Austrelatus zimmermanni (Gschwendtner, 1934) comb. nov. (Coleoptera: Dytiscidae: Copelatinae) with two new subspecies from Japan, and notes on the species biology and ecology
Abstract
Museum specimens and the newly collected material of the species known so far as Copelatus zimmermanni Gschwendtner, 1934 from Japan were examined, and based on our molecular analysis and morphological study, the species is recognised to belong to the genus Austrelatus Shaverdo, Hájek, Hendrich, Surbakti, Panjaitan & Balke, 2023: Austrelatus zimmermanni (Gschwendtner, 1934) comb. nov. Although showing almost no differences in the shape of the male genitalia, specimens from the Minamidaitô-jima and Miyako-jima islands have a very distinct external morphology from those of other localities and, therefore, are described here as two new subspecies: A. zimmermanni daitoensis Watanabe, Kato & Shaverdo subsp. nov. and A. zimmermanni miyakoensis Watanabe, Kato & Shaverdo subsp. nov., which is also supported by the received molecular data. Their important diagnostic characters are compared with those of the nominotypical subspecies and illustrated. In comparison with the Chinese and Korean material, a taxonomic problem of A. zimmermanni is postulated and discussed. Data on the species ecology and distribution, including its two subspecies, are provided as well as the newly received data on its biology and phenology.
References
Aimoto A. 2018. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from Yamaguchi Prefecture]. Sayabane, New Series (29): 27−28. [In Japanese.]
Aoyagi M. 2022. Family Dytiscidae. In: Matsumura M., Kusui Y., Kohama T., Sasaki T., Aoyagi M. & Yoshitake H. (Eds) Beetles of Okinawa: 117–121. Okinawa Jiji Publishing Co., Naha. [In Japanese.]
Bergsten J. & Miller K.B. 2007. Phylogeny of diving beetles reveals a coevolutionary arms race between the sexes. PloS ONE 2 (6): e522. https://doi.org/10.1371/journal.pone.0000522
Bolger A.M., Lohse M. & Usadel B. 2014. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30 (15): 2114–2120. https://doi.org/10.1093/bioinformatics/btu170
Bryant D. & Moulton V. 2004. Neighbor-net: an agglomerative method for the construction of phylogenetic networks. Molecular Biology and Evolution 21 (2): 255–265. https://doi.org/10.1093/molbev/msh018
Cho Y.B. & Kim D.S. 1998. Insect fauna of some wetlands in Cheju Island, Korea. Bulletin of the KACN 17: 57–74. [In Korean.]
Chung H.Y., Yeom C.M., Kim J.H., Park S., Lee Y.W., Pyo G. & Kim S.H. 2020. Species diversity and community characteristics of benthic macroinvertebrates from irrigation ponds in the western CCZ area, Korea. Korean Journal of Ecology and Environment 53: 173–184. https://doi.org/10.11614/KSL.2020.53.2.173
Colgan D.J., McLauchlan A., Wilson G.D.F., Livingston S.P., Edgecombe G.D., Macaranas J., Cassis G. & Gray M.R. 1998. Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology 46 (5): 419–437. https://doi.org/10.1071/ZO98048
ESK (The Entomological Society of Korea) & KSAE (Korean Society of Applied Entomology) 1994. Check List of Insects from Korea. Kon-Kuk University Press, Seoul.
Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3 (5): 294–299.
Fukagawa G. 2015. Record of water beetles (Noteridae, Dytiscidae, Gyrinidae, Hydrochidae, Hydraenidae and Hydrophilidae), in main land of Nagasaki Prefecture, Japan. Transactions of the Nagasaki Biological Society 76: 24–44. [In Japanese.]
Fukagawa G., Imasaka S., Yamamoto N., Noda M., Abiru N., Matsuo T. & Tanaka K. 2016. A fauna of the beetles (Coleoptera) in Nishisonogi peninsula, Nagasaki Pref., Japan. Koganemushi 81: 1–152. [In Japanese.]
Guéorguiev V.B. 1968. Essai de classification des coléoptères Dytiscidae. I. Tribus Copelatini (Colymbetinae). Izvestija na Zoologitjeskija Instituts Musei Sofia 28: 5–45.
Guéorguiev V.B. 1970. Dytiscidae (Coleoptera) nouveaux ou peu connus de l’Asie. Izvestija na Zoologitjeskija Instituts Musei Sofia 32: 253–260.
Hájek J. & Shaverdo H. 2024. Two new species of Austrelatus diving beetles from continental Southeast Asia, linking the distribution area of the genus (Coleoptera: Dytiscidae: Copelatinae). Acta Entomologica Musei Nationalis Pragae 64: 243–247. https://doi.org/10.37520/aemnp.2024.016
Hájek J., Shaverdo H., Hendrich L. & Balke M. 2021. A review of Copelatus diving beetles from the Solomon Islands, reporting the discovery of six new species (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 1023: 81–118. https://doi.org/10.3897/zookeys.1023.61478
Hájek J., Alarie Y., Benetti C.J., Hamada N., Springer M., Hendrich L., Villastrigo A., Torres R.O., Basantes M.S. & Balke M. 2024. Underestimated diversity and range size of diving beetles in tank bromeliads — Coleoptera of ‘hygrofloric’ lifestyle (Dytiscidae). Zoological Journal of the Linnean Society 200: 720–735. https://doi.org/10.1093/zoolinnean/zlad093
Han M.S., Kim M.H., Bang H.S., Na Y.E., Lee D.B. & Kang K.K. 2011. Geographical distribution of diving beetles (Dytiscidae) in Korean paddy ecosystem. Korean Journal of Environmental Agriculture 30: 209−215. https://doi.org/10.5338/KJEA.2011.30.2.209
Hirano T., Kameda Y. & Chiba S. 2014. Phylogeny of the land snails Bradybaena and Phaeohelix (Pulmonata: Bradybaenidae) in Japan. Journal of Molluscan Studies 80 (2): 177−183. https://doi.org/10.1093/mollus/eyu004
Hosoya T. & Tanahashi M. 2017. New record of dytiscid beetle (Coleoptera, Dytiscidae) from Kuchino-shima Island in the Tokara Islands, the Ryukyu Archipelago, Japan. Elytra, New Series 7 (1): 15–16.
Hosoya T., Kiyoshi T. & Kawashimo Y. 2009. Records on aquatic Coleoptera in Takara Island, Tokara Islands, Kagoshima Prefecture, Japan. Coleopterists’ News 165: 5–9. [In Japanese.]
Huson D.H. & Bryant D. 2024. The SplitsTree App: interactive analysis and visualization using phylogenetic trees and networks. Nature Methods 21 (10): 1773−1774. https://doi.org/10.1038/s41592-024-02406-3
Imasaka S., Hosoya T., Kokubu K., Ito R. & Arima K. 2021. [Collecting travelogue in Koshiki-shima Island, part 3 (July 2020)]. Korasana 96: 67–97. [In Japanese.]
Iwasaki I. & Kinoda T. 1995. New records of aquatic insects from Higashi-morokata district, Miyazaki Prefecture (Hemiptera & Coleoptera). In: Hirashima Y. (ed.) Contribution to the Fauna and Flora of Miyazaki Higashimorokata District: New Records and Observations of Noteworthy Insects, Spiders and Plants Based on Systematic and Biologic Studies: 101–111. Miyazaki Municipal University, Miyazaki, Japan.
Jiang Z.Y., Zhao S., Yang Z.Y., Jia F.L. & Hájek J. 2022. A review of Copelatus Erichson, 1832 of Mainland China, with description of ten new species from the japonicus complex (Coleoptera: Dytiscidae: Copelatinae). Zootaxa 5124 (3): 251–295. https://doi.org/10.11646/zootaxa.5124.3.1
Jiang Z.Y., Hájek J., Zheng Y.L., Jia F.L. & Wang B.X. 2025. Two new species of Oriental Copelatus Erichson, 1832, with additional records of the genus from China (Coleoptera: Dytiscidae: Copelatinae). Zootaxa 5632 (1): 42–50. https://doi.org/10.11646/zootaxa.5632.1.2
Jiang Z.Y., Zheng Y.L., Deng Z., Jia F.L. & Wang B.X. 2026. Two new species of Copelatus Erichson, 1832 from urban low mountains of eastern China, with new province records of the other Copelatinae in China (Coleoptera: Dytiscidae). European Journal of Taxonomy 1052: 105–124. https://doi.org/10.5852/ejt.2026.1052.3271
Kaito T. & Toda M. 2016. The biogeographical history of Asian keelback snakes of the genus Hebius (Squamata: Colubridae: Natricinae) in the Ryukyu Archipelago, Japan. Biological Journal of the Linnean Society 118 (2): 187–199. https://doi.org/10.1111/bij.12726
Kalyaanamoorthy S., Minh B.Q., Wong T.K., Von Haeseler A. & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14 (6): 587–589. https://doi.org/10.1038/nmeth.4285
Kamite Y. 2003. [New records of dytiscid species from Minamidaitô-jima Island]. Coleopterists’ News 143: 12. [In Japanese.]
Kamite Y., Aoyagi M. & Kamite N. 2019. Records on aquatic Coleoptera of Aka-jima and Geruma-jima, Okinawa Prefecture, Japan. Sayabane, New Series 36: 21–26. [In Japanese.]
Karlsson Green K., Kovalev A., Svensson E.I. & Gorb S.N. 2013. Male clasping ability, female polymorphism and sexual conflict: fine-scale elytral morphology as a sexually antagonistic adaptation in female diving beetles. Journal of the Royal Society Interface 10: e20130409. https://doi.org/10.1098/rsif.2013.0409
Kato M. & Watanabe K. 2025a. Records of eight aquatic insect species collected from Zamami–jima Island, the Ryukyus, Japan, including six new records (Hemiptera and Coleoptera). Fauna Ryukyuana 74: 1–6. [In Japanese.]
Kato M. & Watanabe K. 2025b. New distributional records of aquatic Coleoptera (Noteridae, Dytiscidae, Hydrophilidae) from Amami-Ôshima and Kakeroma-jima Islands, Kagoshima Prefecture, Japan. Elytra, New Series 15: 153–157.
Lee D.H. & Ahn K.J. 2018. Insect Fauna of Korea, Volume 12, Number 22, Aquatic Coleoptera I, Arthropoda: Insecta: Coleoptera: Dytiscidae. National Institute of Biological Resources, Incheon.
Lee S.H. 1991. A systematic study and distribution of Korean Dytiscidae. Science Fair 1991: 16–20. [In Korean].
Leigh J.W. & Bryant D. 2015. POPART: full-feature software for haplotype network construction. Methods in Ecology & Evolution 6 (9): 1110–1116. https://doi.org/10.1111/2041-210X.12410
Li Y.L. & Liu J.X. 2018. StructureSelector: a web-based software to select and visualize the optimal number of clusters using multiple methods. Molecular Ecology Resources 18 (1): 176–177. https://doi.org/10.1111/1755-0998.12719
Matsui E. 1987. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from Kyushu, Japan]. Gekkan-Mushi (202): 5–6. [In Japanese.]
Matsui E. 1988. [Aquatic insect collection record in 1987 (Tanega-shima Island)]. Satsuma 37 (99): 1–10. [In Japanese.]
Matsui E. 1989. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from Kume-jima Island, Okinawa Prefecture, Japan]. Gekkan-Mushi (226): 20. [In Japanese.]
Matsui E., Takai Y. & Tanabe T. 1988. Aquatic Coleoptera in Kagoshima Prefecture. Satsuma 37 (100): 61–115. [In Japanese.]
Matsui M. & Tominaga A. 2020. Distinct species status of a Microhyla from the Yaeyama Group of the Southern Ryukyus, Japan (Amphibia, Anura, Microhylidae). Current Herpetology 39 (2): 120–136. https://doi.org/10.5358/hsj.39.120
Miller K.B. & Nilsson A.N. 2003. Homology and terminology: Communicating information about rotated structures in water beetles. Latissimus 17: 1–4.
Mitamura T., Hirasawa K. & Yoshii S. 2017. The Handbook of Japanese Aquatic Insect, Volume 1: Coleoptera. Bun-ichi Sogo Shuppan, Tokyo. [In Japanese.]
Miyake T. 2021. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from Ôita Prefecture, Japan]. Nihônomushi 59: 100. [In Japanese.]
Mori M. & Kitayama A. 2002. Dytiscoidea of Japan. Bun-ichi Sogo Shuppan, Tokyo. [In Japanese.]
Morii T. 2018. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from Amami-ôshima Island, Japan]. Sayabane, New Series 29: 5. [In Japanese.]
Nakajima J. & Inoue D. 2004. [Records of Copelatus zimmermanni Gschwendtner, 1934 from Fukuoka Prefecture]. Coleopterists’ News 148: 14. [In Japanese.]
Nakajima J., Hayashi M., Ishida K., Kitano T. & Yoshitomi H. 2020. Aquatic Coleoptera and Hemiptera of Japan. Bun-ichi Sogo Shuppan, Tokyo. [In Japanese.]
Nakamine A. & Nakamine K. 2021. Records of aquatic insects collected on Tanegashima Island in 2020. Bulletin of the Kagoshima Prefectural Museum 40: 1–8. [In Japanese.]
Nakane T. 1963. New or little-known Coleoptera from Japan and its adjacent regions XVIII. Fragmenta Coleopterologica 6: 23–26.
Narukawa N., Ichihashi H., Nakanishi M. & Kawakita H. 2006. [Insect Fauna of the Kumanonada Costal Laurel Forest.] Mie Konchû Danwakai, Matsusaka. [In Japanese.]
Nilsson A.N. & Hájek J. 2026. A World Catalogue of the Family Dytiscidae, or the Diving Beetles (Coleoptera, Adephaga) Version 1.I.2026. Available from http://www.waterbeetles.eu [accessed 12 May 2026].
Nomura S., Mizokami S. & Ôtsuka K. 1996. [Aquatic Coleoptera of Saga Prefecture, Japan]: 249–262. Sagaken no Seibutsu.
Ôtsuka I. 1991. [A new record of Copelatus zimmermanni Gschwendtner, 1934 from the Amakusa Islands, Japan]. Gekkan-Mushi 240: 39. [In Japanese.]
Papadopoulou A., Anastasiou I. & Vogler A.P. 2010. Revisiting the insect mitochondrial molecular clock: the mid-Aegean trench calibration. Molecular Biology and Evolution 27 (7): 1659–1672. https://doi.org/10.1093/molbev/msq051
Park H.C., Lee Y.B. & Lee H.A. 2008. Taxonomic review of the genus Copelatus Erichson (Coleoptera: Dytiscidae) in Korea. Entomological Research 38 (1): 73–76. https://doi.org/10.1111/j.1748-5967.2008.00134.x
Pritchard J.K., Wen X. & Falush D. 2009. Documentation for Structure Software: Version 2.3. University of Chicago, Chicago.
QGIS Development Team 2024. QGIS Geographic Information System. Open Source Geospatial Foundation Project. Available from https://qgis.org/ [accessed 22 Dec. 2025].
R Core Team 2020. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from https://www.r-project.org/ [accessed 22 Dec. 2025].
Rambaut A. 2009. FigTree version 1.4.4. Available from https://tree.bio.ed.ac.uk/software/figtree/ [accessed 4 Mar. 2026].
Ranarilalatiana T. & Bergsten J. 2019. Discovery of a specialist Copelatinae fauna on Madagascar: highly ephemeral tropical forest floor depressions as an overlooked habitat for diving beetles (Coleoptera, Dytiscidae). ZooKeys 871: 89–118. https://doi.org/10.3897/zookeys.871.36337
Ranarilalatiana T., Harivelo L., Ravaomanarivo R. & Bergsten J. 2019. Taxonomic revision of the Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non-erichsonii group species. ZooKeys 871: 89–118. https://doi.org/10.3897/zookeys.869.33997
Régimbart M. 1899. Révision des Dytiscidae de la Région Indo-Sino-Malaise. Annales de la Société entomologique de France 68: 186–367. Available from https://www.biodiversitylibrary.org/page/8431385 [accessed 22 Dec. 2025].
Rochette N.C., Rivera-Colón A.G. & Catchen J.M. 2019. Stacks 2: analytical methods for paired-end sequencing improve RADseq-based population genomics. Molecular Ecology 28 (21): 4737–4754. https://doi.org/10.1111/mec.15253
Sakurai S. 1992. [Records of three Copelatus species from Chôkaisan, Honshu, Japan]. Gekkan-Mushi 262: 20–21. [In Japanese.]
Sato F. 2004. [Records of Copelatus zimmermanni Gschwendtner, 1934 from Oga Peninsula, Akita Prefecture]. Gekkan-Mushi 406: 8. [In Japanese.]
Satô M. 1983. Distribution of the genus Copelatus in Japan (Coleoptera: Dytiscidae). In: Satô M. (ed.) Special Issue Concerning the Aquatic Coleoptera Presented at the Workshop of the XVI International Congress of Entomology in Kyoto, Japan in 1980: 35–41. Nagoya.
Satô M. 1984. Taxonomic notes on the aquatic Coleoptera of Japan, II. Coleopterists’ News 66: 1–4. [In Japanese.]
Satô M. 1985. The genus Copelatus of Japan (Coleoptera: Dytiscidae). Transactions of the Shikoku Entomological Society 17: 57–67.
Satô M. 1998. Aquatic Coleoptera of the Ryukyu archipelago, III. Coleopterists’ News 121: 7–13. [In Japanese.]
Shaverdo H. & Balke M. 2022. A species-group key and notes on phylogeny and character evolution in New Guinean Exocelina Broun, 1886 diving beetles (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 1131: 31–58. https://doi.org/10.3897/zookeys.1131.94205
Shaverdo H., Surbakti S., Hendrich L. & Balke M. 2012. Introduction of the Exocelina ekari-group with descriptions of 22 new species from New Guinea (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 250: 1–76. https://doi.org/10.3897/zookeys.250.3715
Shaverdo H., Hájek J., Hendrich L., Surbakti S., Panjaitan R. & Balke M. 2023. Austrelatus gen. nov., a new genus of Australasian diving beetles (Coleoptera, Dytiscidae, Copelatinae), with the discovery of 31 new species from New Guinea. ZooKeys 1170: 1–164. https://doi.org/10.3897/zookeys.1170.103834
Shaverdo H., Hendrich L., Surbakti S., Panjaitan R. & Balke M. 2024. Revision of the Austrelatus papuensis group with descriptions of 42 new species from New Guinea (Coleoptera, Dytiscidae, Copelatinae). ZooKeys 1201: 1–165. https://doi.org/10.3897/zookeys.1201.116131
Shaverdo H., Wewalka G., Hendrich L. & Balke M. 2026 (in preparation). Revision of the Austrelatus species of Fiji (Coleoptera, Dytiscidae, Copelatinae), with descriptions of 29 new species and notes on Copelatinae species of the Pacific Islands.
Sheth S.D., Ghate H.V. & Hajek J. 2018. Copelatus Erichson, 1832 from Maharashtra, India, with description of three new species and notes on other taxa of the genus (Coleoptera: Dytiscidae: Copelatinae). Zootaxa 4459 (2): 235–260. https://doi.org/10.11646/zootaxa.4459.2.2
Suyama Y. & Matsuki Y. 2015. MIG-seq: an effective PCR-based method for genome-wide single-nucleotide polymorphism genotyping using the next-generation sequencing platform. Scientific Reports 5 (1): 16963. https://doi.org/10.1038/srep16963
Suyama Y., Hirota S.K., Matsuo A., Tsunamoto Y., Mitsuyuki C., Shimura A. & Okano K. 2022. Complementary combination of multiplex high-throughput DNA sequencing for molecular phylogeny. Ecological Research 37 (1): 171–181. https://doi.org/10.1111/1440-1703.12270
Tahira Y. & Kitano T. 2000. [Records of Copelatus zimmermanni Gschwendtner, 1934 from Nirayama-cho, Shizuoka Prefecture]. Gekkan-Mushi 357: 44−45. [In Japanese.]
Takahashi M. 1994. [Records of Copelatus zimmermanni Gschwendtner, 1934]. Coleopterists’ News 105: 5–6. [In Japanese.]
Tamu N. & Tsukamoto K. 1955. [On the Coleoptera in Kammuri Island (Kyoto, Japan) I (Studies on the insects in small islands, I)]. Akitu 4: 69–76. [In Japanese.]
Tamura K., Stecher G. & Kumar S. 2021. MEGA11: molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution 38 (7): 3022–3027. https://doi.org/10.1093/molbev/msab120
Thompson J.D., Higgins D.G. & Gibson T.J. 1994. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22 (22): 4673–4680. https://doi.org/10.1093/nar/22.22.4673
Watanabe K. 2022. Biological notes on immature stages of Platambus ussuriensis (Nilsson, 1997) and Copelatus nakamurai Guéorguiev, 1970 (Coleoptera: Dytiscidae). The Coleopterists Bulletin 76 (2): 233–236. https://doi.org/10.1649/0010-065X-76.2.233
Watanabe K. 2025. Review of Dytiscidae (Coleoptera) eggs: focus on Japanese species. Journal of Insect Conservation 29: 1. https://doi.org/10.1007/s10841-024-00636-6
Watanabe K. & Hayashi M. 2019. Reproductive ecology and immature stages of Copelatus masculinus Régimbart, 1899 (Coleoptera, Dytiscidae). Elytra, New Series 9 (2): 269–278.
Watanabe K. & Ohba S. 2022. Life history of Copelatus zimmermanni Gschwendtner, 1934 (Coleoptera: Dytiscidae) and the ecological significance of the larval period of three Copelatus species. Entomological Science 25 (2): e12505. https://doi.org/10.1111/ens.12505
Watanabe K. & Yoshitomi H. 2022 [2023]. Family Dytiscidae. In: the Editorial Committee of Catalogue of the Insects of Japan (ed.) Catalogue of the Insects of Japan, 6. Coleoptera, Part 1: 8–27. Touka Shobo, Fukuoka. [In Japanese.]
Watanabe K., Hayashi M. & Kato M. 2017. Immature stages and reproductive ecology of Copelatus parallelus Zimmermann, 1920 (Coleoptera, Dytiscidae). Elytra, New Series 7 (2): 361–374.
Watanabe K., Hayashi M. & Nagashima S. 2024. Life history of Copelatus kammuriensis Tamu and Tsukamoto, 1955 (Coleoptera: Dytiscidae: Copelatinae) and biological implications. Aquatic Insects 45 (2): 285–297. https://doi.org/10.1080/01650424.2023.2253250
Watanabe N., Arai K., Otsubo M., Toda M., Tominaga A., Chiyonobu S., Sato T., Ikeda T., Takahashi A., Ota H. & Iryu Y. 2023. Geological history of the land area between Okinawa Jima and Miyako Jima of the Ryukyu Islands, Japan, and its phylogeographical significance for the terrestrial organisms of these and adjacent islands. Progress in Earth and Planetary Science 10 (1): 40. https://doi.org/10.1186/s40645-023-00567-x
Wong T.K.F., Ly-Trong N., Ren H., Demotte P., Baños H., Roger A.J., Susko E., Bielow C., de Maio N., Goldman N., Hahn M.W., dos Reis M., Vinh L.S., Huttley G., Lanfear R. & Minh B.Q. 2026. IQ-TREE 3: phylogenomic inference software using complex evolutionary models. Molecular Biology and Evolution 43 (5): msag117. https://doi.org/10.1093/molbev/msag117
Zimmermann A. 1920. Bemerkenswerte Neuerwerbungen des Zoologischen Museums in Hamburg. Haliplidae, Dytiscidae et Gyrinidae. Entomologische Blätter 16: 224–234. Available from https://www.biodiversitylibrary.org/page/34024158 [accessed 22 Dec. 2025].
Zimmermann A. 1934. Monographie der paläarktischen Dytisciden. V. Colymbetinae (1. Teil). Koleopterologische Rundschau 20: 138–214.
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