{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,6]],"date-time":"2026-03-06T12:33:01Z","timestamp":1772800381815,"version":"3.50.1"},"reference-count":160,"publisher":"Magnolia Press","issue":"1","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"<jats:p>The genus Thraulus is widespread throughout much of the Eastern Hemisphere. Since Eaton established Thraulus in 1881, 62 species have, at one time or another, been placed in this genus. Thirty-eight of those species were eventually moved to other genera. Any comprehensive study of the remaining species, based on the published literature, is difficult as they were described by many authors, using different criteria, over a period of 142 years. The purpose of this study was to redescribe this genus, based on previously described species and nine new species, and to provide a format for future taxonomic and morphological studies of Thraulus. Redescriptions of most species were based on direct examination of external morphological characters. Descriptions or diagnoses of species, whose types were unavailable for study, were made using the original published description and additional information provided by authors of several of those species. The following species were studied: Thraulus amravati Vasanth, Subramanian &amp; Selvakumar, 2022; T. bellus Eaton, 1881; T. bishopi Peters &amp; Tsui, 1972; T. cuspidatus Vasanth, Subramanian &amp; Selvakumar, 2022; T. demoulini Peters &amp; Tsui, 1973; T. fasciatus (Kimmins, 1956); T. fatuus Kang &amp; Yang, 1994; T. femoratus Li, Liu &amp; Zhou, 2006; T. gopalani Grant &amp; Sivaramakrishnan, 1985; T. jacobusi Isack, Srinivasan, Sivaruban &amp; Barathy, 2022; T. macilentus Kang &amp; Yang, 1994; T. malabarensis Vasanth, Subramanian &amp; Selvakumar, 2022; T. mudumalaiensis Soman, 1991; T. plumeus Selvakumar, Vasanth &amp; Subramanian, 2022; T. semicastaneus (Gillies, 1951); T. thiagarajani Balasubramanian &amp; Muthukatturaja, 2019; T. thraker Jacob, 1988; T. torrentis (Gillies, 1964); T. turbinatus (Ulmer, 1909); T. umbrosus Kang &amp; Yang, 1994; and T. vellimalaiensis Vasanth, Subramanian &amp; Selvakumar, 2022. Nine new species of Thraulus are described: T. connubialis sp. nov., Malaysia; T. cursus sp. nov., Japan; T. eatoni sp. nov., Indonesia; T. ishiwatai sp. nov., Japan; T. madagasikarensis sp. nov., Madagascar; T. nihonensis sp. nov., Japan; T. opifer sp. nov., Australia; T. parentalis sp. nov., Malaysia; and T. petersorum sp. nov., Malaysia. Thraulus can be distinguished from all other genera of Leptophlebiidae by the following combination of characters: In the imagos, 1) upper portion of eyes oval-suboval, major axes diverge anteriorly; 2) vein MA fork of fore wings symmetrical; 3) vein MP fork of fore wings asymmetrical\u2014a cross vein connects base of MP2 to MP1, MP fork closer to base of wing than Rs fork; 4) strongly oblique cross vein extends between veins R4+5 and MA1 just apical to fork of vein MA; 5) 2 cubital intercalary veins in fore wings; 6) costal projection on hind wings well-developed, bluntly rounded to acutely pointed; 7) claws dissimilar\u2014one blunt and pad-like, the other apically hooked; 8) penes long, relatively straight, narrow, parallel, usually contiguous mesally but not fused, apex may have lateral projections; 9) sternum 7 of female with posterior margin straight or shallowly concave or convex mesally; and 10) sternum 9 of females rounded apically. In addition, penile spines occur on most species. In the nymphs, 1) lateral margins of clypeus parallel; 2) width of labrum subequal to width of clypeus; 3) 2 dorsal rows of setae on labrum; 4) venter of labrum with 1 row of short stout setae on either side of midline near anterior margin, rows curve mesally; 5) hypopharynx with small, rounded, posterolateral projections on arms of superlingua; 6) large spine on posterolateral corners of terga 6\u20139, 7\u20139 or 8\u20139; 7) gills 1\u20137 dissimilar: gill 1 composed of 1 or 2 subulate lamellae or a dorsal subulate lamella and a ventral fimbriate oval lamella, and gills 2\u20137 composed of dorsal and ventral oval lamellae with fimbriate margins. Two species continue to be nomen dubium\u2014T. siewertii (Weyenbergh, 1883) and T. vogleri (Weyenbergh, 1883). Thraulus grandis Gose, 1980 is considered nomen nudum. A review of published phylogenetic studies involving Thraulus is provided. With the species discussed in this paper, along with reports of additional new species to be described, Thraulus has the potential to be included among the more specious genera of Ephemeroptera.\n\u00a0<\/jats:p>","DOI":"10.11646\/zootaxa.5407.1.1","type":"journal-article","created":{"date-parts":[[2024,2,8]],"date-time":"2024-02-08T23:01:59Z","timestamp":1707433319000},"page":"1-87","source":"Crossref","is-referenced-by-count":3,"title":["Revision of Thraulus Eaton 1881 (Ephemeroptera: Leptophlebiidae: Choroterpinae)"],"prefix":"10.11646","volume":"5407","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4932-2209","authenticated-orcid":false,"given":"PETER M.","family":"GRANT","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"4913","published-online":{"date-parts":[[2024,2,9]]},"reference":[{"key":"277944","doi-asserted-by":"crossref","unstructured":"Alhou, B., Micha, J.-C., Dodo, A. & Awaiss, A. 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(1990) The Australian mayfly fauna: composition, distribution and convergence. In: Campbell, I.C. (Ed.), Mayflies and stoneflies: life histories and biology. Kluwer Academic Publishers, Dordrecht, pp. 149\u2013153. https:\/\/doi.org\/10.1007\/978-94-009-2397-3_19","DOI":"10.1007\/978-94-009-2397-3_19"},{"key":"277962","doi-asserted-by":"crossref","unstructured":"Campbell, I.C. & Suter, P.J. (1988) Three new genera, a new subgenus and a new species of Leptophlebiidae (Ephemeroptera) from Australia. Journal of the Australian Entomological Society, 27, 259\u2013273. https:\/\/doi.org\/10.1111\/j.1440-6055.1988.tb01172.x","DOI":"10.1111\/j.1440-6055.1988.tb01172.x"},{"key":"277963","doi-asserted-by":"crossref","unstructured":"Cozzarolo, C.-S., Balke, M., Buerki, S., Arrigo, N., Pitteloud, C., Gueuning, M., Salamin, N., Sartori, M. & Alvarez, N. (2019) Biogeography and ecological diversification of a mayfly clade in New Guinea. Frontiers in Ecology and Evolution, 7, 233. https:\/\/doi.org\/10.3389\/fevo.2019.00233","DOI":"10.3389\/fevo.2019.00233"},{"key":"277964","unstructured":"Dean, J.C. (1999) Preliminary Keys for the Identification of Australian Mayfly Nymphs of the Family Leptophlebiidae Identification Guide No. 20. Cooperative Research Centre for Freshwater Ecology, Thurgoona, 91 pp."},{"key":"277965","unstructured":"Degrange, C. (1960) Recherches sur la reproduction des \u00c9ph\u00e9m\u00e9ropt\u00e8res. Travaux du Laboratoire d\u2019Hydrobiologie et de Pisciculture de l\u2019Universit\u00e9 de Grenoble, 50\/51, 7\u2013193."},{"key":"277966","unstructured":"Demoulin, G. (1954) Les \u00c9ph\u00e9m\u00e9ropt\u00e8res Leptophlebiidae de Born\u00e9o. Bulletin et Annales de la Soci\u00e9t\u00e9 Entomologique de Belgique, 90 (5\u20136), 123\u2013131."},{"key":"277967","unstructured":"Demoulin, G. (1955a) Note sur deux nouveaux genres de Leptophlebiidae d\u2019Australie (Ephemeroptera). Bulletin et Annales de la Soci\u00e9t\u00e9 Royale Entomologique de Belgique, 91 (9\u201310), 227\u2013229."},{"key":"277968","unstructured":"Demoulin, G. (1955b) Une mission biologique belge au Br\u00e9sil. \u00c9ph\u00e9m\u00e9ropt\u00e8res. Bulletin de l\u2019Institut Royal des Sciences Naturelles de Belgique, 31 (20), 1\u201332."},{"key":"277969","unstructured":"Demoulin, G. (1969) Les Eph\u00e9m\u00e9ropt\u00e8res r\u00e9colt\u00e9s par la mission danoise du Noona Dan aux \u00eeles Philippines et Bismarck. Entomologiske Meddelelser, 37 3, 225\u2013241."},{"key":"277970","unstructured":"Diarra, B., Konan, K.J., Yapo, L.M. & Kouassi, K.P. (2018) Aquatic macroinvertebrates associated with free-floating macrophytes in a marginal lentic ecosystem (Ono Lagoon, C\u020fte d\u2019Ivoire). Journal of Entomology and Zoology Studies, 6, 1432\u20131441."},{"key":"277971","doi-asserted-by":"crossref","unstructured":"Dom\u00ednguez, E. (1999) Systematics, cladistics and biogeography of the American genus Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae). Zoological Journal of the Linnean Society, 126, 155\u2013189. https:\/\/doi.org\/10.1111\/j.1096-3642.1999.tb00152.x","DOI":"10.1111\/j.1096-3642.1999.tb00152.x"},{"key":"277972","doi-asserted-by":"crossref","unstructured":"Dom\u00ednguez, E. & Flowers, R.W. (1989) A revision of Hermanella and related genera (Ephemeroptera: Leptophlebiidae; Atalophlebiinae) from Subtropical South America. Annals of the Entomological Society of America, 82, 555\u2013573. https:\/\/doi.org\/10.1093\/aesa\/82.5.555","DOI":"10.1093\/aesa\/82.5.555"},{"key":"277973","unstructured":"Dom\u00ednguez, D., Hubbard, M.D. & Pescador, M.L. (1994) Los Ephemeroptera en Argentina. In: Castellanos, Z.A. de (Ed.), Fauna de agua dulce de la Rep\u00fablica Argentina. Vol. 33. Fasc. 1. Profadu, La Plata, pp. 1\u2013142."},{"key":"277974","unstructured":"Dom\u00ednguez, E., Molineri, C., Pescador, M.L., Hubbard, M.D. & Nieto, C. (2006) Ephemeroptera of South America. In: Adis, J., Arias, J.R., Rueda-Delgado, G. & Wantzen, K.M. (Eds.), Aquatic Biodiversity in Latin America (ABLA). Vol. 2. Pensoft, Sofia-Moscow, pp. 1\u2013646."},{"key":"277975","doi-asserted-by":"crossref","unstructured":"Dom\u00ednguez, E., Peters, W.L., Peters, J.G. & Savage, H.M. (1997) The imago of Simothraulopsis Demoulin with a redescription of the nymph (Ephemeroptera: Leptophlebiidae: Atalophlebiinae). Aquatic Insects, 19, 141\u2013150. https:\/\/doi.org\/10.1080\/01650429709361648","DOI":"10.1080\/01650429709361648"},{"key":"277976","unstructured":"Eaton, A.E. (1881) An announcement of new genera of the Ephemeridae. The Entomologist\u2019s Monthly Magazine, 17, 191\u2013197."},{"key":"277977","doi-asserted-by":"crossref","unstructured":"Eaton, A.E. (1883\u20131888) A revisional monograph of recent Ephemeridae or mayflies. The Transactions of the Linnean Society of London, Series 2, Zoology, 3, 1\u2013352. https:\/\/doi.org\/10.1111\/j.1096-3642.1883.tb01550a.x","DOI":"10.1111\/j.1096-3642.1883.tb01550a.x"},{"key":"277978","unstructured":"Eaton, A.E. (1887) Notes on the entomology of Portugal.-IX. Ephemeridae. The Entomologist\u2019s Monthly Magazine, 24, 4\u00ad\u20136."},{"key":"277979","unstructured":"Eaton, A.E. (1892) Fam. Ephemeridae. Biologia Centrali-Americana, 38, 1\u201316."},{"key":"277980","unstructured":"Edmunds, G.F. Jr. (1948) A new genus of mayflies from western North America (Leptophlebiinae). Proceedings of the Biological Society of Washington, 61,141\u2013148."},{"key":"277981","unstructured":"Edmunds, G.F. Jr. (1950) Notes on Neotropical Ephemeroptera. I. New and little known Leptophlebiidae. Revista de Entomologia, 21, 551\u00ad\u2013554."},{"key":"277982","unstructured":"Edmunds, G.F. Jr. (1955) Ephemeroptera. In: Kessel, E.L. (Ed.), A Century of Progress in the Natural Sciences 1853\u20131953. California Academy of Sciences, San Francisco, California, pp. 509\u2013512."},{"key":"277983","doi-asserted-by":"crossref","unstructured":"El Alami, M., El Yaagoubi, S., Gattolliat, J.-L., Sartori, M. & Dakki, M. (2022) Diversity and distribution of mayflies from Morocco (Ephemeroptera, Insecta). Diversity, 14, 498. https:\/\/doi.org\/10.3390\/d14060498","DOI":"10.3390\/d14060498"},{"key":"277984","doi-asserted-by":"crossref","unstructured":"Esben-Petersen, M. (1912a) New and little-known species of Ephemerida from Argentine. (Neuropt.) Deutsche entomologische Zeitschrift, 1912, 333\u2013342.","DOI":"10.1002\/mmnd.48019120310"},{"key":"277985","unstructured":"Esben-Petersen, M. (1912b) Addition to the knowledge of the neuropterous insect fauna of Corsica. 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(1975) Segnalazione di Thraulus bellus Eaton in Italia (Ephemeroidea). Bollettino della Societ\u00e0 Entomologica Italiana, 107, 25\u201331."},{"key":"277989","unstructured":"Garc\u00eda, L., Pardo, I. & Nebra, A. (2007) Un nuevo Leptophlebiidae (Ephemeroptera) para las Islas Baleares: Thraulus bellus (Eaton, 1881). Bolet\u00edn de la Asociaci\u00f3n Espa\u00f1ola de Entomologia, 3, 193\u2013195."},{"key":"277990","unstructured":"Gauthier, M. (1952) Stations d\u2019\u00c9ph\u00e9m\u00e9ropt\u00e8res dans les Alpes du Dauphin\u00e9. Travaux du Laboratoire d\u2019Hydrobiologie et de Pisciculture de l\u2019Universit\u00e9 de Grenoble, 43\u201344, 9\u201322."},{"key":"277991","doi-asserted-by":"crossref","unstructured":"Gillies, M. (1951) Further notes on Ephemeroptera from India and South East Asia. 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