A new genus of river snails, Bakyietaia (Mollusca, Viviparidae), from South China and the Indochinese Peninsula
Abstract
A new genus of the river snail family Viviparidae, Bakyietaia Zhang, Yen & von Rintelen gen. nov., is described based on a comprehensive study of its morphology, anatomy, mtDNA (COI), ecology and distribution. This genus is widely distributed in South China and the Indochinese Peninsula. It can be distinguished from all other viviparid genera, especially Angulyagra, Sinotaia and Anulotaia, by characters of the columellar lip of the shell, the operculum and the marginal teeth of radula. Based on an integrative taxonomic approach, 17 species are assigned to this new genus and described here, including 7 new combinations and 10 new species: Bakyietaia subcostata gen. et comb. nov., B. polyzonata gen. et comb. nov., B. boettgeri gen. et comb. nov., B. duchieri gen. et comb. nov., B. guangdungensis gen. et comb. nov., B. mutica gen. et comb. nov., B. wilhelmi gen. et comb. nov., B. chenghuang Zhang, Yen & von Rintelen gen. et sp. nov., B. jingweiae Yen, Zhang & von Rintelen gen. et sp. nov., B. liusanjieae Zhang, Yen & von Rintelen gen. et sp. nov., B. avisvenatoris Yen, Zhang & von Rintelen gen. et sp. nov., B. liangzhuorum Yen, Zhang & von Rintelen gen. et sp. nov., B. luikongi Yen, Zhang & von Rintelen gen. et sp. nov., B. fontinalis Zhang, Yen & von Rintelen gen. et sp. nov., B. naiadica Zhang, Yen & von Rintelen gen. et sp. nov., B. luuemxlong Zhang, Yen & von Rintelen gen. et sp. nov. and B. indrapura Zhang, Yen & von Rintelen gen. et sp. nov. The monophyly of all morphospecies is supported by the COI phylogeny. Radular characters, especially the number of cusps of the outer marginal teeth, are considered valuable in the classification of species of Bakyietaia with similar shells.
References
Annandale N. 1921. The aquatic and amphibious Mollusca of Manipur. The Prosobranchia. Records of the Indian Museum 22: 538–564. https://doi.org/10.5962/bhl.part.1480
Annandale T.N. & Seymour Sewell R.B. 1920. Progress report on a Survey of the freshwater gastropod molluscs of the Indian Empire and of their trematode parasites. Indian Journal of Medical Research 8: 93–124.
Benson W.H. 1836. Descriptive catalogue of a collection of land and fresh-water shells, chiefly contained in the Museum of the Asiatic Society. Journal of the Asiatic Society of Bengal 5: 741–750. Available from https://www.biodiversitylibrary.org/page/37188918 [accessed 9 Jun. 2023].
Chan S.Y. 1996. Some freshwater gastropods of Singapore. Of Sea and Shore 18: 184–187.
Clements R., Koh L.P., Lee T.M., Meier R. & Li D. 2006. Importance of reservoirs for the conservation of freshwater molluscs in a tropical urban landscape. Biological Conservation 128: 136–146. https://doi.org/10.1016/j.biocon.2005.09.023
Dautzenberg P. & Fischer H. 1908. Contribution à la faune malacologique de l’Indo-Chine. Journal de Conchyliologie 54: 145–226. Available from https://www.biodiversitylibrary.org/page/16298507 [accessed 9 Jun. 2023].
Do T.A., Do V.K., Do A D. 2014. The species composition, distribution, biomass of mollusk (Class: Gastropoda, Bivalvia, Cephalopoda) in coral reef in 19 surveyed islands in Vietnam. Vietnam Journal of Marine Science and Technology 14 (4): 358–367. https://doi.org/10.15625/1859-3097/14/4/5822
Fischer H. 1898. Notes sur la faune du Haut Tonkin. III. Liste des mollusques recueillis par le Dr. A. Billet. Bulletin scientifique de la France et de Belgique 28: 310–338. Available from https://www.biodiversitylibrary.org/page/10724680 [accessed 9 Jun. 2023].
Fischer H. & Dautzenberg P. 1904. Catalogue des mollusques terrestres et fluviatiles de ľIndo-Chine orientale cités jusqu’à ce jour. Mission Pavie, Études diverses 3: 390–450.
Frauenfeld G. 1862. Versuch einer Aufzählung der Arten der Gattung Bithynia Lch. und Nematura Bns. nach der kaiserlichen und Cuming’s Sammlung. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 12: 1145–1170. Available from https://www.biodiversitylibrary.org/page/30333254 [accessed 9 Jun. 2023].
Ghilardi R.P., Rodrigues S.C., Simone L.R.L., Carbonaro F.A. & Nava W.R. 2011. Moluscos fósseis do Grupo Bauru. In: Souza Carvalho I.D., Kumar Srivastava N., Strohschoen Jr. O., Cunha Lana C. (eds) Paleontologia: Cenários de Vida Vol. 4: 239–250. Interciência, Rio de Janeiro.
Gredler V.M. 1889. Zur Conchylien-Fauna von China. XIV. Stück. Nachrichtsblatt der Deutschen malakozoologischen Gesellschaft 21: 155–163. Available from https://www.biodiversitylibrary.org/page/15599998 [accessed 9 Jun. 2023].
Griffith E. & Pidgeon E. 1833–1834. The Mollusca and Radiata, Vol. 12. In: Griffith E. 1824−1835. The Animal Kingdom Arranged in Conformity with its Organization, by the Baron Cuvier. Whittaker and Co., London. https://doi.org/10.5962/bhl.title.45021
Haas F. 1939. Malacological notes. Zoological Series of Field Museum of Natural History 24: 93–103. Available from https://www.biodiversitylibrary.org/page/2772848 [accessed 9 Jun. 2023].
Heude P.M. 1882–1890. Notes sur les mollusques terrestres de la Vallée du Fleuve Bleu. Mémoires concernant l’histoire naturelle de l’Empire chinois 1: 1–177. https://doi.org/10.5962/bhl.title.50365
Hilgers L., Hartmann S., Pfaender J., Lentge-Maaß N., Marwoto R.M., von Rintelen T. & Hofreiter M. 2022. Evolutionary divergence and radula diversification in two ecomorphs from an adaptive radiation of freshwater snails. Genes 13: 1029. https://doi.org/10.3390/genes13061029
Hirano T., Saito T., Tsunamoto Y., Koseki J., Prozorova L., Do V.T., Matsuoka K., Nakai K., Suyama Y. & Chiba S. 2019a. Role of ancient lakes in genetic and phenotypic diversification of freshwater snails. Molecular Ecology 28: 5032–5051. https://doi.org/10.1111/mec.15272
Hirano T., Saito T., Tsunamoto Y., Koseki J., Ye B., Do V.T., Miura O., Suyama Y. & Chiba S. 2019b. Enigmatic incongruence between mtDNA and nDNA revealed by multi-locus phylogenomic analyses in freshwater snails. Scientific Reports 9: e6223. https://doi.org/10.1038/s41598-019-42682-0
Hirano T., Saito T., Ito S., Ye B., Linscott T.M., Do V.T., Dong Z. & Chiba S. 2023. Phylogenomic analyses reveal incongruences between divergence times and fossil records of freshwater snails in East Asia. Molecular Phylogenetics and Evolution 182: 107728. https://doi.org/10.1016/j.ympev.2023.107728
Johnson D.S. 1973. Freshwater life. In: Chuang S.H. (ed.) Animal Life and Nature in Singapore. Singapore University Press, Singapore.
Kobelt W. 1906–1909. Die Gattung Paludina Lam. (Vivipara Montfort) in Abbildungen nach der Natur mit Beschreibungen. Sytematisches Conchylien-Cabinet von Martini und Chemnitz, Neue Folge Band 1, Abt. 21a. Bauer & Raspe, Nürnberg. https://doi.org/10.5962/bhl.title.119758
Kozlov A.M., Darriba D., Flouri T., Morel B. & Stamatakis A. 2019. RAxML-NG: A fast, scalable and user-friendly tool for maximum likelihood phylogenetic inference. Bioinformatics 35: 4453–4455. https://doi.org/10.1093/bioinformatics/btz305
Kumar S., Stecher G., Li M., Knyaz C. & Tamura K. 2018. MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35: 1547–1549. https://doi.org/10.1093/molbev/msy096
Liu Y.Y., Zhang W.Z. & Wang Y.X. 1995. Distribution of the Viviparidae from China (Mollusk: Gastropoda). Transactions of the Chinese Society of Malacology 5–6: 8–16. [In Chinese.]
Maassen W.J.M. 2001. A Preliminary Checklist of the Non-marine Mollusks of Malaysia: “A hand list”. De Kreukel Extra Editie.
Madsen H. & Hung N.M. 2014. An overview of freshwater snails in Asia with main focus on Vietnam. Acta Tropica 140: 105–17. https://doi.org/10.1016/j.actatropica.2014.08.005
MolluscaBase eds. 2023. MolluscaBase. Viviparidae J.E. Gray, 1847. Available from https://www.molluscabase.org/aphia.php?p=taxdetails&id=154000 [accessed 9 Jun. 2023].
Ng P.K.L. 1992. A Guide to Freshwater Life in Singapore. Science Centre Singapore, Singapore.
Ng P.K.L., Chou L.M. & Lam T.J. 1993. The status and impact of introduced freshwater animals in Singapore. Biological Conservation 64: 19–24. https://doi.org/10.1016/0006-3207(93)90379-F
Ng T.H., Tan S.K. & Yeo D.C. 2014. The taxonomy, distribution and introduction history of the earliest reported alien freshwater mollusc in Singapore — Sinotaia guangdungensis (Gastropoda: Viviparidae). Malacologia 57: 401–408. https://doi.org/10.4002/040.057.0211
Ng T.H., Dulipat J., Foon J.K., Lopes-Lima M., Zieritz A. & Liew T.S. 2017. A preliminary checklist of the freshwater snails of Sabah (Malaysian Borneo) deposited in the BORNEENSIS collection, Universiti Malaysia Sabah. ZooKeys 673: 105–123. https://doi.org/10.3897/zookeys.673.12544
Palmieri M., Palmeiri J. & Sullivan J. 1980. A chemical analysis of the habitat of nine commonly occurring Malaysia freshwater snails. The Malayan Nature Journal 34: 1–7.
Philippi R.A. 1842–1850. Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien unter Mithülfe mehrerer deutscher Conchyliologen. T. Fischer, Cassel. https://doi.org/10.5962/bhl.title.10589
Prashad B. 1928. Recent and fossil Viviparidae. A study in distribution, evolution and palaeogeography. Memoirs of the Indian Museum 8: 153–253.
Preece R.C., White T.S., Raheem D.C., Ketchum H., Ablett J., Taylor H., Webb K. & Naggs F. 2022. William Benson and the golden age of malacology in British India. Biography, illustrated catalogue and evaluation of his molluscan types. Tropical Natural History Supplement 6: 1–434. https://doi.org/10.58837/tnh.22.6.257073
Preston H.B. 1915. Freshwater Gastropoda & Pelecypoda. In: Blanford W.T. (ed.) The Fauna of British India, Including Ceylon and Burma. Mollusca Vol. 4. Taylor & Francis, London. https://doi.org/10.5962/bhl.title.13091
Puillandre N., Brouillet S. & Achaz G. 2020. ASAP: Assemble Species by Automatic Partitioning. Molecular Ecology Resources 21: 609–620. https://doi.org/10.1111/1755-0998.13281
Qian Z.X., Fang Y.F. & He J. 2014. A conchological review of Bellamyinae (Gastropoda: Viviparidae) of China. Shell Discoveries 1: 3–12.
Rao H.S. 1925. On the comparative anatomy of Oriental Viviparidae. Records of the Zoological Survey of India 27: 129–135. https://doi.org/10.26515/rzsi/v27/i2/1925/163463
Rao H.S. 1931. Angulyagra oxytropis (Benson) nom. nov. – A correction. Records of the Zoological Survey of India 33: 301. https://doi.org/10.26515/rzsi/v33/i3/1931/162505
Richter R. 2015. Die Evolution und Biogeographie der südostasiatischen Sumpfdeckelschnecken (Viviparidae): ein molekularer und morphologischer Ansatz. PhD thesis, Humboldt Universität, Berlin.
Ronquist F., Teslenko M., Van Der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
Schultheiß R., Wilke T., Jørgensen A. & Albrecht C. 2011. The birth of an endemic species flock: Demographic history of the Bellamya group (Gastropoda, Viviparidae) in Lake Malawi. Biological Journal of the Linnean Society 102: 130–143. https://doi.org/10.1111/j.1095-8312.2010.01574.x
Schultheiß R., Van Bocxlaer B., Riedel F., von Rintelen T. & Albrecht C. 2014. Disjunct distributions of freshwater snails testify to a central role of the Congo system in shaping biogeographical patterns in Africa. BMC Evolutionary Biology 14: 1–2. https://doi.org/10.1186/1471-2148-14-42
Sengupta M.E., Kristensen T.K., Madsen H. & Jørgensen A. 2009. Molecular phylogenetic investigations of the Viviparidae (Gastropoda: Caenogastropoda) in the lakes of the Rift Valley area of Africa. Molecular Phylogenetics and Evolution 52: 797–805. https://doi.org/10.1016/j.ympev.2009.05.007
Shea M. 1994. The Chinese viviparid snail Bellamya heudei guangdungensis (Kobelt, 1906) in Australia (Prosobranchia: Viviparidae). Molluscan Research 15: 3–11. https://doi.org/10.1080/13235818.1994.10673652
Sivan N., Heller J. & Van Damme D. 2006. Fossil Viviparidae (Mollusca: Gastropoda) of the Levant. Journal of Conchology 39: 207–219.
Starobogatov Y.I. 1970. Fauna mollyuskov i zoogeograficheskoe raionirovanie kontinental’nykh vodoemov zemnogo shara. Nauka, Leningrad.
Stelbrink B., von Rintelen T., Albrecht C., Clewing C. & Naga P.O. 2019. Forgotten for decades: Lake Lanao and the genetic assessment of its mollusc diversity. Hydrobiologia 843: 31–49. https://doi.org/10.1007/s10750-018-3666-0
Stelbrink B., Richter R., Köhler F., Riedel F., Strong E.E., van Bocxlaer B., Albrecht C., Hauffe T., Page T.J., Aldridge D.C., Bogan A.E., Du L.N., Manuel-Santos M.R., Marwoto R.M., Shirokaya A.A. & von Rintelen T. 2020. Global diversification dynamics since the Jurassic: Low dispersal and habitat-dependent evolution explain hotspots of diversity and shell disparity in river snails (Viviparidae). Systematic Biology 69: 944–961. https://doi.org/10.1093/sysbio/syaa011
Strong E.E., Gargominy O., Ponder W.F. & Bouchet P. 2008. Global diversity of gastropods (Gastropoda; Mollusca) in freshwater. Hydrobiologia 595: 149–166. https://doi.org/10.1007/s10750-007-9012-6
Subba Rao N.V. 1989. Handbook: freshwater molluscs of India. Zoological Survey of India, Calcutta.
Tan S.K. & Woo H.P.M. 2010. A Preliminary Checklist of the Molluscs of Singapore. Raffles Museum of Biodiversity Research, Singapore.
Tan S.K., Chan S.Y. & Clements G.R. 2012. A Guide to Snails and other Non-marine Mollucs of Singapore. Science Centre Singapore, Singapore.
Thanh D.N., Hai H.T. & Cuong D. 2004. Snail species of Viviparidae in Vietnam. Academia Journal of Biology 26: 1–5.
Tracey S., Todd J.A. & Erwin D.H. 1993. Mollusca: Gastropoda. In: Benton M.J. (ed.) The Fossil Record 2: 131–167. Chapman and Hall, London.
von Rintelen T., Wilson A.B., Meyer A. & Glaubrecht M. 2004. Escalation and trophic specialization drive adaptive radiation of freshwater gastropods in ancient lakes on Sulawesi, Indonesia. Proceedings of the Royal Society of London. Series B: Biological Sciences 271: 2541–2549. https://doi.org/10.1098/rspb.2004.2842
Winnepenninckx B., Backeljau T. & De Wachter R. 1993. Extraction of high molecular weight DNA from molluscs. Trends in Genetics 9: e407. https://doi.org/10.1016/0168-9525(93)90102-N
Ye B., Saito T., Hirano T., Dong Z., Do V.T. & Chiba S. 2020. Human-geographic effects on variations in the population genetics of Sinotaia quadrata (Gastropoda: Viviparidae) that historically migrated from continental East Asia to Japan. Ecology and Evolution 10: 8055–8072. https://doi.org/10.1002/ece3.6456
Ye B., Hirano T., Saito T., Dong Z., Do V.T. & Chiba S. 2021. Molecular and morphological evidence for a unified, inclusive Sinotaia quadrata (Caenogastropoda: Viviparidae: Bellamyinae). Journal of Molluscan Studies 87: eyab013. https://doi.org/10.1093/mollus/eyab013
Yen T.C. 1938 Notes on the gastropod fauna of Szechwan Province. Mitteilungen aus dem Zoologischen Museum in Berlin 23: 438–457.
Yen T.C. 1939. Die chinesischen Land- und Süsswasser-Gastropoden des Natur-Museums Senckenberg. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 444: 1–233.
Yen T.C. 1943. A preliminary revision of the recent species of Chinese Viviparidae. The Nautilus 56: 124–130. Available from https://www.biodiversitylibrary.org/page/8520239 [accessed 9 Jun. 2023].
Yeo D.C.J. & Chia C.S.W. 2010. Introduced species in Singapore: An overview. COSMOS 6: 23–37. https://doi.org/10.1142/S0219607710000486
Zhang L.J. & von Rintelen T. 2021. The neglected operculum: A revision of the opercular characters in river snails (Caenogastropoda: Viviparidae). Journal of Molluscan Studies 87: eyab008. https://doi.org/10.1093/mollus/eyab008
Zhang L.J., Chen S.C., Yang L.T., Jin L. & Köhler F. 2015. Systematic revision of the freshwater snail Margarya Nevill, 1877 (Mollusca: Viviparidae) endemic to the ancient lakes of Yunnan, China, with description of new taxa. Zoological Journal of the Linnean Society 174: 760–800. https://doi.org/10.1111/zoj.12260
Zhang L.J., Du L.N. & von Rintelen T. 2023. A new genus of river snails, Dalipaludina (Gastropoda, Viviparidae), endemic to the Yunnan Plateau of SW China. Zoosystematics and Evolution 99: 285–297. https://doi.org/10.3897/zse.99.102586
Zhu X.G. 1994. Jurassic-Cretaceous nonmarine gastropods from northern Xinjiang. Acta Palaeontologica Sinica 33: 85–105.
Zilch A. 1938. Das Vorkommen von Buliminopsis incerta (Pfr.) auf Formosa und in China. Archiv für Molluskenkunde 7: 3.
Zilch A. 1955. Die Typen und Typoide des Natur-Museums Senckenberg, 14: Mollusca, Viviparidae. Archiv für Molluskenkunde 84: 45–87.
Zilch A. 1974. Zur Geschichte der deutschen Malakozoologie, XIV. Vinzenz Gredler und die Erforschung der Weichtiere Chinas durch Franziskaner aus Tirol. Archiv für Molluskenkunde 104: 171–228.
Copyright (c) 2025 Le-Jia Zhang, Yu-Hsiu Yen, Zhe-Yu Chen, Li-Na Du, Ting Hui Ng, Thomas von Rintelen
This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
