{"status":"ok","message-type":"work-list","message-version":"1.0.0","message":{"facets":{},"total-results":17,"items":[{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T14:09:37Z","timestamp":1785420577102,"version":"3.56.0"},"reference-count":0,"publisher":"Entomological Society of Indochina","issue":"44","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["ICE"],"abstract":"A new species of torrent beetle, Satonius jiadongi sp. nov., is described from Hengqin Island, Guangdong Province, China, which is the southernmost species of the genus Satonius Endr\u00f6dy-Younga, 1997 in the world, and the 13th known species of the family Torridincolidae in China. The new species resembles Satonius panghongae Liang, Jiang, Bergsten, Jia & Ge, 2024 and S. xinghuiensis Liang, Jiang, Bergsten, Jia & Ge, 2024 in having a body length of less than 1.7 mm and male genitalia with strongly reduced paramere bearing only one apical seta. The adult of the new species is diagnosed and illustrated in detail. A brief description and illustrations of its larvae are also provided.<\/jats:p>","DOI":"10.70590\/ice.2026.02.44","type":"journal-article","created":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T13:18:35Z","timestamp":1785417515000},"page":"475-484","source":"Crossref","is-referenced-by-count":0,"title":["Satonius jiadongi sp. nov., a new species of torrent beetle from Hengqin Island, China (Coleoptera: Myxophaga: Torridincolidae)"],"prefix":"10.70590","volume":"2","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3124-2021","authenticated-orcid":false,"given":"Zu-Qi","family":"Mai","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"51386","published-online":{"date-parts":[[2026,7,30]]},"container-title":["The Indochina Entomologist"],"deposited":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T13:18:36Z","timestamp":1785417516000},"score":12.812983,"resource":{"primary":{"URL":"https:\/\/www.indochinaentomologist.com\/articles\/389.html"}},"issued":{"date-parts":[[2026,7,30]]},"references-count":0,"journal-issue":{"issue":"44","published-online":{"date-parts":[[2026,7,30]]}},"URL":"https:\/\/doi.org\/10.70590\/ice.2026.02.44","ISSN":["3065-1107"],"issn-type":[{"value":"3065-1107","type":"electronic"}],"published":{"date-parts":[[2026,7,30]]}},{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T16:33:42Z","timestamp":1790181222322,"version":"4.1.0"},"reference-count":17,"publisher":"Magnolia Press","issue":"1","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"Species of the genus Satonius Endr\u00f6dy-Younga from China are briefly reviewed. Seven new species are described: Satonius cheni sp. nov. from Guangdong, Satonius nigriventralis sp. nov. from Shaanxi, Satonius panghongae sp. nov. from Jiangxi, Hunan, Guangxi and Zhejiang, Satonius quzhouensis sp. nov. from Zhejiang and Anhui, Satonius xinhuiensis sp. nov. from Guangdong, Satonius zhangi sp. nov. from Hunan, and Satonius zhenhuai sp. nov. from Hubei. Species diagnoses of the new species are provided, along with images of the habitus and illustrations of the aedeagus and metatrochanters for all available Chinese species. Additionally, further faunistic data of known Chinese species is supplemented. A key to all known species of the genus is also included. \n\u00a0<\/jats:p>","DOI":"10.11646\/zootaxa.5528.1.8","type":"journal-article","created":{"date-parts":[[2024,10,22]],"date-time":"2024-10-22T23:54:05Z","timestamp":1729641245000},"page":"89-105","source":"Crossref","is-referenced-by-count":2,"title":["A review of the genus Satonius Endr\u00f6dy-Younga from China with description of seven new species (Coleoptera: Myxophaga: Torridincolidae)"],"prefix":"10.11646","volume":"5528","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1289-2697","authenticated-orcid":false,"given":"ZU-LONG","family":"LIANG","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6825-4283","authenticated-orcid":false,"given":"ZHUO-YIN","family":"JIANG","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6153-4431","authenticated-orcid":false,"given":"JOHANNES","family":"BERGSTEN","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2391-5038","authenticated-orcid":false,"given":"FENG-LONG","family":"JIA","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5924-3400","authenticated-orcid":false,"given":"SI-QIN","family":"GE","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4913","published-online":{"date-parts":[[2024,10,23]]},"reference":[{"key":"364938","doi-asserted-by":"crossref","unstructured":"Alarie, Y., Michat, M.C., Jia, F.-L. & H\u00e1jek, J. (2019) Hydrotrupes chinensis Nilsson, 2003 (Coleoptera: Dytiscidae): new records, (re)description of adult and larva, and notes on its biology. Aquatic Insects, 40 (3), 236\u2013256.Benetti, C. J. & Hamada, N. (2016) Three new species of Claudiella Reichardt & Vanin, 1976 (Coleoptera, Torridincolidae) from Brazil. Zootaxa, 4205 (2), 151\u2013161. https:\/\/doi.org\/10.1080\/01650424.2019.1601229","DOI":"10.1080\/01650424.2019.1601229"},{"key":"364939","doi-asserted-by":"crossref","unstructured":"Bergsten, J. (2022) Coleoptera: Torridincolidae, torrent beetles. In: Goodman, S.M. (Ed.), The new natural history of Madagascar. Princeton University Press, Princeton, pp. 1047\u20131049. https:\/\/doi.org\/10.2307\/j.ctv2ks6tbb.135","DOI":"10.2307\/j.ctv2ks6tbb.135"},{"key":"364940","unstructured":"Beutel, R.G. & Vanin, S.A. (2016) Torridincolidae Steffan, 1964. In: Beutel, R.G. & Leschen, R.A.B. (Eds.), Handbook of zoology. A natural history of the phyla of the animal kingdom. Vol. IV. Arthropoda: Insecta. Part 38. Coleoptera, Beetles. Vol. 1. Morphology and systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim.). 2nd Edition. Walter de Gruyter, Berlin and New York, pp. 67\u201370."},{"key":"364941","doi-asserted-by":"crossref","unstructured":"Bilton, D.T. & Mlambo, M.C. (2023) A revision of Delevea Reichardt, 1976: a Southern African endemic water beetle genus including the largest known extant myxophagans (Coleoptera: Myxophaga: Torridincolidae). Zootaxa, 5285 (2), 311\u2013324. https:\/\/doi.org\/10.11646\/zootaxa.5285.2.5","DOI":"10.11646\/zootaxa.5285.2.5"},{"key":"364942","doi-asserted-by":"crossref","unstructured":"Chen, L., Song, Y. & Xu, S. (2008) The boundary of palaearctic and oriental realms in western China. Progress in Natural Science, 18, 833\u2013841. https:\/\/doi.org\/10.1016\/j.pnsc.2008.02.004","DOI":"10.1016\/j.pnsc.2008.02.004"},{"key":"364943","unstructured":"Endr\u00f6dy-Younga, S. (1997) Active extraction of water-dissolved oxygen and descriptions of new taxa of Torridincolidae (Coleoptera: Myxophaga). Annals of the Transvaal Museum, 36 (24), 313\u2013332."},{"key":"364944","unstructured":"H\u00e1jek, J. & Fik\u00e1\u010dek, M. (2008) A review of the genus Satonius (Coleoptera: Myxophaga: Torridincolidae): taxonomic revision, larval morphology, notes on wing polymorphism, and phylogenetic implications. Acta Entomologica Musei Nationalis Pragae, 48 (2), 655\u2013676."},{"key":"364945","doi-asserted-by":"crossref","unstructured":"H\u00e1jek, J., Yoshitomi, H., Fik\u00e1\u010dek, M., Hayashi, M. & Jia, F.-L. (2011) Two new species of Satonius Endr\u00f6dy-Younga from China and notes on the wing polymorphism of S. kurosawai Sat\u00f4 (Coleoptera: Myxophaga: Torridincolidae). Zootaxa, 3016 (1), 51\u201362. https:\/\/doi.org\/10.11646\/zootaxa.3016.1.4","DOI":"10.11646\/zootaxa.3016.1.4"},{"key":"364946","doi-asserted-by":"crossref","unstructured":"Holt, B.G., Lessard, J.-P., Borregaard, M.K., Fritz, S.A., Ara\u00fajo, M.B., Dimitrov, D., Fabre, P.H., Graham, C.H., Graves, G.R., J\u00f8nsson, K.A. & Nogu\u00e9s-Bravo, D. (2013) An Update of Wallace\u2019s Zoogeographic Regions of the World. Science, 339, 74\u201378. https:\/\/doi.org\/10.1126\/science.1228282","DOI":"10.1126\/science.1228282"},{"key":"364947","unstructured":"J\u00e4ch, M.A. (1998) Torridincolidae: I. First record of Torridincolidae from China (Coleoptera). In: J\u00e4ch, M.A. & Ji, L. (Eds.), Water beetles of China. Vol. II. Zoologisch-Botanische Gesellschaft in \u00d6sterreich and Wiener Coleopterologenverein, Wien, pp. 51\u201352."},{"key":"364948","doi-asserted-by":"crossref","unstructured":"Perkins, P.D. & Bergsten, J. (2019) New Myxophagan water beetles from Madagascar (Coleoptera: Torridincolidae, Hydroscaphidae). Zootaxa, 4657 (1), 57\u201396. https:\/\/doi.org\/10.11646\/zootaxa.4657.1.2","DOI":"10.11646\/zootaxa.4657.1.2"},{"key":"364949","doi-asserted-by":"crossref","unstructured":"Reichardt, H. (1973) A critical study of the suborder Myxophaga, with a taxonomic revision of the Brazilian Torridincolidae and Hydroscaphidae (Coleoptera). Arquivos de Zoologia, S\u00e3o Paulo, 24, 73\u2013162. https:\/\/doi.org\/10.11606\/issn.2176-7793.v24i2p73-162","DOI":"10.11606\/issn.2176-7793.v24i2p73-162"},{"key":"364950","doi-asserted-by":"crossref","unstructured":"Sampaio, B.H.L. & Ferreira, N. Jr. (2014) A new species of Iapir Py-Daniel, Fonseca & Barbosa (Coleoptera: Myxophaga: Torridincolidae) from Brazil with key to species of the genus. Zootaxa, 3753 (2), 196\u2013200. https:\/\/doi.org\/10.11646\/zootaxa.3753.2.9","DOI":"10.11646\/zootaxa.3753.2.9"},{"key":"364951","doi-asserted-by":"crossref","unstructured":"Sampaio, B.H.L. & Ferreira, N. Jr. (2018) Five new species of Ytu Reichardt (Coleoptera: Myxophaga: Torridincolidae) and new records from Brazil. Zootaxa, 4402 (3), 508\u2013524. https:\/\/doi.org\/10.11646\/zootaxa.4402.3.5","DOI":"10.11646\/zootaxa.4402.3.5"},{"key":"364952","doi-asserted-by":"crossref","unstructured":"Sampaio, B.H.L. & Short, A.E.Z. (2018) Families Hydroscaphidae and Torridincolidae. In: Hamada, N., Thorp, J.H. & Rogers, C. (Eds.), Thorp and Covich\u2019s Freshwater Invertebrates. Keys to Neotropical Hexapoda. Vol. 3. 4th Edition. Academic Press, Elsevier Inc., London, San Diego, Cambridge, Oxford, pp. 519\u2013525. https:\/\/doi.org\/10.1016\/B978-0-12-804223-6.00022-6","DOI":"10.1016\/B978-0-12-804223-6.00022-6"},{"key":"364953","unstructured":"Sat\u00f4, M. (1982) Discovery of Torridincolidae (Coleoptera) in Japan. Annotationes Zoologicae Japonenses, 55, 276\u2013283."},{"key":"364954","unstructured":"Shepard, W.D., Barr, C.B. & Aguilar Julio, C.A. (2013) The occurrence of Torridincolidae (Coleoptera: Myxophaga) in Paraguay and a world checklist of species. Bolet\u00edn del Museo Nacional de Historia Natural del Paraguay, 17 (1), 76\u201382."}],"container-title":["Zootaxa"],"link":[{"URL":"https:\/\/mapress.com\/zt\/article\/download\/zootaxa.5528.1.8\/54637","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/mapress.com\/zt\/article\/download\/zootaxa.5528.1.8\/54637","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,22]],"date-time":"2024-10-22T23:54:20Z","timestamp":1729641260000},"score":11.738806,"resource":{"primary":{"URL":"https:\/\/mapress.com\/zt\/article\/view\/zootaxa.5528.1.8"}},"issued":{"date-parts":[[2024,10,23]]},"references-count":17,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2024,10,23]]}},"URL":"https:\/\/doi.org\/10.11646\/zootaxa.5528.1.8","ISSN":["1175-5334","1175-5326"],"issn-type":[{"value":"1175-5334","type":"electronic"},{"value":"1175-5326","type":"print"}],"published":{"date-parts":[[2024,10,23]]}},{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T16:33:46Z","timestamp":1790181226951,"version":"4.1.0"},"reference-count":13,"publisher":"Magnolia Press","issue":"1","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"Two new species of Torridincolidae from China, Satonius fui sp. nov. (Hubei) and S. jaechi sp. nov. (Fujian) are described, illustrated and compared with other species of the genus. The polymorphism of metathoracic wings was studied in the Japanese S. kurosawai (Sat\u00f4, 1982), which macropterous form is presented here for the first time. In addition, new records of S. stysi H\u00e1jek & Fik\u00e1\u010dek, 2008 in China are also briefly mentioned.<\/jats:p>","DOI":"10.11646\/zootaxa.3016.1.4","type":"journal-article","created":{"date-parts":[[2019,3,4]],"date-time":"2019-03-04T06:38:45Z","timestamp":1551681525000},"source":"Crossref","is-referenced-by-count":10,"title":["Two new species of Satonius Endr\u00f6dy-Younga from China and notes on the wing polymorphism of S. kurosawai Sat\u00f4 (Coleoptera: Myxophaga: Torridincolidae)"],"prefix":"10.11646","volume":"3016","author":[{"given":"JI\u0158\u00cd","family":"H\u00c1JEK","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"HIROYUKI","family":"YOSHITOMI","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"MARTIN","family":"FIK\u00c1\u010cEK","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"MASAKAZU","family":"HAYASHI","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"FENG-LONG","family":"JIA","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4913","published-online":{"date-parts":[[2011,9,7]]},"reference":[{"key":"#cr-split#-34918.1","unstructured":"

Akiyama, Y. (2008) Miscellaneous notes of Coleoptera from Hiroshima Prefecture (8). Miscellaneous Reports of the Hiwa Museum for Natural History<\/em>, 49, 119-146, 2 pls. (in Japanese with English title).<\/p>

Beutel, R.G. &"},{"key":"#cr-split#-34918.2","unstructured":"Vanin, S.A. (2005) 6.2. Torridincolidae Steffan, 1964, pp. 46-48. In: Beutel, R.G. &"},{"key":"#cr-split#-34918.3","unstructured":"Leschen, R.A.B. (Eds.) Handbook of zoology. A natural history of the phyla of the animal kingdom. Volume IV. Arthropoda: Insecta. Part 38<\/em>. Coleoptera, Beetles. Volume 1: Morphology and systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim.). <\/em>Berlin, New York: Walter de Gruyter, xi + 567 pp.<\/p>

Endr\u00f6dy-Younga, S. (1997) Active extraction of water-dissolved oxygen and descriptions new taxa of Torridincolidae (Coleoptera: Myxophaga). Annale van die Transvaal Museum<\/em>, 36 (24), 313-332.<\/p>

Falamarzi, S., P\u00fctz, A., Heidari, M. &"},{"key":"#cr-split#-34918.4","unstructured":"Nasserzadeh, H. (2010) Confirmed occurrence of Hydroscapha granulum<\/em> in Iran, with notes on its biology (Coleoptera: Myxophaga: Hydroscaphidae). Acta Entomologica Musei Nationalis Pragae<\/em>, 50, 97-106.<\/p>

Fik\u00e1\u010dek, M. &"},{"key":"#cr-split#-34918.5","doi-asserted-by":"crossref","unstructured":"\u0160\u00edpkov\u00e1, H. (2009) New Asian Hydroscapha<\/em>, with comments on male-female association of co-occuring species (Coleoptera, Myxophaga, Hydroscaphidae). Zootaxa<\/em>, 2286, 31-48.<\/p>

H\u00e1jek, J. &","DOI":"10.11646\/zootaxa.2286.1.2"},{"key":"#cr-split#-34918.6","unstructured":"Fik\u00e1\u010dek, M. (2008) A review of the genus Satonius <\/em>(Coleoptera: Myxophaga: Torridincolidae): taxonomic revision, larval morphology, notes on wing polymorphism, and phylogenetic implications. Acta Entomologica Musei Nationalis Pragae<\/em>, 48, 655-676.<\/p>

Hammer, \u00d8., Harper, D.A.T. &"},{"key":"#cr-split#-34918.7","unstructured":"Ryan, P.D. (2001) Paleontological Statistics software package for education and data analysis. Palaeontologica Electronica<\/em>, <\/em>4(1), 1-9.<\/p>

Hayashi, M. (2007). Distributional records and ecological notes on aquatic Coleoptera of Shimane Prefecture. Bulletin of the Hoshizaki Green Fondation<\/em>, 10, 77-113 (in Japanese with English title).<\/p>

Hayashi, M. &"},{"key":"#cr-split#-34918.8","unstructured":"Kadowaki, H. (2007) Records on aquatic Coleoptera from rivers in and around Mt. Daisen, Tottori Prefecture, Japan. Bulletin of the Hoshizaki Green Fondation<\/em>, 10, 149-168 (in Japanese with English title).<\/p>

Hayashi, M. &"},{"key":"#cr-split#-34918.9","unstructured":"Fujiwara, J. (2007) Records on aquatic Coleoptera of Yakushima, Kagoshima Prefecture, Japan. Coleopterists' News<\/em>, 159, 7-10 (in Japanese with English title).<\/p>

J\u00e4ch, M.A. (1998) Torridincolidae: I. First record of Torridincolidae from China (Coleoptera), pp. 51-52. In: J\u00e4ch, M.A. &"},{"key":"#cr-split#-34918.10","unstructured":"Ji, L. (Eds.) Water beetles of China. <\/em>Vol. II. Wien: Zoologisch-Botanische Gesellschaft in \u00d6sterreich and Wiener Coleopterologenverein, 371 pp.<\/p>

Kamite, Y. (2003) [Records of Satonius kurosawai<\/em> from Shikoku]. Heriguro<\/em>, 24, 82 (in Japanese).<\/p>

Ohta, Y., Kobayashi, N., Suzuki, S., Kato, T., Hori, S., Yamauchi, S. &"},{"key":"#cr-split#-34918.11","doi-asserted-by":"crossref","unstructured":"Katakura, H. (2009) Evolution of flight-muscle polymorphism in the dung beetle Phelotrupes laevistriatus Coleoptera: Geotrupidae): A phylogeographic analysis. Annals of the Entomological Society of America<\/em>, 102, 826-834.<\/p>

Roff, D.A. (1990) The evolution of flightlessness in insects. Ecological Monographs<\/em>, 60, 389-421.<\/p>

Saito, T. (2006) [Record of Satonius kurosawai<\/em> from Hyogo Prefecture]. Gekkan-mushi<\/em>, 430, 26 (in Japanese).<\/p>

Saito, M. &","DOI":"10.1603\/008.102.0509"},{"key":"#cr-split#-34918.12","unstructured":"Narukawa, N. (1986) [Remarkable beetles collected from Mie Prefecture]. Hirakura<\/em>, 30, 36-37 (in Japanese).<\/p>

Sat\u00f4, M. (1982) Discovery of Torridincolidae (Coleoptera) in Japan. Annotationes Zoologicae Japonenses<\/em>, 55, 276-283.<\/p>

Takai, Y. (1999) [Record of Satonius kurosawai<\/em> from Toyama Prefecture]. Gekkan-mushi<\/em>, 345, 47 (in Japanese).<\/p>

Yoshitomi, H. (1997) [Records of Delevea kurosawai<\/em> from Ch\u00fbbu district, Japan]. Coleopterists' News<\/em>, 117, 7 (in Japanese).<\/p>

Zera, A.J. &"},{"key":"#cr-split#-34918.13","doi-asserted-by":"crossref","unstructured":"Denno, R.F. (1997) Physiology and ecology of dispersal polymorphism in insects. Annual Review of Entomology<\/em>, <\/em>42, 207-230.<\/p>","DOI":"10.1146\/annurev.ento.42.1.207"}],"container-title":["Zootaxa"],"link":[{"URL":"https:\/\/mapress.com\/zt\/article\/download\/zootaxa.3016.1.4\/20577","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/mapress.com\/zt\/article\/download\/zootaxa.3016.1.4\/20577","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,3,18]],"date-time":"2024-03-18T03:44:29Z","timestamp":1710733469000},"score":10.850109,"resource":{"primary":{"URL":"https:\/\/mapress.com\/zt\/article\/view\/zootaxa.3016.1.4"}},"issued":{"date-parts":[[2011,9,7]]},"references-count":13,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2011,9,7]]}},"URL":"https:\/\/doi.org\/10.11646\/zootaxa.3016.1.4","ISSN":["1175-5334","1175-5326"],"issn-type":[{"value":"1175-5334","type":"electronic"},{"value":"1175-5326","type":"print"}],"published":{"date-parts":[[2011,9,7]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:56:33Z","timestamp":1791266193712,"version":"4.1.0"},"reference-count":79,"publisher":"Pensoft Publishers","license":[{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["ASP"],"abstract":"\n Abstract<\/jats:bold>\n <\/jats:p>\n \n For the first time we provide embryological information for Archostemata and Myxophaga, two very small but phylogenetically crucial suborders of Coleoptera. For this purpose we used\n Tenomerga mucida<\/jats:italic>\n of Cupedidae, the largest archostematan family, and\n Satonius kurosawai<\/jats:italic>\n of Torridincolidae, the largest family of Myxophaga. The egg of\n T. mucida<\/jats:italic>\n is cylindrical with a well-developed micropylar projection on the anterior pole of the egg. A long embryo forms on the ventral side of the egg, and with its progressive growth the segmentation rapidly proceeds, with an embryogenesis of the long-germ type. The elongated embryo then immerses into the egg, and taking a position in the egg along its anteroposterior axis, it continues to develop until blastokinesis occurs. Undergoing this process, the embryo appears on the egg surface, and with the progressive definitive dorsal closure, it acquires its definitive configuration. The first instar larva, which lacks an egg tooth, hatches, biting the chorion open with its robust mandibles. The egg of\n S. kurosawai<\/jats:italic>\n is oval and lacks a micropylar projection. A small embryo forms on the ventral surface. It stretches along the egg surface, increasing its length, and successively forming the segments, with its caudal end passing the posterior end of the egg and anteriorly almost reaching near the anterior pole. In contrast to\n T. mucida<\/jats:italic>\n , the embryogenesis is of the short-germ type. With the blastokinesis taking place and the definitive dorsal closure proceeding, the embryo acquires its definitive configuration. The first instar larva hatches by tearing the chorion longitudinally. Comparing the information obtained such as regarding the micropylar projections and germ types with those of other groups of beetles and Neuropterida, we discuss the phylogenetic relationships of coleopteran suborders. Our embryological study does not fit well with the sister group relationship between Archostemata and Myxophaga as suggested by recent large-scale molecular phylogenetic studies, i.e., requiring additional evolutionary steps and differing profoundly in the developmental pattern. In contrast, they are fully compatible with patterns suggested based on morphology, especially \u201cArchostemata + (Adephaga + (Myxophaga + Polyphaga))\u201d.\n <\/jats:p>","DOI":"10.3897\/asp.84.e198736","type":"journal-article","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"page":"847-866","source":"Crossref","is-referenced-by-count":0,"title":["Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship"],"prefix":"10.3897","volume":"84","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[{"name":"Kayo Frontier High School, Gifu, Japan"},{"name":"Sugadaira Research Station, Mountain Science Center, University of Tsukuba, Nagano, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[{"name":"Unaffiliated, Sayama, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[{"name":"Institute of Zoology and Evolutionary Research, Jena, Germany"},{"name":"Entomology II Section, Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Frankfurt, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[{"name":"Institute of Evolution and Ecology, University of T\u00fcbingen, T\u00fcbingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[{"name":"IDEA Consultants Inc., Yokohama, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[{"name":"Sugadaira Research Station, Mountain Science Center, University of Tsukuba, Nagano, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"reference":[{"key":"198736_B1","unstructured":"Anderson DT (1972a) The development of hemimetabolous insects. 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[in Russian].","DOI":"10.1172\/JCI105825"}],"container-title":["Arthropod Systematics & Phylogeny"],"link":[{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/download\/pdf\/","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/download\/xml\/","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article_preview.php?id=198736&skip_redirect=1","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:34Z","timestamp":1791265114000},"score":10.360025,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":79,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736","ISSN":["1864-8312","1863-7221"],"issn-type":[{"value":"1864-8312","type":"electronic"},{"value":"1863-7221","type":"print"}],"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:56:33Z","timestamp":1791266193992,"version":"4.1.0"},"description":"Figure 15 Embryonic development of Satonius kurosawai (III). A, A\u2019, A\u201d Stage 9: A ventral view, A\u2019 lateral view, A\u2019\u2019 dorsal view. B, B\u2019, B\u2019\u2019 Stage 10: B ventral view, B\u2019 lateral view, B\u2019\u2019 dorsal view. C, C\u2019, C\u2019\u2019 Stage 11: C ventral view, C\u2019 lateral view, C\u2019\u2019 dorsal view. D, D\u2019 D\u2019 Stage 12: D ventral view, D\u2019 lateral view, D\u2019\u2019 dorsal view. \u2014 Abbreviations: Ab \u2014 abdomen; An \u2014 antenna; Cl \u2014 clypeus; DDC \u2014 definitive dorsal closure; E \u2014 eye; HC \u2014 head capsule; Hp \u2014 hypopharynx; L1\u20143 \u2014 first to third legs; Lb \u2014 labium; Lr \u2014 labrum; Md \u2014 mandible; Mx \u2014 maxilla; Pd \u2014 proctodaeum; Pp \u2014 pleuropodium; SDO \u2014 secondary dorsal organ; Se \u2014 serosa; SG \u2014 spiracular gill; T1\u20143 \u2014 first to third thoracic segments; I\u2014XI \u2014 abdominal segments I to XI. \u2014 Scale: 100 \u00b5m.","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure15","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 15 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. Arthropod Systematics &amp; Phylogeny 84: 847-866. https:\/\/doi.org\/10.3897\/asp.84.e198736"],"prefix":"10.3897","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"score":6.413023,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/element\/2\/111\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":0,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736.figure15","relation":{"is-component-of":[{"id-type":"doi","id":"10.3897\/asp.84.e198736","asserted-by":"object"}]},"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:55:49Z","timestamp":1791266149647,"version":"4.1.0"},"description":"Figure 9 SEMs of Tenomerga mucida embryos (III). Stage 8 just before hatching. A, A\u2019, A\u2019\u2019 Embryo just before hatching. A dorsal view, A\u2019 ventral view, A\u2019\u2019 lateral view. B Embryo tearing through the chorion. \u2014 Abbreviations: An \u2014 antenna; Ca \u2014 cardo; Ch \u2014 chorion; Cl \u2014 clypeus; HC \u2014 head capsule; L1\u20143 \u2014 first to third legs; LbP \u2014 labial palp; Lr \u2014 labrum; MP \u2014 micropylar projection; Mx \u2014 maxilla; MxP \u2014 maxillary palp; St \u2014 sitpes; T1\u20143 \u2014 first to third thoracic segments; TP \u2014 tergal process; I\u2014IX \u2014 abdominal segments I to IX. \u2014 Scales: 200 \u00b5m.","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure9","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 9 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. Arthropod Systematics &amp; Phylogeny 84: 847-866. https:\/\/doi.org\/10.3897\/asp.84.e198736"],"prefix":"10.3897","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:26Z","timestamp":1791265106000},"score":6.352232,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/element\/2\/16\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":0,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736.figure9","relation":{"is-component-of":[{"id-type":"doi","id":"10.3897\/asp.84.e198736","asserted-by":"object"}]},"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:55:22Z","timestamp":1791266122043,"version":"4.1.0"},"description":"Figure 10 First instar larva of Tenomerga mucida. \u2014 Abbreviations: HC \u2014 head capsule; L1\u20143 \u2014 first to third legs; TP \u2014 tergal process. \u2014 Scale: 500 \u00b5m.","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure10","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 10 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. Arthropod Systematics &amp; Phylogeny 84: 847-866. https:\/\/doi.org\/10.3897\/asp.84.e198736"],"prefix":"10.3897","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:27Z","timestamp":1791265107000},"score":6.321249,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/element\/2\/17\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":0,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736.figure10","relation":{"is-component-of":[{"id-type":"doi","id":"10.3897\/asp.84.e198736","asserted-by":"object"}]},"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:55:57Z","timestamp":1791266157746,"version":"4.1.0"},"description":"Figure 17 Mapping the embryological information on the proposed phylogenies of coleopteran suborders. A Most widely accepted topology based on comparative morphology and biological information, formulated as \u201cArchostemata + (Adephaga + (Myxophaga + Polyphaga))\u201d. B A topology suggested in a study on the prothoracic features, formulated as \u201c(Archostemata + Adephaga) + (Myxophaga + Polyphaga)\u201d. C A topology based on a comparative morphology of the hindwings, formulated as \u201cPolyphaga + (Archostemata + (Myxophaga + Adephaga)\u201d. D, E Topologies based on recent large-scale molecular phylogenetic studies, formulated as \u201cAdephaga + ((Archostemata + Myxophaga) + Polyphaga)\u201d or as \u201cPolyphaga + ((Archostemata + Myxophaga) + Adephaga)\u201d. See the text (4.3.).","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure17","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 17 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. Arthropod Systematics &amp; Phylogeny 84: 847-866. https:\/\/doi.org\/10.3897\/asp.84.e198736"],"prefix":"10.3897","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:26Z","timestamp":1791265106000},"score":6.31398,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/element\/2\/113\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":0,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736.figure17","relation":{"is-component-of":[{"id-type":"doi","id":"10.3897\/asp.84.e198736","asserted-by":"object"}]},"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:56:34Z","timestamp":1791266194121,"version":"4.1.0"},"description":"Figure 8 SEMs of Tenomerga mucida embryos (II). A. A\u2019 Stage 5: A ventral view, A\u2019 lateral view. B, B\u2019 Stage 6: B ventral view, B\u2019 lateral view. Arrowheads show the spiracles. As for a star, see the text. \u2014 Abbreviations: An \u2014 antenna; Cllr \u2014 clypeolabrum; DDC \u2014 definitive dorsal closure; Ga \u2014 galea; HC \u2014 head capsule; HL \u2014 head lobe; L1\u20143 \u2014 first to third legs; La \u2014 lacinia; Lb \u2014 labium; LbP \u2014 labial palp; Lr \u2014 labrum; Md \u2014 mandible; Mx \u2014 maxilla; MxP \u2014 maxillary palp; Pd \u2014 proctodaeum; Pp \u2014 pleuropodium; Sd \u2014 stomodaeum; Se \u2014 serosa; St \u2014 sitpes; T1\u20143 \u2014 first to third thoracic segments; I\u2014XI \u2014 abdominal segments I to XI. \u2014 Scale: 200 \u00b5m.","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure8","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 8 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. Arthropod Systematics &amp; Phylogeny 84: 847-866. https:\/\/doi.org\/10.3897\/asp.84.e198736"],"prefix":"10.3897","author":[{"given":"Kazuki","family":"Kojima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukimasa","family":"Kobayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margarita I.","family":"Yavorskaya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Tsutsumi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichiro","family":"Machida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"2258","published-online":{"date-parts":[[2026,10,6]]},"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:26Z","timestamp":1791265106000},"score":6.3039913,"resource":{"primary":{"URL":"https:\/\/arthropod-systematics.arphahub.com\/article\/198736\/element\/2\/15\/"}},"issued":{"date-parts":[[2026,10,6]]},"references-count":0,"URL":"https:\/\/doi.org\/10.3897\/asp.84.e198736.figure8","relation":{"is-component-of":[{"id-type":"doi","id":"10.3897\/asp.84.e198736","asserted-by":"object"}]},"published":{"date-parts":[[2026,10,6]]}},{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:55:57Z","timestamp":1791266157750,"version":"4.1.0"},"description":"Figure 16 First instar larvae of Satonius kurosawai. A Just hatched first instar larva, with the egg exuvia. Arrow shows the crack opened in the chorion from which the larva emerged. B Dorsal view of the first instar larva. \u2014 Abbreviations: An \u2014 antenna; Ch \u2014 chorion; E \u2014 eye; L1 \u2014 first leg; SG \u2014 spiracular gill; T1\u20143 \u2014 first to third thoracic segments; Ug \u2014 urogomphus; I\u2014IX \u2014 abdominal segments I to IX. \u2014 Scales: 200 \u00b5m.","reference-count":0,"publisher":"Pensoft Publishers","content-domain":{"domain":[],"crossmark-restriction":false},"DOI":"10.3897\/asp.84.e198736.figure16","type":"component","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T05:38:25Z","timestamp":1791265105000},"source":"Crossref","is-referenced-by-count":0,"title":["Figure 16 from: Kojima K, Kobayashi Y, Beutel RG, Yavorskaya MI, Tsutsumi K, Machida R (2026) Embryonic development of Tenomerga mucida and Satonius kurosawai \u2013 First embryological information on Archostemata and Myxophaga (Coleoptera) does not support their sister group relationship. 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