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Because pterosaurs were the earliest vertebrate lineage capable of powered flight and included the largest volant animal in the history of the earth, understanding how they evolved their flight apparatus, the wing, is an important issue in evolutionary biology. Herein, I speculate on the potential basis of pterosaur wing evolution using recent advances in the developmental biology of flying and non\u2010flying vertebrates. The most significant morphological features of pterosaur wings are: (<jats:italic>i<\/jats:italic>) a disproportionately elongated fourth finger, and (<jats:italic>ii<\/jats:italic>) a wing membrane called the brachiopatagium, which stretches from the posterior surface of the arm and elongated fourth finger to the anterior surface of the leg. At limb\u2010forming stages of pterosaur embryos, the zone of polarizing activity (<jats:styled-content style=\"fixed-case\">ZPA<\/jats:styled-content>) cells, from which the fourth finger eventually differentiates, could up\u2010regulate, restrict, and prolong expression of 5\u2032\u2010located <jats:italic>Homeobox D<\/jats:italic> (<jats:italic>Hoxd<\/jats:italic>) genes (e.g. <jats:italic>Hoxd11<\/jats:italic>, <jats:italic>Hoxd12<\/jats:italic>, and <jats:italic>Hoxd13<\/jats:italic>) around the <jats:styled-content style=\"fixed-case\">ZPA<\/jats:styled-content> through pterosaur\u2010specific exploitation of sonic hedgehog (<jats:styled-content style=\"fixed-case\">SHH<\/jats:styled-content>) signalling. 5\u2032<jats:italic>Hoxd<\/jats:italic> genes could then influence downstream bone morphogenetic protein (<jats:styled-content style=\"fixed-case\">BMP<\/jats:styled-content>) signalling to facilitate chondrocyte proliferation in long bones. Potential expression of <jats:italic>Fgf10<\/jats:italic> and <jats:italic>Tbx3<\/jats:italic> in the primordium of the brachiopatagium formed posterior to the forelimb bud might also facilitate elongation of the phalanges of the fourth finger. To establish the flight\u2010adapted musculoskeletal morphology shared by all volant vertebrates, pterosaurs probably underwent regulatory changes in the expression of genes controlling forelimb and pectoral girdle musculoskeletal development (e.g. <jats:italic>Tbx5<\/jats:italic>), as well as certain changes in the mode of cell\u2013cell interactions between muscular and connective tissues in the early phase of their evolution. Developmental data now accumulating for extant vertebrate taxa could be helpful in understanding the cellular and molecular mechanisms of body\u2010plan evolution in extinct vertebrates as well as extant vertebrates with unique morphology whose embryonic materials are hard to obtain.<\/jats:p>","DOI":"10.1111\/brv.12150","type":"journal-article","created":{"date-parts":[[2014,10,31]],"date-time":"2014-10-31T12:11:00Z","timestamp":1414757460000},"page":"1163-1178","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":14,"title":["How the pterosaur got its wings"],"prefix":"10.1111","volume":"90","author":[{"given":"Masayoshi","family":"Tokita","sequence":"first","affiliation":[{"name":"Department of Organismic and Evolutionary Biology Harvard University  16 Divinity Avenue Cambridge MA 02138 U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2014,10,31]]},"reference":[{"key":"e_1_2_8_2_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.22547"},{"key":"e_1_2_8_3_1","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:osobl\/9780199207114.001.0001"},{"key":"e_1_2_8_4_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.187807.112"},{"key":"e_1_2_8_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2014.03.030"},{"key":"e_1_2_8_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature12336"},{"key":"e_1_2_8_7_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0022336000034946"},{"key":"e_1_2_8_8_1","doi-asserted-by":"publisher","DOI":"10.1080\/08912969709386566"},{"key":"e_1_2_8_9_1","first-page":"191","volume-title":"Evolution and Palaeobiology of Pterosaurs","author":"Bennett S. 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