ZooKeys 958: 107–141 (2020) A peer-reviewed open-access journal doi: 10.3897/zookeys.958.53865 CHECKLIST https://zookeys.pensoft.net Launched to accelerate biodiversity research Annotated checklist of freshwater molluscs from the largest freshwater lake in Southeast Asia Ting Hui Ng1, Ekgachai Jeratthitikul2, Chirasak Sutcharit3, Samol Chhuoy4,5, Kakada Pin5, Arthit Pholyotha3, Warut Siriwut2, Ruttapon Srisonchai6, Zeb S. Hogan5,7, Peng Bun Ngor4,5 1 Lee Kong Chian Natural History Museum, Faculty of Science, National University of Singapore, 2 Con- servatory Drive, 117377, Singapore 2 Animal Systematics and Molecular Ecology Laboratory, Department of Biology, Faculty of Science, Mahidol University, Bangkok 10400, Thailand3 Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, 254 Phayathai Road, Pathumwan, Bangkok 10330, Thailand 4 Inland Fisheries Research and Development Institute (IFReDI), Fisheries Admin- istration, No. 86, Norodom Blvd., PO Box 582, Phnom Penh, Cambodia 5 Wonders of the Mekong Project, c/o IFReDI, No. 86, Norodom Blvd., PO Box 582, Phnom Penh, Cambodia 6 Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand7 Department of Biology, University of Nevada, 1664 N. Virginia Street, Reno, NV 89557, USA Corresponding author: Peng Bun Ngor ([email protected]) Academic editor: Graham Oliver | Received 13 May 2020 | Accepted 3 June 2020 | Published 11 August 2020 http://zoobank.org/AB196008-1542-49D4-B23E-1892D2191C18 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 Abstract 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, freshwa- ter 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, in- cluding 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 Copyright Ting Hui Ng et al. 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. 108 Ting Hui Ng et al. / ZooKeys 958: 107–141 (2020) 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. Keywords alien species, bivalves, Cambodia, diversity, economic species, Lower Mekong basin, snails, Tonle Sap Lake Introduction “Clams, cockles, snails, et cetera can be obtained just by cupping one’s hands into the Fresh Water Lake” (Zhou in Uk and Uk 2016). Being the largest natural lake in Southeast Asia, the Tonle Sap Lake is a cru- cial freshwater habitat for various animals, including fish and birds (Campbell et al. 2006), but besides the above mentioned brief account by the Chinese diplomat, Zhou Daguan, in the 13th century (Uk and Uk 2016), little else is known about the freshwater molluscs of the lake. Although poorly studied, freshwater molluscs in the Tonle Sap basin, as with the fauna in the surrounding Indo-Burmese region, occupy a range of habitats and perform vital ecological roles, interacting with other fauna (Köhler et al. 2012). For its size, Cambodia is the most speciose country in East and Southeast Asia for unionid bivalves; although the data on bivalves in the country requires updating (Zieritz et al. 2018). The most comprehensive checklist of freshwater molluscs for Cambodia was based on material collected almost half a century ago from the main stem of the Mekong River, and did not include the Tonle Sap ecosystem (Brandt and Temcharoen 1971). Recent surveys and ecological stud- ies in the fresh waters of Cambodia have targeted various taxa, and when molluscs were included, all were, at most, identified only to genera (Vongsombath et al. 2009; Ohtaka et al. 2011; Sor et al. 2017). In addition, there has been interest in economically-important molluscs, includ- ing freshwater apple snails (Ampullariidae) and Asian clams (Cyrenidae) (Ngor et al. 2014, 2016, 2018d). The composition of ampullariids in the Tonle Sap Lake appears to vary seasonally, probably owing to the vastly different climatic and hydro- logical conditions (Ngor et al. 2018a, d). There is also growing concern about the economic and ecological impacts of invasive apple snails, Pomacea species, which were not established in Cambodia two decades ago (Cowie 1995), but are now spreading rapidly, including in the Tonle Sap Lake (Ngor et al. 2014; Khay et al. 2018; Ngor PB, pers. obs.). The threat of invasive species, in addition to climate change, flow modification within the Mekong River basin, overharvesting, pollu- tion, and land use change, would most probably impact the ecosystem and biodi- versity of the Tonle Sap Lake (Arias et al. 2014, 2019; McCann et al. 2016; Ngor et al. 2018a, b, d; Uk et al. 2018). Freshwater molluscs of Tonle Sap basin 109 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, -al lowing non-native species to establish and spread unnoticed (e.g., Ng et al. 2015, 2018). 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 pro- vide 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. Materials and methods Historical information 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 (IUCN 2018), MolluscaBase (2019), MUS- SEL Project (Graf and Cummings 2019), Google and Web of Science using key- words (Cambodia, Cambodge, freshwater, fluviatiles, mollusc, mollusk, mollusques, clam, bivalve, snail, gastropod) or applying relevant filters. Major references included Mabille and Le Mesle (1866), Crosse and Fischer (1876), Morlet (1889), Fischer and Dautzenberg (1904), Brandt and Temcharoen (1971), and Brandt (1974). Only original descriptions and key references, including first record in Cambodia, if appli- cable, Brandt (1974), and major revisionary work, are listed in the annotated species checklist. Historical records from the Tonle Sap basin were noted based on specific mentions of ‘Tonle Sap Lake’ or ‘grand lac’, and localities within the provinces around the Lake, i.e., Kampong Chhnang, Pursat, Battambang, Banteay Meanchey, Siem Reap and Kampong Thom. Sampling 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 Prov- 110 Ting Hui Ng et al. / ZooKeys 958: 107–141 (2020) ince), 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. 1, Table 1). Specimens were collected by hand and with long-handled nets. Some specimens were also purchased from local markets. Species identification All the specimens were identified to genus or species level based on shell characteris- tics by referring to the historical literature with original species descriptions, Brandt (1974), Morton and Dinesen (2010), Jeratthitikul