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Within Gerromorpha, we explore the evolutionary transformations of tarsomeres based on detailed morphological documentation, to clarify the complex morphology and function of the legs. In this infraorder, plesiomorphic conditions of the tarsi are maintained in Mesoveliidae, Hebridae, Hydrometridae, \u201cMacroveliidae\u201d and Hermatobatidae, with apical claws and without specialized tarsal or pretarsal structures. In contrast, highly specialized deeply and symmetrically cleft apical mesotarsomeres, with blade-like claws and a feather-shaped empodium, are apomorphic features of Chenevelia stridulans and Rhagovelia antilleana (Veliidae). Using X-ray microtomography, we document to anatomy and infer the function of middle leg tarsomere swimming apparatus for Rhagovelia antilleana, Callivelia conata (Veliidae), Chenevelia stridulans, and Gerris buenoi (Gerridae). We also propose a hypothetical two-component explanation for the release of the fan in Rhagovelia and Chenevelia based on video and scan observations. For Rhagovelia and Chenevelia, the contraction of the femuro-pretarsalis muscle, which is attached by a very long and thin tendon to the conspicuously curved unguitractor plate, releases actively the leaf-like claws from the cleft, as well as the fan-shaped structure. The latter is folded in its resting position but unfolds passively outside of the cleft. The combination of deployable fan and modified claws is a locomotory system driven by a hybrid active-passive mechanism enabling species of these genera to improve their propulsion and colonize fast-running water habitats. These structural changes were a unique evolutionary innovation in Gerromorpha, enabling them to cope with specialized habitats and to access new ecological niches.<\/jats:p>","DOI":"10.1093\/icb\/icag139","type":"journal-article","created":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T11:44:00Z","timestamp":1785498240000},"source":"Crossref","is-referenced-by-count":0,"title":["Evolution of Distal Swimming Structures in Gerromorpha: Functional Use of Claws and Fan"],"prefix":"10.1093","volume":"66","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5398-6756","authenticated-orcid":false,"given":"Jordan","family":"Drapin","sequence":"first","affiliation":[{"name":"Institut de Recherche sur la Biologie de l\u2019Insecte (IRBI), CNRS UMR 7261, Universit\u00e9 Fran\u00e7ois Rabelais, Facult\u00e9 des Sciences et Techniques , 37200 Tours ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0433-7626","authenticated-orcid":false,"given":"Rolf","family":"Beutel","sequence":"additional","affiliation":[{"name":"Institut f\u00fcr Zoologie und Evolutionsforschung, FSU Jena , 07743 Jena ,","place":["Germany"]},{"name":"Entomology II, Senckenberg Forschungsinstitut und Naturmuseum Frankfurt , Senckenberganlage 25, 60325 Frankfurt ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mathieu","family":"Venin","sequence":"additional","affiliation":[{"name":"Laboratoire de M\u00e9canique Gabriel Lam\u00e9 (LaM\u00e9), EA 7494, Universit\u00e9 de Tours, Universit\u00e9 d\u2019Orl\u00e9ans, INSA CVL , 37200 Tours ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mirjam","family":"Urb","sequence":"additional","affiliation":[{"name":"\u00c9cole Normale Sup\u00e9rieure de Lyon , CNRS IGFL UMR 5242, Universit\u00e9 Lyon 1, 69000 Lyon ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Arnaud","family":"Badiane","sequence":"additional","affiliation":[{"name":"\u00c9cole Normale Sup\u00e9rieure de Lyon , CNRS IGFL UMR 5242, Universit\u00e9 Lyon 1, 69000 Lyon ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Quang","family":"Ngo\u00a0Xuan","sequence":"additional","affiliation":[{"name":"Graduate University of Science and Technology, Vietnam Academy of Science and Technology , 18 Hoang Quoc Viet Street, 11307 Ha Noi ,","place":["Vietnam"]},{"name":"Institute of Life Science, Vietnam Academy of Science and Technology , 85 Tran Quoc Toan Street, Ho Chi Minh City 700000 ,","place":["Vietnam"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abderrahman","family":"Khila","sequence":"additional","affiliation":[{"name":"\u00c9cole Normale Sup\u00e9rieure de Lyon , CNRS IGFL UMR 5242, Universit\u00e9 Lyon 1, 69000 Lyon ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1666-295X","authenticated-orcid":false,"given":"J\u00e9r\u00f4me","family":"Casas","sequence":"additional","affiliation":[{"name":"Institut de Recherche sur la Biologie de l\u2019Insecte (IRBI), CNRS UMR 7261, Universit\u00e9 Fran\u00e7ois Rabelais, Facult\u00e9 des Sciences et Techniques , 37200 Tours ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,7,31]]},"reference":[{"key":"2026082917411525400_bib1","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1093\/isd\/ixac002","article-title":"The skeletomuscular system of the mesosoma of formica rufa workers (Hymenoptera: formicidae)","volume":"6","author":"Aibekova","year":"2022","journal-title":"Insect Syst Diver"},{"key":"2026082917411525400_bib2","first-page":"7","article-title":"Fine structure of the body hair layers and morphology of the spiracles of semi-aquatic bugs (Hemiptera, Heteroptera, Gerromorpha) in relation to life on the water surface","volume":"140","author":"Andersen","year":"1976","journal-title":"Vidensk Medd fra Dansk Naturhist Foren"},{"key":"2026082917411525400_bib3","first-page":"337","article-title":"A comparative study of locomotion on the water surface in semi-aquatic bugs (Hemiptera, Heteroptera, Gerromorpha)","volume":"139","author":"Andersen","year":"1976","journal-title":"Vidensk Medd fra Dansk Naturhist Foren"},{"key":"2026082917411525400_bib4","article-title":"Water-striders (Hemiptera: gerridae, Veliidae, etc.)","volume-title":"Marine insects","author":"Andersen","year":"1976"},{"key":"2026082917411525400_bib5","doi-asserted-by":"publisher","DOI":"10.1163\/9789004631267","volume-title":"The semiaquatic bugs (Hemiptera: Gerromorpha)","author":"Andersen","year":"1982"},{"key":"2026082917411525400_bib6","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1163\/187631283X00254","article-title":"The old world microveliinae (Hemiptera: veliidae) I. 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