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Today, only male chondrichthyans possess claspers, which extend from posterior pelvic fins and function as intromittent organs. Here we report that clasper development from pelvic fins of male skates is controlled by hormonal regulation of the <jats:italic>Sonic hedgehog<\/jats:italic> (<jats:italic>Shh<\/jats:italic>) pathway. We show that Shh signalling is necessary for male clasper development and is sufficient to induce clasper cartilages in females. Androgen receptor (AR) controls the male-specific pattern of <jats:italic>Shh<\/jats:italic> in pelvic fins by regulation of <jats:italic>Hand2<\/jats:italic>. We identify an androgen response element (ARE) in the <jats:italic>Hand2<\/jats:italic> locus and present biochemical evidence that AR can directly bind the <jats:italic>Hand2<\/jats:italic> ARE. Together, our results suggest that the genetic circuit for appendage development evolved an androgen regulatory input, which prolonged signalling activity and drove clasper skeletogenesis in male fins.<\/jats:p>","DOI":"10.1038\/ncomms7698","type":"journal-article","created":{"date-parts":[[2015,4,14]],"date-time":"2015-04-14T09:46:27Z","timestamp":1429004787000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":52,"title":["Molecular development of chondrichthyan claspers and the evolution of copulatory organs"],"prefix":"10.1038","volume":"6","author":[{"given":"Katherine L.","family":"O\u2019Shaughnessy","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Randall D.","family":"Dahn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin J.","family":"Cohn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2015,4,14]]},"reference":[{"key":"BFncomms7698_CR1","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1038\/nature08176","volume":"460","author":"P Ahlberg","year":"2009","unstructured":"Ahlberg, P., Trinajstic, K., Johanson, Z. & Long, J. 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