Abstract
The present study provides an integrative taxonomic assessment of the burrowing mayfly family Ephemeridae (Ephemeroptera) from the Western Ghats, India, through a combination of morphological and molecular approaches. Four new species viz., Ephemera kapilaensis sp. nov., Ephemera kodai sp. nov., Ephemera kudremukha sp. nov., and Ephemera subramanyai sp. nov., are described based on both larval and imaginal characters. This work provides the first molecular characterization of Ephemera species from southern India and generates the first DNA barcodes for the subgenus Ephemera in this region. The finding highlights the importance of integrative taxonomy in resolving species boundaries and evolutionary relationships among Oriental Ephemera. Additionally, the study provides distributional data, habitat ecology of the new species, and an identification key to all known species of the Western Ghats.
References
- Bae, Y.J. (1995) Ephemera separigata, a new species of Ephemeridae (Insecta: Ephemeroptera) from Korea. Korean Journal of Systematic Zoology, 11 (2), 159–166.
- Balasubramanian, C., Venkataraman, K. & Sivaramakrishnan, K.G. (1996) Life stages of a south Indian burrowing mayfly, Ephemera (Aethephemera) nadinae McCafferty and Edmunds 1973 (Ephemeroptera: Ephemeridae). Aquatic Insects, 13 (4), 223–228. https://doi.org/10.1080/01650429109361449
- Burian, S.K. (2001) A revision of the genus Leptophlebia Westwood in North America (Ephemeroptera: Leptophlebiidae: Leptophlebiinae). Bulletin of Ohio Biological Survey, New Series, 13 (3), 1–80.
- Demoulin, G. (1955) Nathanella gen. hove Leptophlebiidae diptera de I'Inde (Ephemeroptera). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, 37 (77), 1–4.
- Eaton, A.E. (1871) A monograph on the Ephemeridae. In: Transactions of the Entomological Society of London. The Entomological Society of London, London, pp. 1–164, pls. 1–6.
- Eaton, A.E. (1883–1888) A revisional monograph of recent Ephemeridae or mayflies. Transactions of the Linnean Society of London, Series 2, Zoology, 3, 1–352, 65 pls. https://doi.org/10.1111/j.1096-3642.1883.tb01550a.x
- Felsenstein, J. (1985) Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39, 783–791. https://doi.org/10.2307/2408678
- 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, 294–299.
- Hafiz, H.A. (1937) The Indian Ephemeroptera (mayflies) of the sub-order Ephemeroidea. Records of the Indian Museum, 39, 351–370, pls. 4–5. https://doi.org/10.26515/rzsi/v39/i4/1937/162279
- Hebert, P.D.N., Cywinska, A., Ball, S.L. & DeWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, Series B: Biological sciences, 270, 313–321. https://doi.org/10.1098/rspb.2002.2218
- Hsu, Y.C. (1936–1937b) The mayflies of China (Order Ephemeroptera). Peking Natural History Bulletin, 11, 433–440.
- Hsu, Y.-C. (1936–1937b) The mayflies of China. Peking Natural History Bulletin, 11 (3), 287–296.
- Hubbard, M.D. (1982) Two new species of Ephemera from South India (Ephemeroptera: Ephemeridae). Pacific Insects, 24 (2), 192–195.
- Hubbard, M.D. (1990) Mayflies of the world. A Catalog of the Family and Genus Group Taxa (Insecta: Ephemeroptera). The Sandhill Crane Press, Inc., Gainesville, Florida, 119 pp.
- Hubbard, M.D. & Peters, W.L. (1978) A catalogue of the Ephemeroptera of the Indian Subregion. Oriental Insects, 9 (Supplement), 1–43. https://doi.org/10.1080/00305316.1978.11745231
- Hwang, J.M. & Bae, Y.J. (2008) Review of the tropical Southeast Asian Ephemera (Ephemeroptera: Ephemeridae). Aquatic Insects, 30 (2), 105–126. https://doi.org/10.1080/01650420701883333
- Hwang, J.M., Bae, Y.J. & McCafferty, W.P. (2008) A checklist of the burrowing mayfly family Ephemeridae. In: Hauer, F.R., Stanford, J.A. & Newell, R.L. (Eds.), International advances in the ecology, zoogeography and systematics of mayflies and stoneflies. University of California Publications in Entomology. No. 128. Proceedings in 11th International Conference on Ephemeroptera, Montana, USA, 22–29 August 2004, pp. 159–172. https://doi.org/10.1525/california/9780520098688.003.0012
- Hwang, J.-M., Yoon, T.-J., Suh, K.-I. & Bae, Y.-J. (2013) Molecular phylogeny evidence of altitudinal distribution and habitat adaptation in Korean Ephemera species (Ephemeroptera: Ephemeridae). Entomological Research, 43, 40–46. https://doi.org/10.1111/1748-5967.12008
- Kang, S.C. & Yang, C.T. (1994) Ephemeroptera of Taiwan (Ephemeroptera). Chinese Journal of Entomology, 14, 391–399.
- Kimura, M. (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120. https://doi.org/10.1007/BF01731581
- Kjærstad, G., Webb, J.M. & Ekrem, T. (2012) A review of the Ephemeroptera of Finnmark–DNA barcodes identify Holarctic relations. Norwegian Journal of Entomology, 59, 182–195.
- Kluge, N.J. (2004) The phylogenetic system of Ephemeroptera. Kluwer Academic Publishers, Dordrecht, Boston and London, 456 pp. https://doi.org/10.1007/978-94-007-0872-3
- Kondratieff, B.C. & Voshell, J.R. (1984) The North and Central American species of Isonychia (Ephemeroptera: Oligoneuriidae). Transactions of the American Entomological Society, 110, 129–244.
- Linnaeus, C. (1758) Systema Naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum Characteribus, Differentiis, Synonymis, Locis. Vol. 1. 10th Edition. Impensis Direct. Laurentii Salvii, holmiae, 824 pp. https://doi.org/10.5962/bhl.title.542
- McCafferty, W.P. (1971) New genus of mayflies from eastern North America (Ephemeroptera: Ephemeridae). Journal of the New York Entomological Society, 79 (4), 45–51.
- McCafferty, W.P. (1973) Systematic and zoogeographic aspects of Asiatic Ephemeridae (Ephemeroptera). Oriental Insects, 7 (1), 49–67. https://doi.org/10.1080/00305316.1973.10434204
- McCafferty, W. (1987) New fossil mayfly in amber and its relationships among extant Ephemeridae (Ephemeroptera). Annals of the Entomological Society of America, 80, 472–474. https://doi.org/10.1093faesaf80.4.472
- McCafferty, W.P. (1991) Toward a phylogenetic classification of the Ephemeroptera (Insecta): A commentary on systematics. Annals of the Entomological Society of America, 84 (4), 343–360. https://doi.org/10.1093/aesa/84.4.343
- McCafferty, W.P. (2004) Contribution to the systematics of Leucrocuta, Nixe, and related genera (Ephemeroptera: Heptageniidae). Transactions of the American Entomological Society, 130, 1–9.
- McCafferty, W.P. & Edmunds, G.F. Jr. (1973) Subgeneric classification of Ephemera (Ephemeroptera: Ephemeridae). Pan-Pacific Entomologist, 49 (4), 300–307.
- McCafferty, W.P. & Gillies, M.T. (1979) The African Ephemeridae (Ephemeroptera). Aquatic Insects, 1, 169–178. https://doi.org/10.1080/01650427909360991
- Muthukatturaja, M., Balasubramanian, C., Selvakumar, C. & Murali, P. (2025) A new species of Choroterpes Eaton, 1881 (Ephemeroptera: Leptophlebiidae) from the River Cauvery, Karnataka, India. Zootaxa, 5609 (2), 151–173. https://doi.org/10.11646/zootaxa.5609.2.1
- Navás, L. (1911) Deux Ephémérides (Ins. Neur.) nouveaux du Congo Belge. Annales de la Société Scientifique de Bruxelles, 35 (1), 221–224.
- Navás, L. (1934) Névroptères et insectes voisins. Chine et pays environnants. 6e Série. Notes d'Entomologie Chinoise (Musée Heude), 1 (14), 1–10.
- Navás, L. (1935) Décadas de insectos nuevos. Década 27. Brotéria (Ciências Naturais), 31, 97–107.
- Saitou, N. & Nei, M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4, 406–425.
- Sivaramakrishnan, K.G. (2016) Systematics of the Ephemeroptera of India: Present status and future prospects. Zoosymposia, 11, 33–52. https://doi.org/10.11646/zoosymposia.11.1.8
- Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA 11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution, 38 (7), 3022–3027. https://doi.org/10.1093/molbev/msab120
- Uéno, M. (1969) Mayflies (Ephemeroptera) from various regions of Southeast Asia. Oriental Insects, 3, 221–238. https://doi.org/10.1080/00305316.1969.10433911
- Ulmer, G. (1926) Trichopteren und Ephemeropteren, Beitrage zur Fauna sinaica. Archiv fur Naturgeschichte, 91, 19–110.
- Ulmer, G. (1939) Eintagsfliegen (Ephemeropteren) von den Sunda–Inselin. Archiv fuer Hydrobiologie, 16 (Supplement), 443–692.
- Walker, F. (1860) Characters of undescribed Neuroptera in the collection of W.W. Saunders, Esq., F.R.S., &c. Transactions of the Entomological Society of London, New Series, 5, 176–199. https://doi.org/10.1111/j.1365-2311.1860.tb01844.x
- Walsh, B.D. (1863) Observations on certain N.A. Neuroptera, by H. Hagen, M.D., of Koenigsberg, Prussia; translated from the original French ms., and published by permission of the author, with notes and descriptions of about twenty new N.A. sp. of Pseudoneuroptera. Proceedings of the Entomological Society of Philadelphia, 2, 167–272.
- Wang, L., Li, B., Jiang, J. & Tong, X.-L. (2022) The complete mitochondrial genome of Ephemera serica (Ephemeroptera: Ephemeridae) and phylogenetic analysis. Mitochondrial DNA, Part B, 7 (3), 461–463. https://doi.org/10.1080/23802359.2022.2044401
- Wang, D., Sun, Y.-X., Wang, N.-N. & Zhou, C.-F. (2025) A new Chinese Ephemera species (Ephemeroptera: Ephemeridae) from a small muddy canal. Journal of Insect Biodiversity, 68 (1), 18–31. https://doi.org/10.12976/jib/2025.68.1.2
- Webb, J.M., Jacobus, L.M., Funk, D.H., Zhou, X., Kondratieff, B., Geraci, C.J., DeWalt, R.E., Baird, D.J., Richard, B., Phillips, I. & Hebert, P.D.N. (2012) A DNA Barcode library for North American Ephemeroptera: progress and prospects. PLoS ONE, 7 (5), e38063. https://doi.org/10.1371/journal.pone.0038063
- Wolf, H.G. & Mort, M.A. (1986) Inter–specific hybridization underlies phenotypic variability in Daphnia populations. Oecologia, Berlin, 68, 507–511. https://doi.org/10.1007/BF00378763
- Yeh, W.-B., Yang, C.-T. & Kang, S.-C. (1997) Identification of two sibling species, Ephemera formosana and E. sauteri (Ephemeroptera: Ephemeidae), based on mitochondrial DNA sequence analysis. Chinese Journal of Entomology, 17, 257–268.
- Zhou, C.-F. & Zheng, L.-Y. (2003) Two synonyms and a new species of the genus Ephemera from China (Ephemeroptera: Ephemeridae). Acta Zootaxonomica Sinica, 28 (4), 665–668.
