{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T01:43:31Z","timestamp":1782265411493,"version":"3.54.5"},"reference-count":83,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2020,3,27]],"date-time":"2020-03-27T00:00:00Z","timestamp":1585267200000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2020,3,27]],"date-time":"2020-03-27T00:00:00Z","timestamp":1585267200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000076","name":"Directorate for Biological Sciences","doi-asserted-by":"publisher","award":["DEB 165766"],"award-info":[{"award-number":["DEB 165766"]}],"id":[{"id":"10.13039\/100000076","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000076","name":"Directorate for Biological Sciences","doi-asserted-by":"crossref","award":["DEB 1555968"],"award-info":[{"award-number":["DEB 1555968"]}],"id":[{"id":"10.13039\/100000076","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Evol Biol"],"published-print":{"date-parts":[[2020,12]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:sec><jats:title>Background<\/jats:title><jats:p>Evolutionary transitions in temporal niche necessitates specialized morphology, physiology, and behaviors. Diurnal, heliothermic squamates (lizards and snakes) that bask require protection from ultraviolet radiation (UV) that can damage internal organs such as the brain, viscera, and gonads. Many smaller squamates have accomplished this protection by hyperpigmentation of the peritoneum and subcutaneous dorsum. Typically, nocturnal species do not require these protections from ultraviolet light. However, some nocturnal species that exhibit extreme crypsis may be exposed to sunlight and UV and require some means of mediating that damage. One such species is<jats:italic>Gekko<\/jats:italic>(<jats:italic>Ptychozoon<\/jats:italic>)<jats:italic>kuhli<\/jats:italic>, a nocturnal, arboreal gecko that uses extreme crypsis to blend in with tree bark. Hiding motionless on tree trunks leaves geckos exposed to sunlight during the day. Thus, we predict that<jats:italic>G. kuhli<\/jats:italic>will have independently evolved a hyperpigmented phenotype. To investigate this hypothesized association between temporal niche, behavior, and morphology, we characterized adult subcutaneous pigment for eight gecko species and embryonic pigment accumulation for a subset of four of these species, exhibiting diverse temporal niche and thermoregulatory behaviors. We predicted that nocturnal\/potentially-heliothermic<jats:italic>G. kuhli<\/jats:italic>would exhibit hyperpigmentation of internal structures like that of diurnal\/heliothermic geckos. We further predicted that embryonic pigment accumulation of<jats:italic>G. kuhli<\/jats:italic>would resemble that of diurnal\/heliothermic as opposed to nocturnal\/thigmothermic geckos.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>We found that temporal niche and thermoregulatory behavior predicted the degree of subcutaneous pigment in the eight gecko species examined. We demonstrate that<jats:italic>G. kuhli<\/jats:italic>accumulates pigment extremely early in embryonic development, unlike a diurnal\/heliothermic gecko species, despite having a similar adult phenotype.<\/jats:p><\/jats:sec><jats:sec><jats:title>Conclusions<\/jats:title><jats:p>The evolution of hyperpigmentation in<jats:italic>G. kuhli<\/jats:italic>is likely an adaptation to limit damage from occasional daytime UV exposure caused by crypsis-associated basking behavior.<jats:italic>Gekko kuhli<\/jats:italic>achieves its hyperpigmented phenotype through a derived developmental pattern, not seen in any other lizard species investigated to date, suggesting novel temporal differences in the migration and\/or differentiation of reptilian neural crest derivatives.<\/jats:p><\/jats:sec>","DOI":"10.1186\/s12862-020-01604-9","type":"journal-article","created":{"date-parts":[[2020,3,27]],"date-time":"2020-03-27T10:02:47Z","timestamp":1585303367000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Distinct patterns of pigment development underlie convergent hyperpigmentation between nocturnal and diurnal geckos (Squamata: Gekkota)"],"prefix":"10.1186","volume":"20","author":[{"given":"Aaron H.","family":"Griffing","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tony","family":"Gamble","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aaron M.","family":"Bauer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2020,3,27]]},"reference":[{"key":"1604_CR1","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1086\/285538","volume":"142","author":"SC Adolph","year":"1993","unstructured":"Adolph SC, Porter WP. Temperature, activity, and lizard life histories. Am Nat. 1993;142:273\u201395.","journal-title":"Am Nat"},{"key":"1604_CR2","doi-asserted-by":"publisher","first-page":"1944","DOI":"10.1111\/evo.13284","volume":"71","author":"SR Anderson","year":"2017","unstructured":"Anderson SR, Wiens JJ. Out of the dark: 350 million years of conservatism and evolution in diel activity patterns of vertebrates. Evolution. 2017;71:1944\u201359.","journal-title":"Evolution"},{"key":"1604_CR3","doi-asserted-by":"publisher","first-page":"103","DOI":"10.2307\/1564015","volume":"23","author":"Z Arad","year":"1989","unstructured":"Arad Z, Raber P, Werner YL. Selected body temperature in diurnal and nocturnal forms of Ptyodactylus (Reptilia: Gekkoninae) in a photothermal gradient. J Herpetol. 1989;23:103\u20138.","journal-title":"J Herpetol"},{"key":"1604_CR4","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1083\/jcb.38.1.67","volume":"38","author":"JT Bagnara","year":"1968","unstructured":"Bagnara JT, Taylor JD, Hadley ME. The dermal chromatophore unit. J Cell Biol. 1968;38:67\u201379.","journal-title":"J Cell Biol"},{"key":"1604_CR5","first-page":"1","volume":"44","author":"T Barbour","year":"1912","unstructured":"Barbour T. A contribution to the zoogeography of the east Indian islands. Mem Mus Comp Zool. 1912;44:1\u2013203.","journal-title":"Mem Mus Comp Zool"},{"key":"1604_CR6","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1080\/21564574.1997.9649986","volume":"46","author":"AM Bauer","year":"1997","unstructured":"Bauer AM. Peritoneal pigmentation and generic allocation in the Chamaeleonidae. Afr J Herpetol. 1997;46:117\u201323.","journal-title":"Afr J Herpetol"},{"key":"1604_CR7","doi-asserted-by":"publisher","first-page":"eaam5979","DOI":"10.1126\/science.aam5979","volume":"362","author":"ZD Blount","year":"2018","unstructured":"Blount ZD, Lenski RE, Losos JB. Contingency and determinism in evolution: Replaying life\u2019s tape. Science. 2018;362:eaam5979.","journal-title":"Science"},{"key":"1604_CR8","doi-asserted-by":"crossref","DOI":"10.5962\/bhl.title.100797","volume-title":"The Fauna of British India, including Ceylon and Burma. Reptilia and Batrachia","author":"GA Boulenger","year":"1890","unstructured":"Boulenger GA. The Fauna of British India, including Ceylon and Burma. Reptilia and Batrachia. London: Taylor & Francis; 1890."},{"key":"1604_CR9","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1016\/0042-6989(77)90117-1","volume":"17","author":"J Bowmaker","year":"1977","unstructured":"Bowmaker J, Knowles A. The visual pigments and oil droplets of the chicken retina. Vis Res. 1977;17:755\u201364.","journal-title":"Vis Res"},{"key":"1604_CR10","volume-title":"Philippine lizards of the family Gekkonidae","author":"CW Brown","year":"1978","unstructured":"Brown CW, Alcala AC. Philippine lizards of the family Gekkonidae. Dumaguete City: Silliman University Press; 1978."},{"key":"1604_CR11","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1016\/j.ympev.2012.06.009","volume":"66","author":"RM Brown","year":"2012","unstructured":"Brown RM, Siler CD, Grismer LL, Das I, McGuire JA. Phylogeny and cryptic diversification in southeast Asian flying geckos. Mol Phylogenet Evol. 2012;66:351\u201361.","journal-title":"Mol Phylogenet Evol"},{"key":"1604_CR12","doi-asserted-by":"publisher","first-page":"94","DOI":"10.2307\/1929864","volume":"24","author":"LC Cole","year":"1943","unstructured":"Cole LC. Experiments on toleration of high temperature in lizards with reference to adaptive coloration. Ecology. 1943;24:94\u2013108.","journal-title":"Ecology"},{"key":"1604_CR13","first-page":"137","volume":"125","author":"B Collette","year":"1961","unstructured":"Collette B. Correlations between ecology and morphology in anoline lizards from Havana, Cuba and southern Florida. Bull Mus Comp Zool. 1961;125:137\u201362.","journal-title":"Bull Mus Comp Zool"},{"key":"1604_CR14","first-page":"28","volume":"48","author":"CJ Conroy","year":"2009","unstructured":"Conroy CJ, Papenfuss T, Parker J, Hahn NE. Use of Tricaine Methanesulfonate (MS222) for euthanasia of reptiles. J. Am. Assoc. lab. Anim. Sci. 2009;48:28\u201332.","journal-title":"Sci"},{"key":"1604_CR15","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/j.semcdb.2008.09.008","volume":"20","author":"CD Cooper","year":"2008","unstructured":"Cooper CD, Raible D. Mechanisms for reaching the differentiated state: insights from neural crested derived melanocytes. Semin Cell Dev Biol. 2008;20:105\u201310.","journal-title":"Semin Cell Dev Biol"},{"key":"1604_CR16","first-page":"265","volume":"85","author":"RB Cowles","year":"1944","unstructured":"Cowles RB, Bogert CM. A preliminary study of the thermal requirements of desert reptiles. Bull Am Mus Nat Hist. 1944;85:265\u201396.","journal-title":"Bull Am Mus Nat Hist"},{"key":"1604_CR17","doi-asserted-by":"crossref","unstructured":"Das I, Vijayakumar SP. New species of\u00a0Ptychozoon (Sauria: Gekkonidae) from the Nicobar archipelago, Indian Ocean. Zootaxa. 2009;2095:8\u201320.","DOI":"10.11646\/zootaxa.2095.1.2"},{"key":"1604_CR18","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1002\/dvdy.38","volume":"248","author":"RE Diaz","year":"2019","unstructured":"Diaz RE, Shylo NA, Roellig D, Bronner M, Trainor PA. Filling in the phylogenetic gaps: induction, migration, and differentiation of neural crest cells in a squamate reptile, the veiled chameleon (Chamaeleo calyptratus). Dev Dyn. 2019;248:709\u201327.","journal-title":"Dev Dyn"},{"key":"1604_CR19","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1146\/annurev.es.21.110190.000421","volume":"21","author":"WE Duellman","year":"1990","unstructured":"Duellman WE, Pianka ER. Biogeography of nocturnal insectivores: historical events and ecological filters. Ann Rev Ecol Syst. 1990;21:57\u201368.","journal-title":"Ann Rev Ecol Syst"},{"key":"1604_CR20","first-page":"421","volume":"115","author":"H-R Duncker","year":"1965","unstructured":"Duncker H-R. Vereilungsmuster der Organpigmentierung bei Echsen (Sauria) und ihre entwicklungsgeschichtlichen Aspspekte. Verh Anat Ges Wien. 1965a;115:421\u20138.","journal-title":"Verh Anat Ges Wien"},{"key":"1604_CR21","first-page":"469","volume":"115","author":"H-R Duncker","year":"1965","unstructured":"Duncker H-R. Organpigmentierungen der Echsen (Sauria), dargestellt bei der Familie der Geckos (Gekkoniden). Verh. Anat. Ges. Wien. 1965b;115:469\u201370.","journal-title":"Verh. Anat. Ges. Wien"},{"key":"1604_CR22","first-page":"1","volume":"40","author":"H-R Duncker","year":"1968","unstructured":"Duncker H-R. Die extracutanen Melanocyten der Echsen (Sauria). Ergeb Anat Entwicklungsgesch. 1968;40:1\u201355.","journal-title":"Ergeb Anat Entwicklungsgesch"},{"key":"1604_CR23","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1034\/j.1600-0749.2000.130502.x","volume":"13","author":"R Fugii","year":"2000","unstructured":"Fugii R. The regulation of motile activity in fish chromatophores. Pigment Cell Res. 2000;13:300\u201319.","journal-title":"Pigment Cell Res"},{"key":"1604_CR24","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1111\/bij.12536","volume":"115","author":"T Gamble","year":"2015","unstructured":"Gamble T, Greenbaum E, Jackman TR, Bauer AM. Into the light: diurnality has evolved multiple times in geckos. Biol J Linn Soc. 2015;115:896\u2013910.","journal-title":"Biol J Linn Soc"},{"key":"1604_CR25","volume-title":"The biology and evolution of Australian lizards","author":"AE Greer","year":"1989","unstructured":"Greer AE. The biology and evolution of Australian lizards. Chipping Norton: Surrey Beatty & Sons Pty; 1989."},{"key":"1604_CR26","doi-asserted-by":"publisher","first-page":"1070","DOI":"10.1002\/dvdy.72","volume":"248","author":"AH Griffing","year":"2019","unstructured":"Griffing AH, Sanger TJ, Daza JD, Nielsen SV, Pinto BJ, Stanley EL, Gamble T. Embryonic development of a parthenogenetic vertebrate, the mourning gecko (Lepidodactylus lugubris). Dev Dyn. 2019;248:1070\u201390.","journal-title":"Dev Dyn"},{"key":"1604_CR27","first-page":"230","volume":"49","author":"AH Griffing","year":"2018","unstructured":"Griffing AH, Sanger TJ, Matamoros IC, Nielsen SV, Gamble T. Protocols for husbandry and embryo collection of a parthenogenetic gecko, Lepidodactylus lugubris (Squamata: Gekkonidae). Herpetol Rev. 2018;49:230\u20135.","journal-title":"Herpetol Rev"},{"key":"1604_CR28","unstructured":"Grismer LL. Amphibians and reptiles of the Seribuat Archipelago, peninsular Malaysia. Frankfurt am Main: Edition Chimaira; 2011a."},{"key":"1604_CR29","volume-title":"Lizards of peninsular Malaysia, Singapore and their adjacent archipelagos. Frankfurt am Main: Edition Chimaira","author":"LL Grismer","year":"2011","unstructured":"Grismer LL. Lizards of peninsular Malaysia, Singapore and their adjacent archipelagos. Frankfurt am Main: Edition Chimaira; 2011b."},{"key":"1604_CR30","unstructured":"Grismer LL, Sukumaran J, Grismer JL, Youmans TM, Wood PL, Johnson R. Report on the herpetofauna of the Temengor Forest Reserve, Perak, West Malaysia. Hamadryad. 2004;29:15\u201332."},{"key":"1604_CR31","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1186\/1471-213X-14-29","volume":"14","author":"RA Guerra-Fuentes","year":"2014","unstructured":"Guerra-Fuentes RA, Daza JD, Bauer AM. The embryology of the retinal pigmented epithelium in dwarf geckos (Gekkota: Sphaerodactylinae): a unique developmental pattern. BMC Dev Biol. 2014;14:29.","journal-title":"BMC Dev Biol"},{"key":"1604_CR32","doi-asserted-by":"publisher","first-page":"994","DOI":"10.1098\/rsbl.2012.0648","volume":"8","author":"MP Heinicke","year":"2012","unstructured":"Heinicke MP, Greenbaum E, Jackman TR, Bauer AM. Evolution of gliding in southeast Asian geckos and other vertebrates is temporally congruent with dipterocarp forest development. Biol Lett. 2012;8:994\u20137.","journal-title":"Biol Lett"},{"key":"1604_CR33","volume-title":"Natural history of west Indian reptiles and amphibians","author":"RW Henderson","year":"2009","unstructured":"Henderson RW, Powell R. Natural history of west Indian reptiles and amphibians. Gainesville: University of Florida Press; 2009."},{"key":"1604_CR34","doi-asserted-by":"publisher","first-page":"210","DOI":"10.2307\/1375075","volume":"23","author":"JE Hill","year":"1942","unstructured":"Hill JE. A supposed adaptation against sunstroke in African diurnal rats. J Mammal. 1942;23:210.","journal-title":"J Mammal"},{"key":"1604_CR35","first-page":"42","volume":"28","author":"M Honda","year":"1997","unstructured":"Honda M, Hikida T, Araya K, Ota H, Nabjitabhata J, Hoi-Sen Y. Cosymbotus craspedotus (frilly gecko) and C. platyurus (flat-tailed gecko). Gliding behavior Herpetol Rev. 1997;28:42\u20133.","journal-title":"Gliding behavior Herpetol Rev"},{"key":"1604_CR36","doi-asserted-by":"publisher","first-page":"326","DOI":"10.1111\/j.1600-0749.2004.00174.x","volume":"17","author":"RN Kelsch","year":"2004","unstructured":"Kelsch RN. Genetics and evolution of pigment patterns in fish. Pigment Cell Res. 2004;17:326\u2013426.","journal-title":"Pigment Cell Res"},{"key":"1604_CR37","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1002\/jez.b.22630","volume":"324","author":"ER Khannoon","year":"2015","unstructured":"Khannoon ER. Developmental stages of the climbing gecko Tarentola annularis with specieal references to the claws, pad lamellae, and subdigital setae. J Exp Zool B. 2015;324:450\u201364.","journal-title":"J Exp Zool B"},{"key":"1604_CR38","first-page":"1","volume":"14","author":"LM Klauber","year":"1939","unstructured":"Klauber LM. Studies of reptile life in the arid southwest. Bull Zool Soc San Diego. 1939;14:1\u2013100.","journal-title":"Bull Zool Soc San Diego"},{"key":"1604_CR39","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1007\/BF01723483","volume":"202","author":"P Kr\u00fcger","year":"1924","unstructured":"Kr\u00fcger P, Kern H. Die physikalische und physioloische Bedeutung des Pigmentes bei Amphibien und Reptilien. Pfl\u00fcgers Arch Gesamte Physiol Menschen Tiere. 1924;202:119\u201338.","journal-title":"Pfl\u00fcgers Arch Gesamte Physiol Menschen Tiere"},{"key":"1604_CR40","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511897948","volume-title":"The neural crest","author":"NM Le Douarin","year":"1999","unstructured":"Le Douarin NM. The neural crest. Cambridge: Cambridge University Press; 1999."},{"key":"1604_CR41","doi-asserted-by":"publisher","first-page":"794","DOI":"10.1086\/665645","volume":"179","author":"O Levy","year":"2012","unstructured":"Levy O, Dayan T, Kronfeld-Schor N, Porter WP. Biophysical modeling of the temporal niche: from first principles to the evolution of activity patterns. Am Nat. 2012;179:794\u2013804.","journal-title":"Am Nat"},{"key":"1604_CR42","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1111\/azo.12245","volume":"100","author":"FC Lima","year":"2017","unstructured":"Lima FC, Py-Daniel TR, Sartori MR, Abe AS, dos Santos OP, Freitas LM, Pereira KF, Sebben A. Developmental staging table of the green iguana. Acta Zool. 2017;100:232\u201344.","journal-title":"Acta Zool"},{"key":"1604_CR43","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1007\/BF03050132","volume":"4","author":"BC Mahendra","year":"1936","unstructured":"Mahendra BC. Contributions to the bionomics, anatomy, reproduction and development of the Indian house-gecko, Hemidactylus flaviviridis R\u00fcppel. Part I Proc Indian Acad Sci B. 1936;4:250\u201381.","journal-title":"Part I Proc Indian Acad Sci B"},{"key":"1604_CR44","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1038\/nchembio.1548","volume":"10","author":"VT Natarajan","year":"2014","unstructured":"Natarajan VT, Ganju P, Ramkumar A, Grover R, Gokhale RS. Multifaceted pathways protect human skin from UV radiation. Nat Chem Biol. 2014;10:542\u201351.","journal-title":"Nat Chem Biol"},{"key":"1604_CR45","unstructured":"Noro M, Uejima A, Abe G, Manabe M, Tamura K. Normal developmental stages of the Madagascar ground gecko Paroedura pictus with special reference to limb morphogenesis. Dev. Dynam. 2009;238:100\u2013109."},{"key":"1604_CR46","doi-asserted-by":"publisher","first-page":"278","DOI":"10.3389\/fphys.2018.00278","volume":"9","author":"J Ollonen","year":"2018","unstructured":"Ollonen J, Da Silva FO, Mahlow K, Di-Po\u00ef N. Skull development, ossification pattern, an adult shape in the emerging lizard model organism Pogona vitticeps: a comparative analysis with other squamates. Front Physiol. 2018;9:278.","journal-title":"Front Physiol"},{"key":"1604_CR47","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1111\/pcmr.12332","volume":"28","author":"DM Parichy","year":"2015","unstructured":"Parichy DM, Spiewak JE. Origins of adult pigmentation: diversity in pigment stem cell lineages and implications for pattern evolution. Pigment Cell Melanoma Res. 2015;28:31\u201350.","journal-title":"Pigment Cell Melanoma Res"},{"key":"1604_CR48","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1002\/9780470987100.ch5","volume-title":"The pigmentary system: physiology and pathophysiology","author":"DM Parichy","year":"2006","unstructured":"Parichy DM, Reedy MV, Erickson CA. Regulation of melanoblast migration and differentiation. In: Nordland JJ, Boissy RE, Hearing VJ, King RA, Oetting WS, Ortonne J-P, editors. The pigmentary system: physiology and pathophysiology. 2nd ed. Oxford: Blackwell Publishing; 2006. p. 108\u201339.","edition":"2"},{"key":"1604_CR49","first-page":"36","volume":"28","author":"TY Pong","year":"1974","unstructured":"Pong TY. Camouflage in the flying gecko, Ptychozoon kuhli Stejn. Malay Nat J. 1974;28:36.","journal-title":"Malay Nat J"},{"key":"1604_CR50","doi-asserted-by":"publisher","first-page":"273","DOI":"10.2307\/1942325","volume":"37","author":"WP Porter","year":"1967","unstructured":"Porter WP. Solar radiation through the living body walls of vertebrates with emphasis on desert reptiles. Ecol Monogr. 1967;37:273\u201396.","journal-title":"Ecol Monogr"},{"key":"1604_CR51","first-page":"17","volume-title":"Biology of the Reptilia, Volume 12, Physiology C. Physiological Ecology","author":"FH Pough","year":"1982","unstructured":"Pough FH, Gans C. Temperature regulation and thermal relations: the vocabulary of reptilian thermoregulation. In: Gans C, Pough FH, editors. Biology of the Reptilia, Volume 12, Physiology C. Physiological Ecology. London, New York, Paris, San Diego, San Francisco, S\u00e3o Paulo, Sydney, Tokyo, and Toronto: Academic Press; 1982. p. 17\u201323."},{"key":"1604_CR52","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1002\/jmor.1051550105","volume":"155","author":"FH Pough","year":"1978","unstructured":"Pough FH, Kwiecinski G, Bemis W. Melanin deposits associated with the venom glands of snakes. J Morphol. 1978;155:63\u201372.","journal-title":"J Morphol"},{"key":"1604_CR53","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1002\/ar.23500","volume":"300","author":"TR Py-Daniel","year":"2017","unstructured":"Py-Daniel TR, Lima AKS, Lima FC, Pic-Taylor A, Pires OR, Sebben A. A staging table of post-ovipositional development for the south American collared lizard Tropidurus torquatus (Squamata: Tropiduridae). Anat Rec. 2017;300:277\u201390.","journal-title":"Anat Rec"},{"key":"1604_CR54","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1186\/1471-213X-10-52","volume":"10","author":"M Reyes","year":"2010","unstructured":"Reyes M, Zandberg K, Desmawati I, de Bellard ME. Emergence and migration of trunk neural crest cells in a snake, the California kingsnake (Lampropeltis getula californiae). BMC Dev Biol. 2010;10:52.","journal-title":"BMC Dev Biol"},{"key":"1604_CR55","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1002\/1097-010X(20001201)287:7<467::AID-JEZ2>3.0.CO;2-8","volume":"287","author":"B R\u00f6ll","year":"2000","unstructured":"R\u00f6ll B. Characterization of retinal oil droplets in diurnal geckos (Reptilia, Gekkonidae). J Exp Zool. 2000;287:467\u201376.","journal-title":"J Exp Zool"},{"key":"1604_CR56","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1007\/s001140100227","volume":"88","author":"B R\u00f6ll","year":"2001","unstructured":"R\u00f6ll B. Multiple origin of diurnality in geckos: evidence from eye lens crystallins. Naturwissenschaften. 2001a;88:293\u20136.","journal-title":"Naturwissenschaften"},{"key":"1604_CR57","doi-asserted-by":"publisher","first-page":"2043","DOI":"10.1016\/S0042-6989(01)00093-1","volume":"41","author":"B R\u00f6ll","year":"2001","unstructured":"R\u00f6ll B. Gecko vision \u2014 retinal organization, foveae and implications for binocular vision. Vis Res. 2001b;41:2043\u201356.","journal-title":"Vis Res"},{"key":"1604_CR58","doi-asserted-by":"crossref","DOI":"10.5962\/bhl.title.24239","volume-title":"The reptiles of the indo-Australian archipelago, volume 1","author":"N de Rooij","year":"1915","unstructured":"de Rooij N. The reptiles of the indo-Australian archipelago, volume 1. Leiden: EJ Brill; 1915."},{"key":"1604_CR59","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1111\/j.1096-3642.1979.tb01093.x","volume":"65","author":"AP Russell","year":"1979","unstructured":"Russell AP. The origin of parachuting locomotion in gekkonid lizards (Reptilia: Gekkonidae). Zool J Linnean Soc. 1979;65:233\u201349.","journal-title":"Zool J Linnean Soc"},{"key":"1604_CR60","doi-asserted-by":"publisher","first-page":"2180","DOI":"10.1111\/evo.12100","volume":"67","author":"TJ Sanger","year":"2013","unstructured":"Sanger TJ, Sherratt E, McGlothlin JW, Brodie ED III, Losos JB, Abzhanov A. Convergent evolution of sexual dimorphism in skull shape using distinct developmental strategies. Evolution. 2013;67:2180\u201393.","journal-title":"Evolution"},{"key":"1604_CR61","volume-title":"Amphibians and reptiles of North Africa","author":"HH Schleich","year":"1996","unstructured":"Schleich HH, K\u00e4stle W, Kabisch K. Amphibians and reptiles of North Africa. Koenigstein: Koeltz Scientific Publishers; 1996."},{"key":"1604_CR62","doi-asserted-by":"publisher","first-page":"9663","DOI":"10.11609\/jott.2866.8.14.9663-9666","volume":"8","author":"A de Silva","year":"2016","unstructured":"de Silva A, Madawala M, Bauer AM, Karunarathna S. Rediscovery of the frilled tail geck Hemidactylus platyurus (Schneider, 1792) in Sri Lanka after more than 160 years. J Threat Taxa. 2016;8:9663\u20136.","journal-title":"J Threat Taxa"},{"key":"1604_CR63","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1111\/j.0022-202X.2004.23528.x","volume":"124","author":"A Slominski","year":"2005","unstructured":"Slominski A, Wortsman J, Plonka PM, Schallreuter KU, Paus R, Tobin DJ. Hair and follicle pigmentation. J Investig Dermatol. 2005;124:13\u201321.","journal-title":"J Investig Dermatol"},{"key":"1604_CR64","unstructured":"Smith MA. The Fauna of British India including Ceylon and Burma. Reptilia & Amphibia, 2: Sauria. London: Taylor & Francis; 1935."},{"key":"1604_CR65","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1152\/physrev.00021.2004","volume":"85","author":"O Strauss","year":"2005","unstructured":"Strauss O. The retinal pigment epithelium in visual function. Physiol Rev. 2005;85:845\u201381.","journal-title":"Physiol Rev"},{"key":"1604_CR66","first-page":"687","volume":"44","author":"EH Taylor","year":"1963","unstructured":"Taylor EH. The lizards of Thailand. Univ Kansas Sci Bull. 1963;44:687\u20131077.","journal-title":"Univ Kansas Sci Bull"},{"key":"1604_CR67","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1111\/j.1096-3642.1950.tb01459.x","volume":"120","author":"MWF Tweedie","year":"1950","unstructured":"Tweedie MWF. The flying gecko, Ptychozoon kuhli Stejn. Proc Zool Soc Lond. 1950;120:13.","journal-title":"Proc Zool Soc Lond"},{"key":"1604_CR68","volume-title":"The Reptile Database","author":"P Uetz","year":"2019","unstructured":"Uetz P, Freed P, Ho\u0161ek J. The Reptile Database. 2019. Available at www.reptile-database.org. Accessed 5 Oct 2019."},{"key":"1604_CR69","first-page":"201","volume":"13","author":"T Ulber","year":"1991","unstructured":"Ulber T, Ulber E. Cosymbotus platyurus (Schneider). Sauria (Supp). 1991;13:201\u20134.","journal-title":"Sauria (Supp)"},{"key":"1604_CR70","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1038\/167183a0","volume":"167","author":"G Underwood","year":"1951","unstructured":"Underwood G. Reptilian retinas. Nature. 1951;167:571\u20135.","journal-title":"Nature"},{"key":"1604_CR71","first-page":"1","volume-title":"Biology of the Reptilia, volume 2, morphology B","author":"G Underwood","year":"1970","unstructured":"Underwood G. The eye. In: Gans C, Prasons TS, editors. Biology of the Reptilia, volume 2, morphology B. London and New York: Academic Press; 1970. p. 1\u201397."},{"key":"1604_CR72","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1007\/s00442-007-0764-5","volume":"153","author":"M Valeix","year":"2007","unstructured":"Valeix M, Chamaill\u00e9-Jammes S, Fritz H. Interference competition and temporal niche shifts: elephants and herbivore communities at waterholes. Oecologia. 2007;153:739\u201348.","journal-title":"Oecologia"},{"key":"1604_CR73","first-page":"464","volume":"33","author":"RS Vetter","year":"1977","unstructured":"Vetter RS, Brodie ED Jr. Background color selection and antipredator behavior of the flying gecko, Ptychozoon kuhli. Herpetologica. 1977;33:464\u20137.","journal-title":"Herpetologica"},{"key":"1604_CR74","first-page":"195","volume":"94","author":"WVD Vos","year":"2018","unstructured":"Vos WVD, Stein K, Di-Po\u00ef N, Bickelmann C, Penner J. 2018. Ontogeny of Hemidactylus (Gekkota, Squamata) with emphasis on the limbs. Zoostyst. Evol. 2018;94:195\u2013209.","journal-title":"Evol"},{"key":"1604_CR75","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.1126\/science.1188545","volume":"331","author":"DB Wake","year":"2011","unstructured":"Wake DB, Wake MH, Specht CD. Homoplasy: from detecting pattern to determining process and mechanism of evolution. Science. 2011;331:1032\u20135.","journal-title":"Science"},{"key":"1604_CR76","volume-title":"The vertebrate eye and its adaptive radiation","author":"GL Walls","year":"1942","unstructured":"Walls GL. The vertebrate eye and its adaptive radiation. Bloomfield: Cranbook Institute of Science, Cranbook Press; 1942."},{"key":"1604_CR77","first-page":"373","volume":"19","author":"F Werner","year":"1912","unstructured":"Werner F. Beitr\u00e4ge zur Anatomie einiger seltenerer Reptilien, mit besonderer Ber\u00fccksichtigung der Atmungsorgane. Arb Zool Inst Univ Wien. 1912;19:373\u2013424.","journal-title":"Arb Zool Inst Univ Wien"},{"key":"1604_CR78","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1002\/jmor.1051330104","volume":"133","author":"YL Werner","year":"1971","unstructured":"Werner YL. The ontogenetic development of the vertebrae in some gekkonoid lizards. J Morphol. 1971;133:41\u201392.","journal-title":"J Morphol"},{"key":"1604_CR79","doi-asserted-by":"publisher","first-page":"408","DOI":"10.2307\/1563624","volume":"12","author":"YL Werner","year":"1978","unstructured":"Werner YL, Goldblatt A. Body temperature in a basking gekkonid lizard, Ptyodactylus hasselquistii (Reptilia, Lacertilia, Gekkonidae). J Herpetol. 1978;12:408\u201311.","journal-title":"J Herpetol"},{"key":"1604_CR80","doi-asserted-by":"publisher","first-page":"1198","DOI":"10.1002\/ar.20945","volume":"292","author":"PAD Wise","year":"2009","unstructured":"Wise PAD, Vickaryous MK, Russell AP. An embryonic staging table for in ovo development of Eublepharis macularius, the leopard gecko. Anat Rec. 2009;292:1198\u2013212.","journal-title":"Anat Rec"},{"key":"1604_CR81","doi-asserted-by":"crossref","unstructured":"Wood PL, Guo X, Travers SL, Su Y-C, Olson KV, Bauer AM, Grismer LL, Siler CD, Moyle RG, Andersen MJ, Brown RM. Parachute geckos free fall into synonymy: Gekko phylogeny, and a new subgeneric classification, inferred from thousands of ultraconserved elements. Mol Phylogenet Evol. 2020;146:106731.","DOI":"10.1016\/j.ympev.2020.106731"},{"key":"1604_CR82","doi-asserted-by":"publisher","first-page":"27557","DOI":"10.1074\/jbc.R700026200","volume":"282","author":"Y Yamaguchi","year":"2007","unstructured":"Yamaguchi Y, Brenner M, Hearing VJ. The regulation of skin pigmentation. J Biol Chem. 2007;282:27557\u201361.","journal-title":"J Biol Chem"},{"key":"1604_CR83","first-page":"987","volume":"52","author":"R-N Zhao","year":"2017","unstructured":"Zhao R-N, Zhang L-Y, Wu PF, Ping J, Jin J-Y, Zhang YP. The morphological characteristics of embryos at different developmental stages in the Japanese gecko, Gekko japonicus. Chin J Zool. 2017;52:987\u201395.","journal-title":"Chin J Zool"}],"container-title":["BMC Evolutionary Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12862-020-01604-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/article\/10.1186\/s12862-020-01604-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12862-020-01604-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,8,2]],"date-time":"2024-08-02T23:32:58Z","timestamp":1722641578000},"score":1,"resource":{"primary":{"URL":"https:\/\/bmcevolbiol.biomedcentral.com\/articles\/10.1186\/s12862-020-01604-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,3,27]]},"references-count":83,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,12]]}},"alternative-id":["1604"],"URL":"https:\/\/doi.org\/10.1186\/s12862-020-01604-9","relation":{},"ISSN":["1471-2148"],"issn-type":[{"value":"1471-2148","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,3,27]]},"assertion":[{"value":"4 November 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 March 2020","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 March 2020","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"All work conducted at Marquette University as it relates to this study was done under approved Marquette University IACUC protocols (AR 279, AR 298, AR288). Marquette University has an AAALAC accredited Animal Care and Use Program and has the PHS Assurance on file with the Office of Laboratory Animal Welfare. All voucher specimens used in this study are stored at Marquette University (Milwaukee, WI USA). All animals, both field-collected and commercially obtained, were handled under Marquette University IACUC approved protocols (AR 279, AR 298, AR288). Individuals of <i>G. kuhli<\/i>, <i>G. badenii<\/i>, and <i>H. platyurus<\/i> used in this study and which produced embryonic material were acquired commercially. All other species were field-collected under the following collecting permits: <i>C. brevis<\/i> (Texas, 2018; SPR-0590-067), <i>H. frenatus<\/i> (Hawaii, 2017; EX-18-02), <i>L. lugubris<\/i> (Hawaii, 2018; EX-18-06), <i>P. laticauda<\/i> (Hawaii, 2018; EX-18-06), <i>S<\/i>. <i>leonardovaldesi<\/i> (Honduras, 2018; Resoluci\u00f3n-DE-MP-213-2018), and <i>S. macrolepis<\/i> (Puerto Rico, 2017; 2016-IC-091).","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"40"}}