{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,4]],"date-time":"2026-11-04T03:12:07Z","timestamp":1793761927954,"version":"4.6.1"},"reference-count":132,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2019,3,15]],"date-time":"2019-03-15T00:00:00Z","timestamp":1552608000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100000001","name":"U.S. National Science Foundation","doi-asserted-by":"crossref","award":["DEB1657662"],"award-info":[{"award-number":["DEB1657662"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["9745-2008"],"award-info":[{"award-number":["9745-2008"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2019,7,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Recently-developed, molecularly-based phylogenies of geckos have provided the basis for reassessing the number of times adhesive toe-pads have arisen within the Gekkota. At present both a single origin and multiple origin hypotheses prevail, each of which has consequences that relate to explanations about digit form and evolutionary transitions underlying the enormous variation in adhesive toe pad structure among extant, limbed geckos (pygopods lack pertinent features). These competing hypotheses result from mapping the distribution of toe pads onto a phylogenetic framework employing the simple binary expedient of whether such toe pads are present or absent. It is evident, however, that adhesive toe pads are functional complexes that consist of a suite of integrated structural components that interact to bring about adhesive contact with the substratum and release from it. We evaluated the competing hypotheses about toe pad origins using 34 features associated with digit structure (drawn from the overall form of the digits; the presence and form of adhesive scansors; the proportions and structure of the phalanges; aspects of digital muscular and tendon morphology; presence and form of paraphalangeal elements; and the presence and form of substrate compliance-enhancing structures). We mapped these onto a well-supported phylogeny to reconstruct their evolution. Nineteen of these characters proved to be informative for all extant, limbed geckos, allowing us to assess which of them exhibit co-occurrence and\/or clade-specificity. We found the absence of adhesive toe pads to be the ancestral state for the extant Gekkota as a whole, and our data to be consistent with independent origins of adhesive toe pads in the Diplodactylidae, Sphaerodactylidae, Phyllodactylidae, and Gekkonidae, with a strong likelihood of multiple origins in the latter three families. These findings are consistent with recently-published evidence of the presence of adhesively-competent digits in geckos generally regarded as lacking toe pads. Based upon morphology we identify other taxa at various locations within the gekkotan tree that are promising candidates for the expression of the early phases of adhesively-assisted locomotion. Investigation of functionally transitional forms will be valuable for enhancing our understanding of what is necessary and sufficient for the transition to adhesively-assisted locomotion, and for those whose objectives are to develop simulacra of the gekkotan adhesive system for biotechnological applications.<\/jats:p>","DOI":"10.1093\/icb\/icz006","type":"journal-article","created":{"date-parts":[[2019,3,14]],"date-time":"2019-03-14T10:50:15Z","timestamp":1552560615000},"page":"131-147","source":"Crossref","is-referenced-by-count":48,"title":["Evolution of the Gekkotan Adhesive System: Does Digit Anatomy Point to One or More Origins?"],"prefix":"10.1093","volume":"59","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6659-6765","authenticated-orcid":false,"given":"Anthony P","family":"Russell","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, University of Calgary, 2500 University Drive NW, Calgary, Alberta, Canada T2N 1N4"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tony","family":"Gamble","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Marquette University, Milwaukee, WI 53201, USA"},{"name":"Bell Museum of Natural History, University of Minnesota, Saint Paul, MN 55113, USA"},{"name":"Milwaukee Public Museum, Milwaukee, WI 53233, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2019,4,27]]},"reference":[{"key":"2019072401000066400_icz006-B1","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1111\/j.1469-7998.1997.tb05755.x","article-title":"Morphogenesis of the digital pad lamellae in the embryo of the lizard Anolis lineatopus","volume":"243","author":"Alibardi","year":"1997","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B2","first-page":"63","article-title":"Structural niche, limb morphology and locomotion in lacertid lizards (Squamata: Lacertidae): a preliminary survey","volume":"64","author":"Arnold","year":"1998","journal-title":"Bull Brit Mus (Nat Hist) Ser"},{"key":"2019072401000066400_icz006-B3","doi-asserted-by":"crossref","first-page":"62","DOI":"10.11646\/zootaxa.1847.1.5","article-title":"A 100 million year old gecko with sophisticated adhesive toe pads, preserved in amber from Myanmar","volume":"1847","author":"Arnold","year":"2008","journal-title":"Zootaxa"},{"key":"2019072401000066400_icz006-B4","doi-asserted-by":"crossref","first-page":"67","DOI":"10.11646\/zootaxa.2984.1.3","article-title":"Homoplasy: from detecting pattern to determining process in evolution, but with a secondary role for morphology?","volume":"2984","author":"Assis","year":"2011","journal-title":"Zootaxa"},{"key":"2019072401000066400_icz006-B5","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1007\/978-3-540-31049-5_12","volume-title":"Biological adhesives","author":"Autumn","year":"2006"},{"key":"2019072401000066400_icz006-B6","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/35015073","article-title":"Adhesive force of a single gecko foot-hair","volume":"405","author":"Autumn","year":"2000","journal-title":"Nature"},{"key":"2019072401000066400_icz006-B7","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1146\/annurev-ecolsys-120213-091839","article-title":"Gecko adhesion as a model system for integrative biology, interdisciplinary science, and bioinspired engineering","volume":"45","author":"Autumn","year":"2014","journal-title":"Annu Rev Ecol Evol Syst"},{"key":"2019072401000066400_icz006-B8","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1002\/(SICI)1097-4687(199801)235:1<41::AID-JMOR4>3.0.CO;2-R","article-title":"Morphology of the adhesive tail tips of carphodactyline geckos (Reptilia: Diplodactylidae)","volume":"235","author":"Bauer","year":"1998","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B9","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1017\/S0952836904006259","article-title":"An Early Eocene gecko from Baltic amber and its implications for the evolution of gecko adhesion","volume":"265","author":"Bauer","year":"2005","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B10","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1111\/j.1558-5646.1959.tb03005.x","article-title":"Preadaptation and multiple evolutionary pathways","volume":"13","author":"Bock","year":"1959","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B11","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1007\/BF02703177","article-title":"Concepts and methods in ecomorphology","volume":"19","author":"Bock","year":"1994","journal-title":"J Biosci"},{"key":"2019072401000066400_icz006-B12","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/j.1558-5646.1965.tb01720.x","article-title":"Adaptation and the form\u2013function complex","volume":"19","author":"Bock","year":"1965","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B13","doi-asserted-by":"crossref","first-page":"69","DOI":"10.3389\/fevo.2018.00069","article-title":"The evolution of tarsal adhesive microstructures in stick and leaf insects (Phasmatoidea","volume":"6","author":"B\u00fcscher","year":"2018","journal-title":"Front Ecol Evol"},{"key":"2019072401000066400_icz006-B14","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1098\/rsif.2011.0254","article-title":"Sliding-induced non-uniform pre-tension governs robust and reversible adhesion: a revisit of adhesion mechanisms of geckos","volume":"9","author":"Cheng","year":"2012","journal-title":"J R Soc Interface"},{"key":"2019072401000066400_icz006-B201"},{"key":"2019072401000066400_icz006-B15","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/S0169-5347(98)01382-2","article-title":"Reconstructing ancestral character states: a critical reappraisal","volume":"13","author":"Cunningham","year":"1998","journal-title":"Trends Ecol Evol"},{"key":"2019072401000066400_icz006-B16","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1111\/j.1463-6395.2011.00521.x","article-title":"Anatomy of an enigma: an osteological investigation of the Namibian festive gecko (Narudasia festiva: Gekkonidae: Gekkota)","volume":"93","author":"Daza","year":"2012","journal-title":"Acta Zool"},{"key":"2019072401000066400_icz006-B17","doi-asserted-by":"crossref","first-page":"e1501080","DOI":"10.1126\/sciadv.1501080","article-title":"Mid-Cretaceous amber fossils illuminate the past diversity of tropical lizards","volume":"2","author":"Daza","year":"2016","journal-title":"Sci Adv"},{"key":"2019072401000066400_icz006-B202"},{"key":"2019072401000066400_icz006-B18","first-page":"15","article-title":"Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales","volume":"57","author":"Dem\u00e9r\u00e9","year":"2008","journal-title":"Syst Zool"},{"key":"2019072401000066400_icz006-B19","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1111\/j.1096-0031.2004.00047.x","article-title":"Convergence and parallelism: is a new life ahead of old concepts?","volume":"21","author":"Desutter-Grandcolas","year":"2005","journal-title":"Cladistics"},{"key":"2019072401000066400_icz006-B20","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1111\/j.1096-0031.2006.00136.x","article-title":"Distinguishing between convergence and parallelism is central to comparative biology: a reply to Williams and Ebach","volume":"23","author":"Desutter-Grandcolas","year":"2007","journal-title":"Cladistics"},{"key":"2019072401000066400_icz006-B21","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1371\/journal.pbio.0040088","article-title":"Relaxed phylogenetics and dating with confidence","volume":"4","author":"Drummond","year":"2006","journal-title":"PLoS Biol"},{"key":"2019072401000066400_icz006-B22","doi-asserted-by":"crossref","first-page":"1969","DOI":"10.1093\/molbev\/mss075","article-title":"Bayesian phylogenetics with BEAUti and the BEAST 1.7","volume":"29","author":"Drummond","year":"2012","journal-title":"Mol Biol Evol"},{"key":"2019072401000066400_icz006-B23","volume-title":"Complex adaptations in evolving populations.","author":"Frazzetta","year":"1975"},{"key":"2019072401000066400_icz006-B24","first-page":"127","article-title":"Modeling complex morphological change in evolution, and a possible ecological analogy","volume":"6","author":"Frazzetta","year":"1982","journal-title":"Evol Theory"},{"key":"2019072401000066400_icz006-B25","first-page":"128","volume-title":"Facsimile reprints in herpetology","author":"Fitzinger","year":"1843"},{"key":"2019072401000066400_icz006-B26","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.cretres.2017.11.003","article-title":"Cretaceous fossil gecko hand reveals a strikingly modern scansorial morphology: qualitative and biometric analysis of an amber-preserved lizard hand","volume":"84","author":"Fontanarrosa","year":"2018","journal-title":"Cret Res"},{"key":"2019072401000066400_icz006-B27","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1111\/j.1463-6409.2008.00330.x","article-title":"Out of the blue: a novel, trans-Atlantic clade of geckos (Gekkota, Squamata)","volume":"37","author":"Gamble","year":"2008","journal-title":"Zool Scripta"},{"key":"2019072401000066400_icz006-B28","doi-asserted-by":"crossref","first-page":"e39429.","DOI":"10.1371\/journal.pone.0039429","article-title":"Repeated origin and loss of adhesive toepads in geckos","volume":"7","author":"Gamble","year":"2012","journal-title":"PLoS ONE"},{"key":"2019072401000066400_icz006-B29","doi-asserted-by":"crossref","first-page":"896","DOI":"10.1111\/bij.12536","article-title":"Into the light: diurnality has evolved multiple times in geckos","volume":"115","author":"Gamble","year":"2015","journal-title":"Biol J Linn Soc"},{"key":"2019072401000066400_icz006-B30","doi-asserted-by":"crossref","first-page":"1429","DOI":"10.1111\/jeb.13097","article-title":"Repeated evolution of digital adhesion in geckos: a reply to Harrington and Reeder","volume":"30","author":"Gamble","year":"2017","journal-title":"J Evol Biol"},{"key":"2019072401000066400_icz006-B31","doi-asserted-by":"crossref","first-page":"2662","DOI":"10.1111\/jeb.12261","article-title":"Key innovations and island colonization as engines of evolutionary diversification: a comparative test with the Australasian diplodactyloid geckos","volume":"26","author":"Garcia-Porta","year":"2013","journal-title":"J Evol Biol"},{"key":"2019072401000066400_icz006-B32","doi-asserted-by":"crossref","first-page":"11278","DOI":"10.1021\/la202085j","article-title":"Shear adhesion strength of thermoplastic gecko-inspired synthetic adhesive exceeds material limits","volume":"27","author":"Gillies","year":"2011","journal-title":"Langmuir"},{"key":"2019072401000066400_icz006-B33","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1111\/j.1558-5646.2008.00505.x","article-title":"On phylogenetic tests of irreversible evolution","volume":"62","author":"Goldberg","year":"2008","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B34","doi-asserted-by":"crossref","first-page":"530","DOI":"10.1007\/s00114-001-0274-y","article-title":"Evolution of locomotory attachment pads in hexapods","volume":"88","author":"Gorb","year":"2001","journal-title":"Naturwissenschaften"},{"key":"2019072401000066400_icz006-B35","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1007\/BF00137351","article-title":"Dollo on Dollo\u2019s law: irreversibility and the status of evolutionary laws","volume":"3","author":"Gould","year":"1970","journal-title":"J Hist Biol"},{"key":"2019072401000066400_icz006-B36","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1111\/j.1096-0031.2004.00037.x","article-title":"Mapping characters on a tree with or without outgroups","volume":"20","author":"Grandcolas","year":"2004","journal-title":"Cladistics"},{"key":"2019072401000066400_icz006-B37","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1002\/jez.b.22614","article-title":"Ancestral state reconstructions require biological evidence to test evolutionary hypotheses: a case study examining the evolution of reproductive mode in squamate reptiles","volume":"324","author":"Griffith","year":"2015","journal-title":"J Exp Zool B Mol Dev Evol"},{"key":"2019072401000066400_icz006-B38","first-page":"273","article-title":"Die Bellgeckos des s\u00fcdlichen Afrika","volume":"194","author":"Haacke","year":"1974","journal-title":"Nat Mus"},{"key":"2019072401000066400_icz006-B39","first-page":"197","article-title":"The burrowing geckos of southern Africa. 1 (Reptilia: Gekkonidae)","volume":"29","author":"Haacke","year":"1975","journal-title":"Ann Transvaal Mus"},{"key":"2019072401000066400_icz006-B40","doi-asserted-by":"crossref","first-page":"2344","DOI":"10.1111\/evo.13318","article-title":"Tempo and mode of performance evolution across multiple independent origins of adhesive toe pads in lizards","volume":"71","author":"Hagey","year":"2017","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B41","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1111\/jeb.13004","article-title":"Rate heterogeneity across Squamata, misleading ancestral state reconstruction and the importance of proper null model specification","volume":"30","author":"Harrington","year":"2017","journal-title":"J Evol Biol"},{"key":"2019072401000066400_icz006-B42","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780198546412.001.0001","volume-title":"The comparative method in evolutionary biology","author":"Harvey","year":"1991"},{"key":"2019072401000066400_icz006-B43","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1186\/s12862-016-0846-2","article-title":"The measure of success: isolation promotes diversification in Pachydactylus geckos","volume":"17","author":"Heinicke","year":"2017","journal-title":"BMC Evol Biol"},{"key":"2019072401000066400_icz006-B44","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1111\/j.1463-6409.2007.00314.x","article-title":"Phylogeny of the Cyprinodontiformes (Teleostei, Atherinomorpha): the contribution of cranial soft tissue characters","volume":"37","author":"Hertwig","year":"2008","journal-title":"Zool Scripta"},{"key":"2019072401000066400_icz006-B45","first-page":"503","article-title":"On the origin of frictional adhesion in geckos: small morphological changes lead to a major biomechanical transition in the genus Gonatodes","volume":"120","author":"Higham","year":"2017","journal-title":"Biol J Linn Soc"},{"key":"2019072401000066400_icz006-B46","doi-asserted-by":"crossref","first-page":"809","DOI":"10.1073\/pnas.1418979112","article-title":"Adaptive simplification and the evolution of gecko locomotion: morphological and biomechanical consequences of losing adhesion","volume":"112","author":"Higham","year":"2015","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2019072401000066400_icz006-B47","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1080\/10635150390192780","article-title":"Stochastic mapping of morphological characters","volume":"52","author":"Huelsenbeck","year":"2003","journal-title":"Syst Biol"},{"key":"2019072401000066400_icz006-B48","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1111\/j.1095-8312.1996.tb01451.x","article-title":"A comparative analysis of clinging ability among pad-bearing lizards","volume":"59","author":"Irschick","year":"1996","journal-title":"Biol J Linn Soc"},{"key":"2019072401000066400_icz006-B49","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1080\/10635150490423962","article-title":"Accepting partnership by submission? Morphological phylogenetics in a molecular millennium","volume":"53","author":"Jenner","year":"2004","journal-title":"Syst Biol"},{"key":"2019072401000066400_icz006-B50","doi-asserted-by":"crossref","first-page":"937","DOI":"10.1111\/j.1469-7580.2009.01075.x","article-title":"Configuration of the setal fields of Rhoptropus (Gekkota; Gekkonidae): functional, evolutionary, ecological and phylogenetic implications of observed pattern","volume":"214","author":"Johnson","year":"2009","journal-title":"J Anat"},{"key":"2019072401000066400_icz006-B51","doi-asserted-by":"crossref","first-page":"1511","DOI":"10.1139\/z05-112","article-title":"Locomotor morphometry of the Pachydactylus radiation of lizards (Gekkota: Gekkonidae): a phylogenetically and ecologically informed analysis","volume":"83","author":"Johnson","year":"2005","journal-title":"Can J Zool"},{"key":"2019072401000066400_icz006-B52","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1111\/j.1096-0031.2006.00110.x","article-title":"Rejecting \u201cthe given\u201d in systematics","volume":"22","author":"Kearney","year":"2006","journal-title":"Cladistics"},{"key":"2019072401000066400_icz006-B53","doi-asserted-by":"crossref","first-page":"1677","DOI":"10.1098\/rspb.2004.2771","article-title":"Repeated evolution of limblessness and digging heads in worm lizards revealed by DNA from old bones","volume":"271","author":"Kearney","year":"2004","journal-title":"Proc R Soc B"},{"key":"2019072401000066400_icz006-B54","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1002\/jez.b.22630","article-title":"Developmental stages of the climbing gecko Tarentola annularis with special reference to the claws, pad lamellae, and subdigital setae","volume":"324","author":"Khannoon","year":"2015","journal-title":"J Exp Zool B Mol Dev Evol"},{"key":"2019072401000066400_icz006-B55","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1093\/icb\/43.3.361","article-title":"Introduction: the evolution of morphology, performance, and fitness","volume":"43","author":"Kingsolver","year":"2003","journal-title":"Integr Comp Biol"},{"key":"2019072401000066400_icz006-B56","volume-title":"Gekkonoid lizard taxonomy","author":"Kluge","year":"1993"},{"key":"2019072401000066400_icz006-B57","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1146\/annurev.ecolsys.27.1.501","article-title":"When does morphology matter?","volume":"27","author":"Koehl","year":"1996","journal-title":"Annu Rev Ecol Syst"},{"key":"2019072401000066400_icz006-B58","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.1111\/j.0014-3820.2006.tb00533.x","article-title":"Evidence for the reversibility of digit loss: a phylogenetic study of limb evolution in Bachia (Gymnophthalmidae: Squamata)","volume":"60","author":"Kohlsdorf","year":"2006","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B59","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1111\/j.1095-8312.2007.00893.x","article-title":"Evolution of novel jaw joints promote trophic diversity in coral reef fishes","volume":"93","author":"Konow","year":"2008","journal-title":"Biol J Linn Soc"},{"key":"2019072401000066400_icz006-B60","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1098\/rspb.2005.3390","article-title":"Footprints in the sand: independent reduction of subdigital lamellae in the Namib\u2013Kalahari burrowing geckos","volume":"273","author":"Lamb","year":"2006","journal-title":"Proc R Soc B"},{"key":"2019072401000066400_icz006-B61","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1002\/jmor.1051610307","article-title":"The extensor assembly in two species of opossum, Philander opossum and Didelphis marsupialis","volume":"161","author":"Landsmeer","year":"1979","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B62","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1002\/jmor.1051680305","article-title":"Digital morphology in Varanus and Iguana","volume":"168","author":"Landsmeer","year":"1981","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B63","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1038\/379812a0","article-title":"Correlated progression and the origin of turtles","volume":"379","author":"Lee","year":"1996","journal-title":"Nature"},{"key":"2019072401000066400_icz006-B64","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1046\/j.1463-6409.2000.00035.x","article-title":"Soft anatomy, diffuse homoplasy, and the relationships of lizards and snakes","volume":"29","author":"Lee","year":"2000","journal-title":"Zool Scripta"},{"key":"2019072401000066400_icz006-B65","doi-asserted-by":"crossref","first-page":"1441","DOI":"10.1111\/j.1558-5646.1998.tb02025.x","article-title":"Reptilian viviparity and Dollo\u2019s law","volume":"52","author":"Lee","year":"1998","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B66","doi-asserted-by":"crossref","first-page":"1827","DOI":"10.1111\/j.1558-5646.2011.01289.x","article-title":"Convergence, adaptation, and constraint","volume":"65","author":"Losos","year":"2011","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B67","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1111\/jzs.12226","article-title":"Systematics of the Saharo-Arabian clade of the Palearctic naked-toed geckos with the description of a new species of Tropiocolotes endemic to Oman","volume":"57","author":"Machado","year":"2018","journal-title":"J Zool Syst Evol Res"},{"key":"2019072401000066400_icz006-B68","doi-asserted-by":"crossref","first-page":"2219","DOI":"10.1111\/evo.13298","article-title":"Malagasy cichlids differentially limit impacts of body shape evolution on oral jaw functional morphology","volume":"71","author":"Martinez","year":"2017","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B69","doi-asserted-by":"crossref","first-page":"942","DOI":"10.1093\/icb\/38.6.942","article-title":"Morphological approaches to phylogeny","volume":"38","author":"Nielsen","year":"1998","journal-title":"Am Zool"},{"key":"2019072401000066400_icz006-B70","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1080\/10635150290102393","article-title":"Mapping mutations on phylogenies","volume":"51","author":"Nielsen","year":"2002","journal-title":"Syst Biol"},{"key":"2019072401000066400_icz006-B71","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1242\/jeb.080085","article-title":"Sticking to the story: outstanding challenges in gecko-inspired adhesives","volume":"219","author":"Niewiarowski","year":"2016","journal-title":"J Exp Biol"},{"key":"2019072401000066400_icz006-B72","doi-asserted-by":"crossref","DOI":"10.7208\/chicago\/9780226090009.001.0001","volume-title":"The comparative approach in evolutionary anthropology and biology","author":"Nunn","year":"2011"},{"key":"2019072401000066400_icz006-B73","doi-asserted-by":"crossref","first-page":"1343","DOI":"10.1111\/j.0014-3820.2003.tb00342.x","article-title":"Arboreality and morphological evolution in the ground beetles (Carabidae: Harpalinae): testing the taxon pulse model","volume":"57","author":"Ober","year":"2003","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B74","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1111\/j.1463-6409.2012.00545.x","article-title":"Phylogeny and systematics of Melanesia\u2019s most diverse gecko lineage (Cyrtodactylus, Gekkonidae, Squamata)","volume":"41","author":"Oliver","year":"2012","journal-title":"Zool Scripta"},{"key":"2019072401000066400_icz006-B75","doi-asserted-by":"crossref","first-page":"1948","DOI":"10.1111\/jeb.12709","article-title":"Repeated evolution of exaggerated dewlaps and other throat morphology in lizards","volume":"28","author":"Ord","year":"2015","journal-title":"J Evol Biol"},{"key":"2019072401000066400_icz006-B76","doi-asserted-by":"crossref","first-page":"662","DOI":"10.2307\/1445147","article-title":"Footprints of the Komodo monitor and the trackways of fossil reptiles","volume":"1984","author":"Padian","year":"1984","journal-title":"Copeia"},{"key":"2019072401000066400_icz006-B203"},{"key":"2019072401000066400_icz006-B77","author":"Paradis","year":"2018"},{"key":"2019072401000066400_icz006-B78","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1002\/ar.20706","article-title":"Subdigital setae of narrow-toed geckos, including a eublepharid (Aeluroscalabotes felinus)","volume":"291","author":"Peattie","year":"2008","journal-title":"Anat Rec"},{"key":"2019072401000066400_icz006-B79","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1007\/s00360-008-0310-8","article-title":"Functional demands of dynamic biological adhesion: an integrative approach","volume":"179","author":"Peattie","year":"2009","journal-title":"J Comp Physiol B"},{"key":"2019072401000066400_icz006-B80","doi-asserted-by":"crossref","first-page":"371","DOI":"10.2307\/1563590","article-title":"The evolution of the subdigital pad of Anolis. 2. Comparisons among the iguanid genera related to the anolines and a view from outside the radiation","volume":"17","author":"Peterson","year":"1983","journal-title":"J Herpetol"},{"key":"2019072401000066400_icz006-B81","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1002\/bies.20237","article-title":"Integrative biology of sticky feet in geckos","volume":"27","author":"Pianka","year":"2005","journal-title":"BioEssays"},{"key":"2019072401000066400_icz006-B82","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1111\/azo.12215","article-title":"Ontogenetic allometry of the digital rays of the Leopard gecko (Gekkota: Eublepharidae: Eublepharis macularius)","volume":"99","author":"Powell","year":"2018","journal-title":"Acta Zool"},{"key":"2019072401000066400_icz006-B83","doi-asserted-by":"crossref","first-page":"20160417","DOI":"10.1098\/rstb.2016.0417","article-title":"Phylogenetic tests for evolutionary innovation: the problematic link between key innovations and exceptional diversification","volume":"372","author":"Rabosky","year":"2017","journal-title":"Phil Trans R Soc B"},{"key":"2019072401000066400_icz006-B204"},{"key":"2019072401000066400_icz006-B85","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1111\/j.0014-3820.2005.tb00986.x","article-title":"Detecting the historical signature of key innovations using stochastic models of character evolution and caldogenesis","volume":"59","author":"Ree","year":"2005","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B86","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1111\/j.2041-210X.2011.00169.x","article-title":"phytools: an R package for phylogenetic comparative biology (and other things)","volume":"3","author":"Revell","year":"2012","journal-title":"Methods Ecol Evol"},{"key":"2019072401000066400_icz006-B87","doi-asserted-by":"crossref","first-page":"2898","DOI":"10.1111\/j.1558-5646.2007.00225.x","article-title":"A phylogenetic test for adaptive convergence in rock-dwelling lizards","volume":"61","author":"Revell","year":"2007","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B88","doi-asserted-by":"crossref","first-page":"e352","DOI":"10.1371\/journal.pbio.0040352","article-title":"Bushes in the tree of life","volume":"4","author":"Rokas","year":"2006","journal-title":"PLoS Biol"},{"key":"2019072401000066400_icz006-B89","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1002\/jmor.1052110302","article-title":"Development of the subdigital adhesive pads of Ptyodactylus guttatus (Reptilia: Gekkonidae)","volume":"211","author":"Rosenberg","year":"1992","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B90","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1111\/j.1469-7998.1975.tb03215.x","article-title":"A contribution to the functional analysis of the foot of the Tokay, Gekko gecko (Reptilia: Gekkonidae)","volume":"176","author":"Russell","year":"1975","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B91","first-page":"217","volume-title":"Morphology and biology of reptiles","author":"Russell","year":"1976"},{"key":"2019072401000066400_icz006-B92","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2307\/1443723","article-title":"Parallelism and integrated design in the foot structure of gekkonine and diplodactyline geckos","volume":"1979","author":"Russell","year":"1979","journal-title":"Copeia"},{"key":"2019072401000066400_icz006-B93","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1002\/jmor.1051690305","article-title":"Descriptive and functional anatomy of the digital vascular system in the Tokay, Gekko gecko","volume":"169","author":"Russell","year":"1981","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B94","doi-asserted-by":"crossref","first-page":"948","DOI":"10.1139\/z86-144","article-title":"The morphological basis of weight-bearing in the scansors of the tokay gecko (Reptilia: Sauria)","volume":"64","author":"Russell","year":"1986","journal-title":"Can J Zool"},{"key":"2019072401000066400_icz006-B95","doi-asserted-by":"crossref","first-page":"1154","DOI":"10.1093\/icb\/42.6.1154","article-title":"Integrative functional morphology of the gekkotan adhesive system (Reptilia: Gekkota)","volume":"42","author":"Russell","year":"2002","journal-title":"Integr Comp Biol"},{"key":"2019072401000066400_icz006-B96","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1111\/azo.12180","article-title":"The structure of anoline (Reptilia: Dactyloidae; Anolis) toe pads in relation to substratum conformity","volume":"98","author":"Russell","year":"2017","journal-title":"Acta Zool"},{"key":"2019072401000066400_icz006-B97","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1002\/jmor.1051970208","article-title":"Paraphalangeal elements of gekkonid lizards: a comparative survey","volume":"197","author":"Russell","year":"1988","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B98","first-page":"453","article-title":"Digit I in pad-bearing gekkonine geckos: alternate designs and the potential constraints of phalangeal number","volume":"29","author":"Russell","year":"1990","journal-title":"Mem Qld Mus"},{"key":"2019072401000066400_icz006-B99","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1111\/j.1469-7998.1990.tb03992.x","article-title":"Hypertrophied phalangeal chondroepiphyses in the gekkonid lizard genus Phelsuma: their structure and relation to the adhesive mechanism","volume":"221","author":"Russell","year":"1990","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B100","first-page":"113","article-title":"Underwood\u2019s classification of the geckos: a 21st century appreciation","volume":"68","author":"Russell","year":"2002","journal-title":"Bull Nat Hist Mus Zool Ser"},{"key":"2019072401000066400_icz006-B101","first-page":"1","volume-title":"Biology of the reptilian","author":"Russell","year":"2008"},{"key":"2019072401000066400_icz006-B102","first-page":"58","article-title":"Digital hyperextension in Anolis sagrei","volume":"57","author":"Russell","year":"2001","journal-title":"Herpetologica"},{"key":"2019072401000066400_icz006-B103","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1111\/jzo.12390","article-title":"Left in the dust: differential effectiveness of the two alternative adhesive pad configurations in geckos (Reptilia: Gekkota)","volume":"301","author":"Russell","year":"2017","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B104","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1002\/jmor.20646","article-title":"A whole lamella perspective on the origin of the epidermal free margin of Anolis (Reptilia: Dactyloidae) toe pads","volume":"278","author":"Russell","year":"2017","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B105","doi-asserted-by":"crossref","first-page":"3705","DOI":"10.1098\/rspb.2009.0946","article-title":"A new angle on clinging in geckos: incline, not substrate, triggers the deployment of the adhesive system","volume":"276","author":"Russell","year":"2009","journal-title":"Proc R Soc B"},{"key":"2019072401000066400_icz006-B106","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1111\/azo.12028","article-title":"Between a rock and a soft place: microtopography of the locomotor substrate and morphology of the setal fields of Namibian day geckos (Gekkota: Gekkonidae; Rhoptropus)","volume":"95","author":"Russell","year":"2014","journal-title":"Acta Zool"},{"key":"2019072401000066400_icz006-B107","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1111\/j.1469-7998.1997.tb05747.x","article-title":"Morphological correlates of the secondarily symmetrical pes of gekkotan lizards","volume":"241","author":"Russell","year":"1997","journal-title":"J Zool Lond"},{"key":"2019072401000066400_icz006-B108","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1002\/jmor.20420","article-title":"The evolution of digit form in Gonatodes (Gekkota: Sphaerodactylidae) and its bearing on the transition from frictional to adhesive contact in gekkotans","volume":"276","author":"Russell","year":"2015","journal-title":"J Morphol"},{"key":"2019072401000066400_icz006-B109","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1023\/B:JOMM.0000041190.83655.1d","article-title":"The tarsal complex of Afro-Malagasy Tenrecoidea: a search for phylogenetically meaningful characters","volume":"11","author":"Salton","year":"2004","journal-title":"J Mammal Evol"},{"key":"2019072401000066400_icz006-B110","doi-asserted-by":"crossref","first-page":"1699","DOI":"10.1111\/j.1558-5646.1997.tb05095.x","article-title":"Likelihood of ancestor states in adaptive radiation","volume":"51","author":"Schluter","year":"1997","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B111","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.tree.2015.10.007","article-title":"The evolving theory of evolutionary radiations","volume":"31","author":"Sim\u00f5es","year":"2016","journal-title":"Trends Ecol Evol"},{"key":"2019072401000066400_icz006-B112","first-page":"216","article-title":"Osteology, phylogeny, and functional morphology of two Jurassic lizard species and the early evolution of scansoriality in geckoes","volume":"180","author":"Sim\u00f5es","year":"2017","journal-title":"Zool J Linn Soc"},{"key":"2019072401000066400_icz006-B113","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1080\/10635150590906000","article-title":"Morphology\u2019s role in phylogeny reconstruction: perspectives from paleontology","volume":"54","author":"Smith","year":"2005","journal-title":"Syst Biol"},{"key":"2019072401000066400_icz006-B114","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1093\/icb\/34.4.502","article-title":"What molecular phylogenies tell us about the evolution of larval forms","volume":"34","author":"Strathmann","year":"1994","journal-title":"Am Zool"},{"key":"2019072401000066400_icz006-B115","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1111\/j.1420-9101.2009.01733.x","article-title":"Survival selection on escape performance and its underlying phenotypic traits: a case of many-to-one mapping","volume":"22","author":"Strobbe","year":"2009","journal-title":"J Evol Biol"},{"key":"2019072401000066400_icz006-B116","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/0025-5564(87)90074-5","article-title":"Reconstructing ancestral character states under Wagner parsimony","volume":"87","author":"Swofford","year":"1987","journal-title":"Math Biosci"},{"key":"2019072401000066400_icz006-B117","doi-asserted-by":"crossref","first-page":"19320","DOI":"10.1073\/pnas.0608841103","article-title":"Adhesion and friction in gecko toe attachment and detachment","volume":"103","author":"Tian","year":"2006","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2019072401000066400_icz006-B118","first-page":"549","article-title":"Ein Eidechsenschwanz mit Saugscheibe","volume":"19","author":"Tornier","year":"1899","journal-title":"Biol Zentralblatt"},{"key":"2019072401000066400_icz006-B119","author":"Uetz","year":"2018"},{"key":"2019072401000066400_icz006-B120","doi-asserted-by":"crossref","first-page":"195","DOI":"10.3897\/zse.94.22289","article-title":"Ontogeny of Hemidactylus (Gekkota, Squamata) with emphasis on the limbs","volume":"94","author":"van der Vos","year":"2018","journal-title":"Zoosyst Evol"},{"key":"2019072401000066400_icz006-B121","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1093\/icb\/45.2.256","article-title":"Many-to-one mapping of form to function: a general principle in organismal design?","volume":"45","author":"Wainwright","year":"2005","journal-title":"Integr Comp Biol"},{"key":"2019072401000066400_icz006-B122","doi-asserted-by":"crossref","DOI":"10.1002\/jmor.20834","article-title":"Are there general laws for digit evolution in squamates? The loss and re-evolution of digits in a clade of fossorial lizards (Brachymeles, Scincidae","author":"Wagner","year":"2018","journal-title":"J Morphol 279:1104\u201319"},{"key":"2019072401000066400_icz006-B123","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.1126\/science.1188545","article-title":"Homoplasy: from detecting pattern to determining process and mechanism of evolution","volume":"331","author":"Wake","year":"2011","journal-title":"Science"},{"key":"2019072401000066400_icz006-B124","doi-asserted-by":"crossref","first-page":"2632","DOI":"10.1111\/evo.13627","article-title":"Clade-specific evolutionary diversification along ontogenetic major axes in avian limb skeleton","volume":"72","author":"Watanabe","year":"2018","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B125","first-page":"123","article-title":"Why does a trait evolve multiple times within a clade? Repeated evolution of snakelike body form in squamate reptiles","volume":"60","author":"Wiens","year":"2006","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B126","doi-asserted-by":"crossref","first-page":"1886","DOI":"10.1111\/j.1558-5646.2007.00159.x","article-title":"Loss and re-evolution of complex life cycles in marsupial frogs: does ancestral state reconstruction mislead?","volume":"61","author":"Wiens","year":"2007","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B127","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1126\/science.215.4539.1509","article-title":"Convergent and alternative designs in the digital adhesive pads of scincid lizards","volume":"215","author":"Williams","year":"1982","journal-title":"Science"},{"key":"2019072401000066400_icz006-B128","doi-asserted-by":"crossref","first-page":"2480","DOI":"10.1111\/j.1558-5646.2007.00210.x","article-title":"Functional equivalence of morphologies enables morphological and ecological diversity","volume":"61","author":"Young","year":"2007","journal-title":"Evolution"},{"key":"2019072401000066400_icz006-B129","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1016\/j.ympev.2015.10.009","article-title":"Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species","volume":"94","author":"Zheng","year":"2016","journal-title":"Mol Phylogenet Evol"}],"container-title":["Integrative and Comparative Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/academic.oup.com\/icb\/advance-article-pdf\/doi\/10.1093\/icb\/icz006\/28537053\/icz006.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/academic.oup.com\/icb\/article-pdf\/59\/1\/131\/28849554\/icz006.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,7,15]],"date-time":"2024-07-15T16:14:29Z","timestamp":1721060069000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/icb\/article\/59\/1\/131\/5381544"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,4,27]]},"references-count":132,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,4,27]]},"published-print":{"date-parts":[[2019,7,1]]}},"URL":"https:\/\/doi.org\/10.1093\/icb\/icz006","relation":{},"ISSN":["1540-7063","1557-7023"],"issn-type":[{"value":"1540-7063","type":"print"},{"value":"1557-7023","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2019,7]]},"published":{"date-parts":[[2019,4,27]]}}}