{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T06:47:35Z","timestamp":1785739655346,"version":"3.56.0"},"reference-count":21,"publisher":"Wiley","issue":"2","license":[{"start":{"date-parts":[[2026,6,7]],"date-time":"2026-06-07T00:00:00Z","timestamp":1780790400000},"content-version":"vor","delay-in-days":6,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP24K09597"],"award-info":[{"award-number":["JP24K09597"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Evolution and Development"],"published-print":{"date-parts":[[2026,6]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    This study presents the first detailed histological description of wing primordium development in Mecoptera and demonstrates a late\u2010forming, non\u2010invaginated developmental mode lacking a morphologically distinct peripodial epithelium. Wing primordium development in holometabolous insects follows two major modes, early\u2010forming and late\u2010forming. Late\u2010forming primordia are epidermal cell populations that remain integrated within the larval epithelium, contribute to cuticle production during each larval molt, and begin morphogenesis only in the final instar. The order Mecoptera remains among the least studied groups of Holometabola with respect to wing primordium development. Using a newly established rearing method and histological analysis, we investigated wing development in the scorpionfly\n                    <jats:italic>Mavropanorpa japonica<\/jats:italic>\n                    and found the following: (1) wing primordia are absent in actively feeding larvae; (2) they originate as localized epidermal thickenings beneath the lateral pinacula approximately 20 days after cocoon formation; (3) thickening of wing primordia begins concurrently with apolysis, and the epithelium extends outward without pouch\u2010like invagination; and (4) no morphologically distinct peripodial epithelium was observed at all stages. Our findings show that wing primordia in Mecoptera develop externally, without invagination and without formation of a morphologically distinct peripodial epithelium. This study provides new comparative insight into heterochronic shifts and evolutionary diversity in holometabolous wing development.\n                  <\/jats:p>","DOI":"10.1111\/ede.70048","type":"journal-article","created":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T04:56:27Z","timestamp":1780894587000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Non\u2010Invaginated Wing Primordia in Holometabolous Insects: First Report From Mecoptera"],"prefix":"10.1111","volume":"28","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5532-7498","authenticated-orcid":false,"given":"Shuhei","family":"Niitsu","sequence":"first","affiliation":[{"name":"Department of Biological Sciences Tokyo Metropolitan University Hachioji Tokyo Japan"},{"name":"Department of Arts and Sciences International Christian University Mitaka Tokyo Japan"},{"name":"The University Museum The University of Tokyo Bunkyo\u2010ku Tokyo Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryo","family":"Ishihara","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Science Shinshu University Matsumoto Nagano Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takehiko","family":"Kamito","sequence":"additional","affiliation":[{"name":"Department of Arts and Sciences International Christian University Mitaka Tokyo Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,6,7]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"crossref","unstructured":"Beutel R. G. andF.Friedrich.2019 Phylogeny In Beutel R. G. and Friedrich F. (eds.) Nannomecoptera and Neomecoptera. DEGRUYTER Berlin. robopp 159\u2013161.","DOI":"10.1515\/9783110272543"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.33338\/ef.84480"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1304332110"},{"key":"e_1_2_9_5_1","first-page":"28","article-title":"When Do Butterfly Wings Form? The Old yet New Problem of Imaginal Discs (in Japanese)","volume":"30","author":"Hidaka T.","year":"1975","journal-title":"Shizen"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10164-022-00753-2"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00464226"},{"issue":"3","key":"e_1_2_9_8_1","first-page":"237","article-title":"Phylogeny of Endoperygota Insects, the Most Successful Lineage of Living Organisms","volume":"96","author":"Kristensen N. P.","year":"1999","journal-title":"European Journal of Entomology"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1111\/imb.12878"},{"issue":"1","key":"e_1_2_9_10_1","first-page":"1","article-title":"The Development of the Wings in Lepidoptera","volume":"8","author":"Mercer W. F.","year":"1900","journal-title":"Journal of the New York Entomological Society"},{"issue":"1","key":"e_1_2_9_11_1","first-page":"5","article-title":"The Development of Wings of Certain Beetles and Some Studies of the Origin of the Wings of Insects","volume":"13","author":"Powell P. B.","year":"1905","journal-title":"Journal of the New York Entomological Society"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/0040-8166(90)90067-J"},{"key":"e_1_2_9_13_1","first-page":"497","article-title":"Uber Zuchtung, Daipause und Postembryonale Entwicklung von Panorpa communis","volume":"83","author":"Rottmar B.","year":"1966","journal-title":"Zoologische Jahrb\u00fccher Abteilung f\u00fcr Anatomie und Ontogenie der Tiere"},{"key":"e_1_2_9_14_1","unstructured":"Sehnal F. P.Svacha andJ.Zrzavy 1996 Evolution of Insect Metamorphosis. In Gilbert L. I. Tata J. R. and Atkinson B. G. (eds.) Metamorphosis. Postembryonic Reprogramming of Gene Expression in Amphibian and Insect Cells. Academic Press San Diego etc. pp. 3\u201358."},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/0012-1606(92)90052-I"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF02161741"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.11646\/palaeoentomology.3.6.16"},{"key":"e_1_2_9_18_1","first-page":"517","article-title":"The Origin and Development of the Wings of Coleoptera","volume":"17","author":"Tower W. L.","year":"1903","journal-title":"Zoologische Jahrb\u00fccher Abteilung f\u00fcr Anatomie und Ontogenie der Tiere"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2019.10.009"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1038\/46737"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ento.47.091201.145230"},{"key":"e_1_2_9_22_1","first-page":"1","article-title":"Die Metamorphose von Corethra plumicornis","volume":"16","author":"Weismann A.","year":"1866","journal-title":"Zeitschrift f\u00fcr Wissenschaftliche Zoologie"}],"container-title":["Evolution &amp; Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ede.70048","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/ede.70048","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ede.70048","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T06:28:40Z","timestamp":1785738520000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/ede.70048"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6]]},"references-count":21,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2026,6]]}},"alternative-id":["10.1111\/ede.70048"],"URL":"https:\/\/doi.org\/10.1111\/ede.70048","archive":["Portico"],"relation":{},"ISSN":["1520-541X","1525-142X"],"issn-type":[{"value":"1520-541X","type":"print"},{"value":"1525-142X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6]]},"assertion":[{"value":"2025-10-16","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-05-27","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-06-07","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70048"}}