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These anatomical characteristics have facilitated the adaptive radiation of chameleons to arboreal niches. To better understand the homology of chameleon carpal and tarsal elements, the process of syndactyly, cleft formation, and how modification of the mesopodial skeleton has played a role in the evolution and diversification of chameleons, we have studied the Veiled Chameleon (<jats:italic>Chamaeleo calyptratus<\/jats:italic>). We analysed limb patterning and morphogenesis through<jats:italic>in situ<\/jats:italic>hybridization,<jats:italic>in vitro<\/jats:italic>whole embryo culture and pharmacological perturbation, scoring for apoptosis, clefting, and skeletogenesis. Furthermore, we framed our data within a phylogenetic context by performing comparative skeletal analyses in 8 of the 12 currently recognized genera of extant chameleons.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>Our study uncovered a previously underappreciated degree of mesopodial skeletal diversity in chameleons. Phylogenetically derived chameleons exhibit a \u2018typical\u2019 outgroup complement of mesopodial elements (with the exception of centralia), with twice the number of currently recognized carpal and tarsal elements considered for this clade. In contrast to avians and rodents, mesenchymal clefting in chameleons commences in spite of the maintenance of a robust apical ectodermal ridge (AER). Furthermore,<jats:italic>Bmp<\/jats:italic>signaling appears to be important for cleft initiation but not for maintenance of apoptosis. Interdigital cell death therefore may be an ancestral characteristic of the autopodium, however syndactyly is an evolutionary novelty. In addition, we find that the pisiform segments from the ulnare and that chameleons lack an astragalus-calcaneum complex typical of amniotes and have evolved an ankle architecture convergent with amphibians in phylogenetically higher chameleons.<\/jats:p><\/jats:sec><jats:sec><jats:title>Conclusion<\/jats:title><jats:p>Our data underscores the importance of comparative and phylogenetic approaches when studying development. Body size may have played a role in the characteristic mesopodial skeletal architecture of chameleons by constraining deployment of the skeletogenic program in the smaller and earliest diverged and basal taxa. Our study challenges the \u2018re-evolution\u2019 of osteological features by showing that \u2018re-evolving\u2019 a \u2018lost\u2019 feature<jats:italic>de novo<\/jats:italic>(contrary to Dollo\u2019s Law) may instead be due to so called \u2018missing structures\u2019 being present but underdeveloped and\/or fused to other adjacent elements (cryptic features) whose independence may be re-established under changes in adaptive selective pressure.<\/jats:p><\/jats:sec>","DOI":"10.1186\/s12862-015-0464-4","type":"journal-article","created":{"date-parts":[[2015,9,14]],"date-time":"2015-09-14T23:23:45Z","timestamp":1442273025000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":53,"title":["Hand\/foot splitting and the \u2018re-evolution\u2019 of mesopodial skeletal elements during the evolution and radiation of chameleons"],"prefix":"10.1186","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9107-124X","authenticated-orcid":false,"suffix":"Jr.","given":"Raul E.","family":"Diaz","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul A.","family":"Trainor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2015,9,18]]},"reference":[{"key":"464_CR1","doi-asserted-by":"crossref","unstructured":"Sahni N, Yi S, Zhong Q, Jailkhani N, Charloteaux B, Cusick ME, et al. 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