A review of Chamalycaeus, Cycloryx and Dicharax species of the Himalaya and Myanmar and seven new species of Alycaeinae from the Blue Mountain, Mizoram, India (Gastropoda, Caenogastropoda, Cyclophoridae)
Abstract
In this paper, we revise the genera Chamalycaeus, Cycloryx and Dicharax of the Himalayas and Myanmar. Another aim of this work was to identify the 13 species of Alycaeinae collected on the Blue Mountain, Mizoram State, northeastern India. Out of the 13 species from the Blue Mountain, seven are new and are described here: Chamalycaeus mizoramensis Páll-Gergely & Aravind sp. nov., Chamalycaeus reflectus Páll-Gergely & Aravind sp. nov., Cycloryx rohiniae Páll-Gergely & Aravind sp. nov., Dicharax praeda Páll-Gergely & Aravind sp. nov., D. semivivus Páll-Gergely & Aravind sp. nov., D. umashaankeri Páll-Gergely & Aravind sp. nov., D. ganeshaiahi Páll-Gergely & Aravind sp. nov. Altogether, 109 species are recognized, and all are figured. Furthermore, 37 new synonyms are reported, 32 of which were described by Godwin-Austen.
References
Aravind N.A. & Páll-Gergely B. 2018. Dicharax (?) bawai n. sp from southern India (Gastropoda: Cyclophoroidea: Alycaeidae). Archiv für Molluskenkunde 147 (1): 55–62. https://doi.org/10.1127/arch.moll/147/055-062
Aravind N.A. & Páll-Gergely B. 2023. Two extraordinary alycaeid species from Northeastern India (Gastropoda: Caenogastropoda: Cyclophoroidea). Acta Zoologica Academiae Scientiarum Hungaricae 69 (4): 353–363. https://doi.org/10.17109/AZH.69.4.353.2023
Bavay A. & Dautzenberg P. 1900. Diagnoses de coquilles nouvelles de l’Indo-Chine. Journal de Conchyliologie 48: 108–122. Available from https://www.biodiversitylibrary.org/page/25146858 [accessed 1 May 2025].
Bavay A. & Dautzenberg P. 1904. Description de coquilles nouvelles de l’Indo-Chine (3e suite). Journal de Conchyliologie 51 (3): 201–236. Available from https://www.biodiversitylibrary.org/page/16311391 [accessed 1 May 2025].
Bavay A. & Dautzenberg P. 1912. Description de coquilles nouvelles de l’Indo-Chine (7e suite). Journal de Conchyliologie 60 (1): 1–54. Available from https://www.biodiversitylibrary.org/page/16358455 [accessed 1 May 2025].
Benson W.H. 1851. Geographical notices, and characters of fourteen new species of Cyclostoma, from the East Indies. Annals and Magazine of Natural History Series 2 8 (45): 184–195. https://doi.org/10.1080/03745486109496203
Benson W.H. 1853. Additional character of the shell of the cyclostomatous genus Alycæus of Gray, with descriptions of its animal inhabitant, –of a fourth species,– and of other new Indian Cyclostomata; also, Remarks on an unrecorded Character in Diplommatina. The Annals and Magazine of Natural History Series 2 9: 283–287. https://doi.org/10.1080/03745485609495767
Benson W.H. 1856. Characters of seventeen new forms of the Cyclostomacea from the British provinces of Burmah, collected by W. Theobald, jun., Esq. The Annals and Magazine of Natural History Series 2 17: 225–233. https://doi.org/10.1080/00222935608697501
Benson W.H. 1857. Characters of Streptaulus a new genus and several species of the Cyclostomacea from Sikkim, the Khasi Hills Ava and Pegu. The Annals and Magazine of Natural History Series 2 19: 201–211. https://doi.org/10.1080/00222935708681840
Benson W.H. 1859. A sectional distribution of the genus Alycæus, Gray, with characters of six new species and of other Cyclostomidæ collected at Darjiling by W.T. Blanford, Esq., Geol. Survey. The Annals and Magazine of Natural History Series 3 3: 176–184. Available from https://www.biodiversitylibrary.org/page/2317897 [accessed 1 May 2025].
Bhosale A., Thackeray T., Yadav O. & Khandekar A. 2025. A new micro-snail species of the genus Dicharax Kobelt & Möllendorff, 1900 (Caenogastropoda: Cyclophoridae: Alycaeinae), from sacred groves in the northern Western Ghats, India. Journal of Conchology 45 (3): 488–497. https://doi.org/10.61733/jconch/4543
Blanford H.F. 1871. On some undescribed species of Camptoceras and other land shells. The Journal of the Asiatic Society of Bengal 40: 39–45. Available from https://www.biodiversitylibrary.org/page/35630849 [accessed 1 May 2025].
Blanford W.T. 1862. Contributions to Indian Malacology, No. III. Descriptions of new operculated land-shells from Pegu, Arakan and the Khasi Hills. Journal of the Asiatic Society of Bengal 31: 135–144. Available from https://www.biodiversitylibrary.org/page/56711841 [accessed 1 May 2025].
Blanford W.T. 1863. Contributions to Indian Malacology. No. IV. Descriptions of new land shells from Ava, and other parts of Burma. Journal of the Asiatic Society of Bengal 32: 319–327. Available from https://www.biodiversitylibrary.org/page/37486655 [accessed 1 May 2025].
Blanford W.T. 1864. On the classification of the Cyclostomacea of Eastern Asia. The Annals and Magazine of Natural History Series 3 13: 441–465. https://doi.org/10.1080/00222936408681635
Blanford W.T. 1865. Contribution to Indian Malacology, No. V. Descriptions of new land shells from Arakan, Pegu, and Ava; with notes on the distribution of described species. Journal of the Asiatic Society of Bengal 34 (2): 66–105. Available from https://www.biodiversitylibrary.org/page/41152769 [accessed 1 May 2025].
Chen D.-N. & Zhang G.-Q. 2001. Five new species of land shells from the Tibet Autonomous Region, China. Acta Zootaxonomica Sinica 26 (1): 184–190.
Chen Z.-Y. & Lin L.-W. 2021. Notes on the genus Messageria Bavay & Dautzenberg, 1904, with descriptions of a new species and a new subspecies (Gastropoda: Caenogastropoda: Cyclophoroidea: Alycaeidae). Zootaxa 5067 (1): 129–134. https://doi.org/10.11646/zootaxa.5067.1.9
Egorov R. 2019. Notes on radula of Boucardicus leopardus Emberton, 2002 (Gastropoda: Cyclophoroidea: Alycaeidae) and systematic position of the genus Boucardicus Fischer-Piette et Bedoucha, 1965. Ruthenica 29 (4): 181–184. https://doi.org/10.35885/ruthenica.2019.29(4).2
Emberton K.C. 2002. The genus Boucardicus, a Madagascan endemic (Gastropoda: Cyclophoridae: Alycaeinae). Archiv für Molluskenkunde 130 (1–2): 1–199. https://doi.org/10.1127/arch.moll/130/2002/1
Fischer-Piette E. & Bedoucha J. 1965. Mollusques terrestres operculés de Madagascar. Mémoires du Muséum national d’Histoire naturelle, Series A, Zoologie 33 (2): 49–91. https://doi.org/10.5281/zenodo.16003273
Foon J.K. & Liew T-S. 2017. A review of the land snail genus Alycaeus (Gastropoda, Alycaeidae) in Peninsular Malaysia. ZooKeys 692: 1–81. https://doi.org/10.3897/zookeys.692.14706
Gittenberger E., Choki Gyeltshen, Kezang Tobgay & Sherub Sherub 2022. The genera Dioryx and Cycloryx (Gastropoda, Caenogastropoda, Alycaeidae) in Bhutan, with a description of four new species. Basteria 86 (2): 72–82.
Gittenberger E., Choki Gyeltshen, Kezang Tobgay & Sherub Sherub S. 2024. The genera Metalycaeus and Dicharax (Caenogastropoda, Cyclophoroidea, Alycaeidae) in Bhutan, with the description of 16 new species. Basteria 88 (2): 187–219.
Godwin-Austen H.H. 1871. Descriptions of the species of Alycæinæ known to inhabit the Khasi Hill ranges. Journal of the Asiatic Society of Bengal 40: 87–93. Available from https://www.biodiversitylibrary.org/page/35630903 [accessed 1 May 2025].
Godwin-Austen H.H. 1874. Descriptions of nine species of Alycæinæ from Assam and the Naga Hills. Journal of the Asiatic Society of Bengal 43 (2): 145–150. Available from https://www.biodiversitylibrary.org/page/35549500 [accessed 1 May 2025].
Godwin-Austen H.H. 1875. Descriptions of new operculated landshells belonging to the genera Craspedotropis, Alycæus, and Diplommatina, from the Nágá Hills and Assam. Journal of the Asiatic Society of Bengal 44 (2): 7–10. Available from https://www.biodiversitylibrary.org/page/37191980 [accessed 1 May 2025].
Godwin-Austen H.H. 1876. On the Cyclostomacea of the Dafla Hills, Assam. Journal of the Asiatic Society of Bengal 45 (2): 171–184. Available from https://www.biodiversitylibrary.org/page/37118189 [accessed 1 May 2025].
Godwin-Austen H.H. 1882–1920. Land and Freshwater Mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon and other islands of the Indian Ocean; Supplementary to Messrs. Theobald and Hanley’s Conchologica Indica. Taylor and Francis, London, Vol. 1 (1): [i–iv] 1–18, pls 1–4 [February 1882]; 1 (2): 19–66, pls 5–12 [July 1882]; 1 (3): 67–94, pls 13–21 [January 1883]; 1 (4): pls 22–42 [September 1883], 95–164 [October 1883]; 1 (5): pls 43–51 [June 1884], 165–206 [May 1886]; 1 (6): pls 52–62 [September 1887], 207–257 [April 1888]. Volume 2 (7): 1–46, pls 63–69 [October 1897]; 2 (8): 47–86, pls 70–82 [January 1898]; 2 (9): 87–146, pls 83–100 [November 1899]; 2 (10): 147–238, pls 101–117 [April 1907]; 2 (11): 239–310, pls 118–132 [March 1910]; 2 (12): 311–442, pls 133–158 [December 1914]. Volume 3: 1–65, pls 159–165.
Godwin-Austen H.H. 1893. On some new species of the land-molluscan genus Alycaeus from the Khasi and Naga Hill Country, Assam, Munipur, and the Ruby Mine District, Upper Burma; and on one species from the Nicobars. Proceedings of the Zoological Society of London 61 (3): 592–595. Available from https://www.biodiversitylibrary.org/page/30981683 [accessed 1 May 2025].
Godwin-Austen H.H. 1922. On a new Alycaeus from the Khasi Hills. Records of the Indian Museum 24 (3): 365. https://doi.org/10.26515/rzsi/v24/i3/1922/162725
Gray J.E. 1847. A list of the genera of recent Mollusca, their synonyma and types. Proceedings of the Zoological Society of London 15: 129–219. Available from https://www.biodiversitylibrary.org/page/12862913 [accessed 1 May 2025].
Gude G.K. 1921. The Fauna of British India including Ceylon and Burma. Mollusca. III. Land operculates (Cyclophoridae, Truncatellidae, Assimineidae, Helicinidae). Taylor and Francis, London. https://doi.org/10.5962/bhl.title.12890
Hanley S.C.T. & Theobald W. 1870–1876. Conchologia Indica; Being Illustrations of the Land and Freshwater Shells of British India. L. Reeve & Co., London. https://doi.org/10.5962/bhl.title.14456
Jirapatrasilp P., Páll-Gergely B., Sutcharit C. & Tongkerd P. 2021. The operculate micro land snail genus Dicharax Kobelt & Möllendorff, 1900 (Caenogastropoda, Alycaeidae) in Thailand, with description of new species. Zoosystematics and Evolution 97 (1): 1–20. https://doi.org/10.3897/zse.97.59143
Kerney M.P. & Cameron R.A.D. 1979. A Field Guide to the Land Snails of Britain and North-west Europe. Collins, London.
Kobelt W. 1902. Das Tierreich. Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen. In Verbindung mit der Deutschen Zoologischen Gesellschaft herausgegeben von der Königlich Preussischen Akademie der Wissenschaften zu Berlin. Lfg 16. Mollusca: Cyclophoridae. R. Friedländer und Sohn, Berlin. Available from https://www.biodiversitylibrary.org/page/1000211 [accessed 1 May 2025].
Kobelt W. & Möllendorff O. von 1900. Zur Systematik der Pneumonomen. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 32: 186. Available from https://www.biodiversitylibrary.org/page/15598533 [accessed 1 May 2025].
Minato H. 1988. A Systematic and Bibliographic List of the Japanese Land Snails. Shirahama, H. Minato.
Möllendorff O.F. von 1886. Materialien zur Fauna for China. Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 13: 156–210. Available from https://www.biodiversitylibrary.org/page/16355855 [accessed 1 May 2025].
Möllendorff O.F. von. 1890. Die Landschnecken-Fauna der Insel Cebu. Bericht über die Senckenbergische Naturforschende Gesellschaft in Frankfurt-am-Main 1890: 189–292. Available from https://www.biodiversitylibrary.org/page/9003882 [accessed 1 May 2025].
Möllendorff O. von 1897a. Diagnosen neuer und kritischer Landdeckelschnecken. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 29: 31–45. Available from https://www.biodiversitylibrary.org/page/28228223 [accessed 1 May 2025].
Möllendorff O. von 1897b. Neue Landschnecken von Java. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 29: 89–97. Available from https://www.biodiversitylibrary.org/page/28228319 [accessed 1 May 2025].
Möllendorff O. von 1902. Binnenmollusken aus Hinterindien. 1. Landschnecken von Kelantan, Ostküste der Halbinsel Malacca. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 34: 135–149. Available from https://www.biodiversitylibrary.org/page/15598858 [accessed 1 May 2025].
MolluscaBase eds 2025. MolluscaBase. Available from https://www.molluscabase.org [accessed 1 May 2025]. https://doi.org/10.14284/448
Nevill G. 1878. Hand List of Mollusca in the Indian Museum, Calcutta. Part I. Gastropoda. Pulmonata and Prosobranchia–Neurobranchia. Order of the Trustees, Calcutta. https://doi.org/10.5962/bhl.title.11957
Nevill G. 1881. New or little-known Mollusca of the Indo-Malayan fauna. The Journal of the Asiatic Society of Bengal 50 (2, art. 3): 125–167. Available from https://www.biodiversitylibrary.org/page/35598060 [accessed 1 May 2025].
Páll-Gergely B. 2017. A new species of Alycaeidae, Pincerna yanseni n. sp. from Sumatra, with the resurrection of the genus Pincerna Preston, 1907 (Gastropoda: Cyclophoroidea). Raffles Bulletin of Zoology 65: 213–219.
Páll-Gergely B. 2023. Revision of the Alycaeidae of China, Laos and Vietnam (Gastropoda: Cyclophoroidea) II: The genera Alycaeus and Pincerna. Zootaxa 5249 (2): 253–276. https://doi.org/10.11646/zootaxa.5249.2.4
Páll-Gergely B. & Asami T. 2017. A new species of Metalycaeus hitherto undistinguished from M. vinctus (Pilsbry, 1902). Venus 75 (1–4): 1–16.
Páll-Gergely B. & Auffenberg K. 2019. A review of the Alycaeidae of the Philippines with descriptions of new species and subspecies (Gastropoda: Caenogastropoda: Cyclophoroidea). Molluscan Research 39 (4): 377–389. https://doi.org/10.1080/13235818.2019.1638541
Páll-Gergely B. & Hunyadi A. 2022. Two new species of Dicharax Kobelt & Möllendorff, 1900 from Myanmar and one from Thailand (Gastropoda: Caenogastropoda: Alycaeidae). Acta Phytopathologica et Entomologica Hungarica 57 (2): 94–105. https://doi.org/10.1556/038.2022.00165
Páll-Gergely B., Fehér Z., Hunyadi A. & Asami T. 2015. Revision of the genus Pseudopomatias and its relatives (Gastropoda: Cyclophoroidea: Pupinidae). Zootaxa 3937 (1): 1–49. https://doi.org/10.11646/zootaxa.3937.1.1
Páll-Gergely B., Naggs F. & Asami T. 2016. Novel shell device for gas exchange in an operculated land snail. Biology Letters 12: 20160151. https://doi.org/10.1098/rsbl.2016.0151
Páll-Gergely B., Hunyadi A., Đỗ Đ.S., Naggs F, & Asami T. 2017. Revision of the Alycaeidae of China, Laos and Vietnam (Gastropoda: Cyclophoroidea) I: The genera Dicharax and Metalycaeus. Zootaxa 4331 (1): 1–124. https://doi.org/10.11646/zootaxa.4331.1.1
Páll-Gergely B., Sajan S., Tripathy B., Meng K., Asami T. & Ablett J.D. 2020. Genus-level revision of the Alycaeidae (Gastropoda, Cyclophoroidea), with an annotated species catalogue. ZooKeys 981: 1–220. https://doi.org/10.3897/zookeys.981.53583
Páll-Gergely B., Hunyadi A., Grego J., Reischütz A. & Auffenberg K. 2021. Nineteen new species of Alycaeidae from Myanmar and Thailand (Gastropoda: Caenogastropoda: Cyclophoroidea). Zootaxa 4973 (1): 1–61. https://doi.org/10.11646/zootaxa.4973.1.1
Páll-Gergely B., Ruthensteiner B., Harl J., Magonyi N.M., Asami T., Krizsik K., Schwaha T. & Fehér Z. 2024. Recurrent evolution of breathing microtunnel system in terrestrial operculate snails (Gastropoda: Caenogastropoda: Cyclophoroidea). Zoological Journal of the Linnean Society 202: zlae158. https://doi.org/10.1093/zoolinnean/zlae158
Pfeiffer L. 1846. Nachtrag zur Revision der Gattung Cyclostoma. Zeitschrift für Malakozoologie 3 (6): 81–87. Available from https://www.biodiversitylibrary.org/page/16291875 [accessed 1 May 2025].
Pilsbry H.A. 1900. Notices of new Japanese land shells. Proceedings of the Academy of Natural Sciences of Philadelphia 52: 381–384. Available from https://www.biodiversitylibrary.org/page/24694261 [accessed 1 May 2025].
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 22 (Supp. 6): 1–434. https://doi.org/10.58837/tnh.22.6.257073
Preston H.B. 1914. Characters of new land and freshwater shells from the Naga Hills, Assam. Proceedings of the Malacological Society of London 11: 19–24. Available from https://www.biodiversitylibrary.org/page/15715517 [accessed 1 May 2025].
Raheem D.C., Taylor, H., Ablett J., Preece R., Aravind N.A. & Naggs F. 2014. A systematic revision of the land snails of the Western Ghats of India. Tropical Natural History 14 (Supp. 4): 1–294. https://doi.org/10.58837/tnh.14.4.103091
Ramakrishna, Mitra S.C. & Dey A. 2010. Annotated Checklist of Indian Land Molluscs. Zoological Survey of India, Kolkata. Available from http://faunaofindia.nic.in/PDFVolumes/occpapers/306/index.pdf [accessed 1 May 2025].
Sowerby G.B. II 1877. Monograph of the genus Alycaeus. In: Conchologia iconica, or Illustrations of the Shells of Molluscous Animals Vol. 20: pls 1–6 and unpaginated text. L.A. Reeve, London. https://doi.org/10.5962/bhl.title.8129
Szekeres M., Grego J., Páll-Gergely B. & Ablett, J.D. 2021. Clausiliidae (Gastropoda: Pulmonata) from India, Myanmar, Pakistan and Sri Lanka in the collection of the Natural History Museum, London. Journal of Conchology 44 (2): 155–187.
Reinhardt O. 1877. Herr Reinhardt legte eine Anzahl japanischer...: 67–70. In: Peters (Director) Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1877: 23–84. Available from https://www.biodiversitylibrary.org/page/8796787 [accessed 1 May 2025].
Rintluanga P. 2009. Mizoram: A Study in Comprehensive Geography. Northern Book Center, New Delhi.
Sajan S., Páll-Gergely B., Tripathy B., Chatterjee P., Chandra K. & Sivakumar K. 2020. Redescription and ecological niche of a land snail Dicharax strangulatus (L. Pfeiffer, 1846) in the Himalaya (Gastropoda: Cyclophoroidea: Alycaeidae). Journal of Conchology 43 (5): 521–530.
Saurin E. 1953. Coquilles nouvelles de l’Indochine. Journal de Conchyliologie 93 (4): 113–120.
Scopoli J.A. 1777. Introductio ad historiam naturalem, sistens genera lapidum, plantarum et animalium hactenus detecta, caracteribus essentialibus donata, in tribus divisa, subinde ad leges naturae. Wolfgangum Gerle, Prague. https://doi.org/10.5962/bhl.title.10827
Theobald W. 1870. Descriptions of some new land shells from the Shan States and Pegu. The Journal of the Asiatic Society of Bengal Part II 39 (4): 395–402. Available from https://www.biodiversitylibrary.org/page/41301998 [accessed 1 May 2025].
Theobald W. & Stoliczka F. 1872. Notes on Barmese and Arakanese land shells, with descriptions of a few species. Journal of the Asiatic Society of Bengal 41 (2): 329–334. Available from https://www.biodiversitylibrary.org/page/37138045 [accessed 1 May 2025].
Thiele J. 1929. Handbuch der systematischen Weichtierkunde, Teil 1. Verlag von Gustav Fischer, Jena.
Yen T.-C. 1939. Die Chinesischen Land-und Süßwasser-Gastropoden des Natur-Museums Senckenberg. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main.
Zilch A. 1957. Die Typen und Typoide der Natur-Museums Senckenberg, 20: Mollusca, Cyclophoridae, Alycaeinae. Archiv für Molluskenkunde 86 (4/6): 141–150.
Copyright (c) 2025 Barna Páll-Gergely, Vukašin Gojšina, Neelavar Ananthram Aravind

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.
