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More than a century of paleobotanical, morphological, and phylogenetic research has still not resolved fundamental questions about leaves. Developmental genetic data are sparse in ferns, and comparative studies of lycophytes and seed plants have reached opposing conclusions on the conservation of a leaf developmental program.<\/jats:p><\/jats:list-item>\n\n<jats:list-item><jats:p>We performed phylogenetic and expression analyses of a leaf developmental regulator (<jats:italic>Class III HD\u2010Zip<\/jats:italic> genes; <jats:italic>C3HDZ<\/jats:italic>s) spanning lycophytes and ferns.<\/jats:p><\/jats:list-item>\n\n<jats:list-item><jats:p>We show that a duplication and neofunctionalization of <jats:italic>C3HDZ<\/jats:italic>s probably occurred in the ancestor of euphyllophytes, and that there is a common leaf developmental mechanism conserved between ferns and seed plants. We show <jats:italic>C3HDZ<\/jats:italic> expression in lycophyte and fern sporangia and show that <jats:italic>C3HDZ<\/jats:italic>s have conserved expression patterns during initiation of lateral primordia (leaves or sporangia). This expression is maintained throughout sporangium development in lycophytes and ferns and indicates an ancestral role of <jats:italic>C3HDZ<\/jats:italic>s in sporangium development.<\/jats:p><\/jats:list-item>\n\n<jats:list-item><jats:p>We hypothesize that there is a deep homology of all leaves and that a sporangium\u2010specific developmental program was coopted independently for the development of lycophyte and euphyllophyte leaves. This provides molecular genetic support for a paradigm shift in theories of lycophyte leaf evolution.<\/jats:p><\/jats:list-item>\n<\/jats:list>\n<\/jats:p>","DOI":"10.1111\/nph.14075","type":"journal-article","created":{"date-parts":[[2016,7,7]],"date-time":"2016-07-07T08:47:06Z","timestamp":1467881226000},"page":"745-758","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":67,"title":["Challenging the paradigms of leaf evolution: Class III HD\u2010Zips in ferns and lycophytes"],"prefix":"10.1111","volume":"212","author":[{"given":"Alejandra","family":"Vasco","sequence":"first","affiliation":[{"name":"The New York Botanical Garden 2900 Southern Blvd Bronx NY 10458\u20105126 USA"},{"name":"Instituto de Biolog\u00eda Universidad Nacional Aut\u00f3noma de M\u00e9xico (UNAM) Mexico DF 04510 Mexico"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tynisha 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