalves that are commercially harvested from the Lake to be sold locally and exported abroad for human consumption and as animal feed (Fig.
Superfamily Unionoidea Rafinesque, 1820
Family Unionidae Rafinesque, 1820
Monocondylaea mouhotii Lea, 1863: 190. Type locality: “Laos Mountains, Cambodia, Siam”.
Pseudodon mouhoti:
Bineurus mouhotii:
MUMNH.UNI.2637, MUMNH.UNI.2646, MUMNH.UNI.2659 MUMNH.UNI.2670.
Phumi Phsar River, Kampong Chnnang Province and Sangkae River, Battambang Province (locality no. 27 and 39); in sandy substrate of streams.
Bineurus mouhotii is widespread along the Mekong basin in Thailand, Laos and Vietnam, and probably in northern Myanmar and Yunnan (
Unio contradens Lea, 1838: 75, pl. 18, fig. 58. Type locality: Java.
Uniandra contradens tumidula:
Uniandra contradens rustica:
Uniandra contradens fischeriana:
CIFI.MOL.017, CIFI.MOL.018, MUMNH.UNI.2621, MUMNH.UNI.2629, MUMNH.UNI.2633, MUMNH.UNI.2648, MUMNH.UNI.2651,
Tonle Sap Lake at Kampong Chhnang and Siem Reap Provinces, and Chhnok Tru landing point (locality no. 8, 11, 18, 36, 37, and 39); in soft muddy substrate.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. Contradens contradens was recently noted for its high varation in shell morphology dued to phenotypic plasticity (
Unio ingallsianus Lea, 1852: 282, pl. 24, fig. 41. Type locality: “Siam”.
Ensidens ingallsianus ingallsianus:
Ensidens ingallsianus:
CIFI.MOL.019, CIFI.MOL.020, CIFI.MOL.021, CIFI.MOL.022, MUMNH.UNI.2617, MUMNH.UNI.2626, MUMNH.UNI.2634, MUMNH.UNI.2642, MUMNH.UNI.2644, MUMNH.UNI.2649, MUMNH.UNI.2655, MUMNH.UNI.2657, MUMNH.UNI.2666, MUMNH.UNI.2668, MUMNH.UNI.2672, MUMNH.UNI.2673,
Tonle Sap Lake at Kampong Chhnang, Siem Reap and Kampong Thom Provinces, Sen River in Kampong Thom Province, Sangkae River in Battambang Province, Chi Kraeng River and Kralanh River in Siem Reap Province, Tonle Sap River in Kandal Province (locality no. 3, 7, 8, 12, 13, 21, 23, 24, 30, 36, 40 and 44); in soft muddy substrate.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. The species previously identified as Ensidens ingallsianus has recently been revealed as a complex of two species (
Unio delphinus Gruner, 1841: 276, pl. 9, fig. 1a–c. (non Spengler, 1793) Type locality: “sungi flumine, Malaccae”.
Hyriopsis bialatus Simpson, 1900: 579. (new replacement name for Unio delphinus).
Hyriopsis (Hyriopsis) bialatus:
CIFI.MOL.007, CIFI.MOL.008, CIFI.MOL.009, MUMNH.UNI.2618, MUMNH.UNI.2622, MUMNH.UNI.2627, MUMNH.UNI.2630, MUMNH.UNI.2635, MUMNH.UNI.2650, MUMNH.UNI.2652, MUMNH.UNI.2658, MUMNH.UNI.2662,
Tonle Sap River in Kampong Chhnang Province and Tonle Sap Lake at Kampong Chhnang and Siem Reap Provinces, Sen River in Kampong Thom Province, Sangkae River, Battambang Province (locality no. 8, 11, 16, 23, 27, 36, 37, 38 and 40); in soft muddy substrate.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. At some parts of the Tonle Sap Lake, shells of Hyriopsis bialata were often covered in mats of Limnoperna fortunei and Sinomytilus harmandi. Hyriopsis bialata is widespread in Indochina, from Thailand to Peninsular Malaysia, and along the middle Mekong basin to the Mekong Delta in southern Vietnam (
Unio (Arconaia) delaportei Crosse & Fischer, 1876: 327–329, pl. 10, fig. 1, pl. 11, fig. 5. Type locality: “Cambodge, dans la province de Compong-Soai”.
Hyriopsis (Hyriopsis) delaportei:
CIFI.MOL.025, CIFI.MOL.026, MUMNH.UNI.2623, MUMNH.UNI.2628, MUMNH.UNI.2631, MUMNH.UNI.2653,
Tonle Sap Lake at Kampong Chhnang, Kampong Thom, Pursat and Siem Reap Provinces (locality no. 11, 16, 37 and 38); in soft muddy substrate.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. Shells of Hyriopsis delaportei were also found to be covered in mats of Limnoperna fortunei and Sinomytilus harmandi. Hyriopsis delaportei was originally described from “Compong Soai; Cambodia” (
Monocondylaea cambodjensis Petit de la Saussaye, 1865: 16, pl. 4, fig. 4. Type locality: “Battabbang, Cambodge”.
Pseudodon cambodjensis cambodjensis:
Monodontina cambodjensis:
CIFI.MOL.023, CIFI.MOL.024, MUMNH.UNI.2637, MUMNH.UNI.2646, MUMNH.UNI.2659, MUMNH.UNI.2670,
Tonle Sap River in Kampong Chhnang Province, Pursat River in Pursat Province and Sen River in Kampong Thom Province, Chi Kraeng River in Siem Reap Province (locality no. 3, 13, 23 and 33); in soft muddy or sandy substrate.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. Monodontina cambodjensis has been recorded from Thailand and Cambodia, in several tributaries of the Mekong River (
Unio micropterus Morelet, 1866: 63, 64. Type locality: “in torrentibus montanis Cambodiae”.
Physunio micropterus:
CIFI.MOL.013, CIFI.MOL.014, CIFI.MOL.015, MUMNH.UNI.2639, MUMNH.UNI.2641, MUMNH.UNI.2656, MUMNH.UNI.2661, MUMNH.UNI.2665, MUMNH.UNI.2667, MUMNH.UNI.2671,
Pursat River in Pursat Province, Sangkae River in Battambang Province, Sen River in Kampong Thom Province, Chi Kraeng River and Sreng River in Siem Reap Province (locality no. 7, 13, 22, 23, 27, 30, 32 and 39); in sandy substrate.
The distribution of Physunio micropterus is restricted to the Tonle Sap basin. There are some reports outside its endemic range, such as in the Ping and Prachinburi rivers in Thailand, but these distributions need to be confirmed (
Anodonta linguaeformis Morelet, 1875: 329, pl. 14, fig. 5. Type locality: “au Cambodje, probablement dans les marécages voisins de Battambang”.
Pilsbryoconcha linguaeformis:
Pilsbryoconcha exilis linguaeformis:
MUMNH.UNI.2616, MUMNH.UNI.2619, MUMNH.UNI.2624, MUMNH.UNI.2625, MUMNH.UNI.2636, MUMNH.UNI.2645
Pond in Kampong Chhnang Province, Tonle Sap Lake in Kampong Chhnang and Kampong Thom Provinces, Banteay Meanchey Province, Sen River in Kampong Thom Province (locality no. 3, 20, 36, 38, 40, and 42); in soft muddy substrate and swampy grounds.
Not sold for food, collected as by-catch of Corbicula and Mekongia harvests, and were often observed to be discarded along with other large unionids. Distribution range of this species seems to be limited to the Tonle Sap basin.
Anodonta lemeslei Morelet, 1875: 328, pl. 14, fig. 1. Type locality: “Cambodge”.
Pilsbryoconcha lemeslei:
MUMNH.UNI.2669
Tributary of Tonle Sap Lake near Preah Tis Bridge in Chi Kraeng District, Siem Reap Province (locality no. 13); in soft muddy substrate of still water.
This species is rare and known only from few places in Thailand, Laos, Cambodia and Vietnam (
Unio mandarinus Morelet, 1864: 159. Type locality: “Cochinchina”.
Scabies crispata:
Scabies mandarinus:
CIFI.MOL.001, CIFI.MOL.002, CIFI.MOL.003, CIFI.MOL.004, CIFI.MOL.004, CIFI.MOL.005, CIFI.MOL.006, MUMNH.UNI.2620, MUMNH.UNI.2632, MUMNH.UNI.2638, MUMNH.UNI.2640, MUMNH.UNI.2643, MUMNH.UNI.2647, MUMNH.UNI.2654, MUMNH.UNI.2663, MUMNH.UNI.2664, MUMNH.UNI.2672,
Tonle Sap Lake open area in Banteay Meanchey and Siem Reap Provinces, Tonle Sap River in Siem Reap Province and Chhnok Tru landing point, Sangkae River in Battambang Province, Pursat River in Pursat Province, Tang Krasang River in Kampong Thom Province, Sreng River in Siem Reap Province (locality no. 2, 3, 7, 8, 12, 24, 27, 30, 32, 36 and 38); in soft muddy and sandy substrate.
Scabies mandarinus was described by Morelet in 1864, based on specimens collected from “Cochinchina”. This name was previously placed as a junior synonym under Scabies scobinatus (Lea, 1856) and Scabies crispata (Gould, 1843) (e.g.,
Superfamily Ampullarioidea Gray, 1824
Family Ampullariidae Gray, 1824
Ampullaria gracilis Lea, 1856: 110. Type locality: “Siam”.
Pila gracilis:
Tonle Sap River, Tonle Sap Lake open areas, and small ponds and paddy fields in the provinces surrounding the Lake (locality no. 4, 7, 9, 10, 24 and 28).
The first record of this species from Cambodia was from the Kampong Svay District in Kampong Thom Province (
Freshwater mollusc fisheries at the Tonle Sap Lake, Cambodia. Small-scale harvesting of freshwater molluscs A Corbicula spp. and B Mekongia rattei, at open areas of the Lake. Freshwater molluscs C Corbicula spp. and D Pila spp., being sorted and packed at main landing sites around the Lake. Photographs by A Pholyotha (A, B), TH Ng (C), PB Ngor (D).
Freshwater gastropods of the Tonle Sap basin, Cambodia A Pila gracilis B Pila pesmei C Pila virescens D Pomacea maculata E Filopaludina martensi cambodjensis F Idiopoma umbilicata G Mekongia rattei H Trochotaia trochoides I Bithynia siamensis goniomphalus J Wattebledia siamensis K Anentome cambojiensis L Anentome helena M Sulcospira housei N Melanoides tuberculata O Radix rubiginosa and PIndoplanorbis exustus. Scale bars: 10 mm, unless stated otherwise. Photographs by TH Ng.
Ampullaria pesmei Morlet, 1899: 185, pl. 8, fig. 2. Type locality: “Phnom-Penh (Cambodge)”.
Pila pesmei:
Tonle Sap open area (locality no. 38).
Although Pila pesmei is widely distributed in Eastern and Northeastern Thailand (
Ampullaria virescens Deshayes, 1824: un-numbered plate. Type locality: unknown.
Pila polita:
CIFI.MOL.010,
Tonle Sap River and Lake, paddy fields and irrigation ponds in Kampong Chhnang, Banteay Meanchey and Siem Reap Provinces (locality no. 3, 13, 14, 18, 35, 38 and 42).
Pila virescens is the largest among the native ampullariids, and commonly harvested for food (Fig.
Pomacea maculata
Perry, 1810: unnumbered plate and text. Type locality: Paraná, Argentina (see discussion in
CIFI.MOL.037,
Tonle Sap River and Lake, and paddy fields in Banteay Meanchey and Kampong Thom Provinces (locality no. 1, 2, 5, 7, 15, 22, 38, 40, and 44).
Unlike Pila species, Pomacea maculata is not native to Southeast Asia. Pomacea maculata (as Pomacea insularum d’Orbigny, 1835 in Hayes 2008), and another species, Pomacea canaliculata (Lamarck, 1822), have both been introduced to Asia from South America (
Family Viviparidae Gray, 1847
Paludina cambodjensis Mabille & Le Mesle, 1866: 135, pl. 7, fig. 4. Type locality: “Moth-Kasa, dans les marais”.
Filopaludina (Siamopaludina) martensi cambodjensis:
CIFI.MOL.033, CIFI.MOL.041,
Tonle Sap River and Lake, and surrounding watershed including rivers, irrigation ponds and paddy fields (locality no. 1, 2, 3, 5, 8, 9, 10, 11, 12, 13, 16, 17, 19, 20, 21, 22, 24, 32, 37, 38, 40, 42 and 44).
Filopaludina martensi cambodjensis is said to be lacking in spiral ridges compared to Filopaludina martensi martensi (Frauenfeld, 1865) (see
Paludina umbilicata Lea, 1856: 109. Type locality: “Takrong River, Siam”.
Idiopoma umbilicata:
Tonle Sap Lake and tributaries at Kampong Chhnang, Battambang, Siem Reap and Kampong Thom Provinces, irrigation ponds and paddy fields at Banteay Meanchey and Siem Reap Provinces (locality no. 7, 13, 24, 29 and 39).
Idiopoma umbilicata is often found together with Filopaludina martensi cambodjensis and can be differentiated from the latter species by its smaller size and shouldered shells. It does not appear to be harvested for food.
Paludina rattei Crocsse & Fischer, 1876: 317. Type locality: “Stung Chinit, Cambodia”.
Mekongia rattei:
CIFI.MOL.042, MUMNH.VIV.001, MUMNH.VIV.002, MUMNH.VIV.003,
Tonle Sap Lake; Chi Kraeng River and Sreng River in Siem Reap Province; Phumi Phsar River in Kampong Chhnang Province (locality no. 7, 8, 11, 13, 16, 20, 24, 27, 36, 37 and 39).
Mekongia rattei is sold in local markets surrounding the Lake. This species along with Corbicula spp. are commercially harvested from the Lake to be sold locally and exported abroad for human consumption and as animal feed in local poultry farms (Fig.
Paludina trochoides Martens, 1860: 12. Type locality: “Siam”.
Trochotaia trochoides:
CIFI.MOL.034,
Paddy fields at Banteay Meanchey Province (locality no. 7).
We found only a few dry shells in the northwestern province of Banteay Meanchey, close to the Thai border. The species was also occasionally encountered for sale at local markets.
Superfamily Truncatelloidea Gray, 1840
Family Bithyniidae Gray, 1857
Paludina goniomphalos Morelet, 1866: 167. Type locality: “Cochinchina”.
Bithynia (Digoniostoma) siamensis goniomphalus:
CIFI.MOL.040,
Found at the edges of Tonle Sap River and Lake, ponds and in paddy fields at Kampong Chhnang, Banteay Meanchey, and Siem Reap Provinces (locality no. 3, 4, 8, 11, 16, 40, 42, 43, and 44).
Bithynia siamensis goniomphalus is of medical importance because it is an intermediate host of the zoonotic parasite, Opisthorchis viverrini (Poirier, 1886) (TROPMED Medical Group 1986). The role of Bithynia siamensis goniomphalus in the transmission of this parasite in Cambodia has not been investigated in detail, although the parasite has been recorded in freshwater fishes at the border of the Kandal-Takeo Provinces in the south (
Wattebledia siamensis Möllendorff, 1902: 160. Type locality: “Siam”.
Wattebledia siamensis:
Found among floating vegetation along the banks of the Tonle Sap River (locality no. 44).
This is the first record of the species in Cambodia, and the species is known throughout Thailand (
Superfamily Buccinoidea Rafinesque, 1815
Family Nassariidae Iredale, 1916 (1835)
Melania cambojiensis Reeve, 1861: Melania species 468, pl. 59. Type locality: “Cambojia”.
Clea (Anentome) cambojiensis:
CIFI.MOL.039,
Tonle Sap Lake; on muddy substrate (locality no. 8, 11, 12, 17 and 37).
The type specimen of Anentome cambojiensis was collected by Henri Mouhot (
Melania helena von dem Busch in Philippi, 1847: 170, pl. 4, fig. 4. Type locality: Java.
Clea (Anentome) helena:
CIFI.MOL.035,
Tonle Sap River, Lake and surrounding tributaries, irrigation ponds and canals (locality no. 2, 7, 8, 10, 11, 13, 16, 17, 21, 22, 24, 25, 26, 34, 38, 40, 41, 42, and 44).
The species recognised as Anentome helena in Indo-China may in fact be a complex of three species (
Superfamily Cerithioidea J. Fleming, 1822
Family Pachychilidae Fischer & Crosse, 1892
Melania housei Lea, 1856: 144–145. Type locality: “Korat, Takrong River, Siam”.
Adamietta housei:
CIFI.MOL.043,
Tonle Sap River, Sangkae River in Battambang Province, canal in Kampong Thom Province (locality no. 1, 2, 26, 38, 41, 43 and 44).
Sulcospira housei is widespread in neighbouring Thailand (
Nerita tuberculata OF Müller, 1774: 191. Type locality: “In littore Coromandel”.
Melanoides tuberculata:
CIFI.MOL.036,
Tonle Sap Lake and Thliem Ma-Orm River in Pursat Province (locality no. 11, 12, 20, 27 and 35).
Melanoides tuberculata has a global distribution but was not found to be common in the Tonle Sap basin during our surveys.
Superorder Hygrophila Férussac, 1822
Family Bulinidae P. Fischer & Crosse, 1880
Planorbis exustus Deshayes, 1833: 417, pl. 1, figs 11–13. Type locality: “Lieux man~cagieux de la cote de Malabar”.
Indoplanorbis exustus:
Dry shells found at a lotus pond in Banteay Meanchey Province (locality no. 5).
Indoplanorbis exustus has been recorded from Cambodia since the 1800s (
Lymnoeus rubiginosus
Michelin, 1831: Moll. no. 22, pl. 22. Type locality: “Indes Orientales” (from
Lymnaea (Radix) auricularia rubiginosa:
At the edges of irrigation ponds and swampy grounds in Banteay Meanchey Province (locality no. 3).
Radix rubiginosa is a cosmopolitan species, widely distributed from Indo-China to Sundaland (
Many of the 19th to early 20th century descriptions and records of freshwater molluscs from Cambodia were collected by expeditions, sponsored or led by the French, including those by the renowned traveller Henri Mouhot and the diplomat Auguste Pavie, and also by missionaries based in the country (e.g.,
Our surveys have revealed that there remains much to be done in resolving the taxonomy and systematics of freshwater molluscs in Cambodia. In addition to the four species of Viviparidae that were highlighted in the Results as requiring taxonomic revision, many other taxa have not been collected or studied closely since the species were described, including the Cyrenidae, Nassariidae, and the speciose Pomatiopsidae (Suppl. material
Aside from the taxonomic confusion, the Ampullariidae of Tonle Sap Lake are extremely unique in terms of the high volume of production for Pila species being harvested from Tonle Sap Lake, compared to elsewhere in Southeast Asia, where the native ampullariids appear to be declining whilst invasive confamilial Pomacea species are increasing (
Although the mytilid mussels are assumed to be native to Cambodia, Limnoperna fortunei in particular, has the potential to be classified as a pest species. The mixed species colonies of Limnoperna fortunei, and to a lesser extent, Sinomytilus harmandi, have byssus threads that form dense mats, not only on hard man-made surfaces, but also on the shells of unionid mussels and gastropods (Fig.
Indeed, the comtemporary Mekong River system is now at a critical point in time, facing challenges of regional development, particularly the increasing numbers of hydropower dams, which alter the timing, magnitude and frequencies of seasonal flow of the tropical flood-pulse system. Such flow alterations have been demonstrated to dampen the seasonal flood pulses (
Our checklist is the first step toward more extensive research on freshwater molluscs in the Tonle Sap basin. It is imperative that more surveys of freshwater molluscs be conducted across different wet and dry seasons to allow for a better representation of the fauna to be captured, along with baseline data of the populations and ecology of the species to be documented. The presence of globally-invasive species like Pomacea maculata, and the prevalence of pest species like Limnoperna fortunei, which have the potential to replace and negatively impact the ecosystem and native species of the Tonle Sap basin is another major concern. In order to combat the combined pressures of invasive species, land cover change, climate change, dams along the main stem and tributaries of the Mekong River, among many other anthropogenic threats (
We thank the USAID-supported Wonders of the Mekong project, Inland Fisheries Research and Development Institute (IFReDI) of the Fisheries Administration Ministry of Agriculture, Forestry and Fisheries of Cambodia, Chulalongkorn University, Mahidol University and National University of Singapore for funding for the fieldwork, and the institutional support for the maintenance of the reference collections of the IFReDI, Mahidol University Museum of Natural History and Lee Kong Chian Natural History Museum. We are grateful to Y. Limpanont for assistance in identifying bithynid species. We thank Arthur Bogan and an anonymous reviewer for their constructive comments that improved the manuscript.
Table S1. Historical records of freshwater molluscs from Cambodia
Data type: records
Explanation note: Historical records of freshwater molluscs from Cambodia based on literature review.