Determinate growth is predominant and likely ancestral in squamate reptiles
Data files
Dec 14, 2020 version files 5.68 GB
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1_Anolis_allisoni.avi
1.09 MB
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10_Anolis_vermiculatus.avi
1.29 MB
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100_Gerrhosaurus_flavigularis.avi
1.41 MB
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101_Timon_tangitanus.avi
1.35 MB
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102_Timon_tangitanius_b.avi
88.44 MB
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102_Timon_tangitanus_b_reslice.avi
85.81 MB
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102_Timon_tangitanus.avi
1.23 MB
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103_Ouroborus_cataphractus.avi
1.06 MB
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104_Uromastyx_acanthinura_nigropunctata.avi
1.49 MB
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105_Uromastyx_acanthinura_nigropunctata.avi
2.10 MB
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106_Uromastyx_acanthinura_nigropunctata.avi
1.28 MB
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107_Uromastyx_acanthinura_nigropuctata.avi
1.09 MB
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108_Shinisaurus_crocodilurus.avi
1.46 MB
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11_Anolis_bartchi.rar
17.59 MB
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110_Cordylus_tropidosternum.avi
14.86 MB
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113_Heloderma_suspectum.avi
1.72 MB
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114_Varanus_indicus.avi
3.52 MB
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115_Varanus_indicus.avi
3.62 MB
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116_Gallotia_stehlini.avi
3.17 MB
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117_Gallotia_stehlini.avi
3.04 MB
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118_Gallotia_stehlini.avi
3.24 MB
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119_Gallotia_stehlini.avi
3.23 MB
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12_Furcifer_lateralis.avi
1.31 MB
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120_Timon_tangitanus_b_reslice.avi
89.76 MB
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120_Timon_tangitanus_b.avi
91.49 MB
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120_Timon_tangitanus.avi
3.41 MB
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121_Eublepharis_macularius.avi
2.94 MB
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122_Eublepharis_macularius.avi
3.08 MB
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123_Eublepharis_macularius.avi
2.82 MB
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124_Eublepharis_macularius.avi
2.79 MB
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125_Varanus_exanthematicus.avi
1.55 MB
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126_Varanus_exanthematicus.avi
1.52 MB
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127_Varanus_exanthematicus.avi
1.61 MB
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128_Varanus_exanthematicus.avi
1.58 MB
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129_Varanus_exanthematicus.avi
1.56 MB
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130_Varanus_exanthematicus.avi
1.31 MB
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131_Callopistes_flavipunctatus.avi
3.27 MB
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132_Varanus_exanthematicus.avi
1.08 MB
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133_Varanus_exanthematicus.avi
1.47 MB
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134_Varanus_exanthematicus.avi
1.34 MB
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135_Varanus_exanthematicus.avi
1.29 MB
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136_Varanus_exanthematicus.avi
1.52 MB
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137_Varanus_exanthematicus.avi
1.58 MB
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138_Leiocephalus_schreibersi.avi
1.28 MB
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139_Trachylepis_perrotetii.avi
1.54 MB
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140_Agama_agama.avi
1.45 MB
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141_Leiocephalus_personatus.avi
1.22 MB
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142_Anolis_roquet_roquet.avi
1.23 MB
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145_Anolis_equestris.avi
1.11 MB
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146_Reippeleon_brevicaudatus.avi
1.69 MB
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147_Trachylepis_quinquetaeniata.avi
1.09 MB
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149_Physignathus_cocincinus.avi
1.18 MB
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150_Intellagama_lesueurii.avi
1.02 MB
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151_Heloderma_suspectum.avi
1.17 MB
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152_Salvator_teguixin.avi
1.40 MB
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153_Iguana_iguana.avi
1.55 MB
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154_Zonosaurus_laticaudatus.avi
1.42 MB
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155_Gerrhosaurus_nigrolineatus.avi
1.38 MB
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156_Broadleysaurus_major.avi
1.33 MB
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157_Cordylus_tropidosternum.avi
1.20 MB
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158_Varanus_prasinus.avi
1.30 MB
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159_Varanus_beccarii.avi
1.20 MB
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16_Abronia_graminea.avi
23.64 MB
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161_Varanus_gouldi.avi
5.60 MB
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162_Varanus_salvator.avi
10.56 MB
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163_Tiliqua_gigas.avi
1.13 MB
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164_Broadleysaurus_major.avi
1.26 MB
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165_Chamaeleo_calyptratus.avi
1.44 MB
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167_Correlophus_cilliatus.bmp
2.63 MB
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17_Anolis_porcus.avi
2.58 MB
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172_Varanus_acanthurus.avi
6.87 MB
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176_Correlophus_ciliatus.avi
5.15 MB
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18_Anolis_porcus.avi
984.23 KB
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180_Hemidactylus_imbricatus.bmp
2.63 MB
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181_Eublepharis_angramainyu.avi
15.21 MB
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183_Gehyra_vorax.avi
14.51 MB
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184_Pogona_vitticeps.png
1.38 MB
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189_Aspidocelis_tigris.avi
6.77 MB
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19_Anolis_porcus.avi
1.38 MB
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190_Agama_aculeata.avi
7.21 MB
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191_Dasia_olivacea.avi
5.46 MB
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194_Furcifer_oustaleti.png
1.50 MB
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195_Acanthosaura_atricolis.avi
7.89 MB
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196_Latastia_longicauda.avi
6.12 MB
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197_Gehyra_vorax.avi
6.43 MB
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198_Chondrodactylus_bibroni.avi
7.86 MB
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199_Acanthosaura_capra.avi
6.57 MB
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2_Anolis_allisoni.avi
1.03 MB
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200_Chlamydosaurus_kingii.png
1.44 MB
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202_Tarentola_delalandi.avi
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203_Physignathus_coccincinus.bmp
2.63 MB
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204_Chamaeleo_calyptratus.png
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205_Takydromus_sexlineatus.avi
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206_Oplurus_cuvieri.avi
8.19 MB
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208_Cyrtodactylus_irianjayaensis.avi
6.83 MB
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209_Gekko_vittatus.avi
29.10 MB
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210_Draco_sumatranus_distal_femur.avi
37.95 MB
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211_Hemidacylus_frenatus.avi
17.66 MB
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212_Eutropis_multifasciatus.avi
15.03 MB
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213_Sceloporus_malachiticus.avi
14.74 MB
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214_Varanus_indicus.avi
5.11 MB
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215_Iguana_iguana_rhinolopha.avi
9.29 MB
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216_Varanus_gilleni_uCT_frontal.AVI
110.44 MB
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216_Varanus_gilleni_uCT_sagital.AVI
112.92 MB
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216_Varanus_gilleni_uCT_transversal.AVI
766.07 MB
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216_Varanus_gilleni_uRTG.AVI
23.48 MB
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217_Trioceros_rudis.tiff
2.27 MB
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218_Trioceros_rudis.png
1.08 MB
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219_Trioceros_johnstoni.avi
14.96 MB
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22_Anolis_porcus.avi
1.07 MB
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220_Chamaeleo_dilepis.tiff
2.32 MB
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221_Trioceros_hoehnelii.png
1.09 MB
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224_Chamaeleo_dilepis.png
1.79 MB
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225_Trioceros_melleri.png
1.61 MB
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226_Furcifer_pardalis.png
1.13 MB
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227_Calumma_parsonii_reslice-2.avi
40.01 MB
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227_Calumma_parsonii-2.avi
50.05 MB
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228_Kinyongia_fischeri.png
1.83 MB
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229_Trioceros_rudis.tiff
2.11 MB
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23_Anolis_barbatus.avi
2.55 MB
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232_Saara_loricata.avi
171.69 MB
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233_Uromastyx_aegyptia.png
1.39 MB
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234_Saara_loricata.png
861.83 KB
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235_Tarentola_chazaliae.avi
15.87 MB
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236_Varanus_acanthurus.png
607.82 KB
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237_Uromastyx_ornata.png
1.34 MB
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238_Uromastyx_ornatus.avi
13.89 MB
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239_Uromastyx_ornatus.tiff
2.27 MB
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24_Anolis_barbatus.avi
2.37 MB
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240_Teratoscincus_keyserlingi.bmp
2.63 MB
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241_Varanus_gilleni.bmp
2.63 MB
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242_Gerrhosaurus_flavigularis.bmp
2.63 MB
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243_Goniurosaurus_lui_b.bmp
2.63 MB
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247_Chamaeleo_calyptratus_reslice.avi
49.84 MB
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247_Chamaeleo_calyptratus.avi
50.85 MB
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249_Aeluroscalabotes_felinus_dist.bmp
2.63 MB
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25_Anolis_barbatus.avi
959.93 KB
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250_Cyrtodactylus_pequensis.avi
17.59 MB
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251_Cyclura_nubila.png
1.19 MB
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254_Oplurus_grandidieri.tiff
2.27 MB
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255_Anolis_garmani.tiff
2.11 MB
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256_Oplurus_fieriensis.png
779.18 KB
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257_Laemanctus_longipes.png
1.46 MB
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258_Cyclura_nubila_prox.bmp
2.63 MB
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259_Rhacodactylus_leachianus.avi
14.18 MB
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26_Anolis_barbatus.avi
2.32 MB
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260_Crotaphytus_collaris.tiff
2.38 MB
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261_Chamaeleo_dilepis.avi
12.02 MB
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264_Calumma_parsonii.avi
13.92 MB
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265_Sauromalus_ater.bmp
2.63 MB
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266_Trioceros_jacksonii.avi
16.10 MB
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267_Trioceros_jacksonii.avi
16.25 MB
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268_Trioceros_deremensis.tiff
2.04 MB
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269_Trioceros_deremensis.avi
15.41 MB
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27_Anolis_smallwoodi.avi
2.72 MB
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270_Furcifer_oustaleti.avi
20.46 MB
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271_Chamaeleo_dilepis.avi
27.29 MB
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273_Furcifer_lateralis.avi
15.40 MB
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275_Cyclura_nubila.zip
1.37 MB
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276_Chamaeleo_calyptratus.avi
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277_Gallotia_stehlini.tif
337.61 KB
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278_Hydrosaurus_weberi.tiff
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28_Anolis_smallwoodi.avi
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280_Bellatorius_frerei.avi
13.22 MB
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281_Paroedura_picta.bmp
2.63 MB
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282_Paroedura_picta.bmp
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284_Paroedura_picta.bmp
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285_Paroedura_picta.bmp
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286_Paroedura_picta.bmp
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287_Paroedura_picta.bmp
2.63 MB
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288_Paroedura_picta.bmp
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289_Paroedura_picta.bmp
2.63 MB
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290_Paroedura_picta.bmp
2.63 MB
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293_Leiolepis_ngovantrii.tiff
2.18 MB
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294_Tribolonotus_gracilis.avi
15.16 MB
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295_Tribolonotus_novaeguinae.avi
14.98 MB
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296_Leiocephalus_personatus.avi
6.86 MB
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297_Phelsuma_standingi.avi
14.58 MB
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298_Phelsuma_grandis.avi
15.99 MB
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299_Zonosaurus_karsteni_reslice.avi
95.78 MB
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299_Zonosaurus_karsteni.avi
97.96 MB
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3_Anolis_baracoae.avi
2.62 MB
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300_Aspidoscelis_deppei.avi
16.58 MB
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301_Gekko_gecko.tif
414.65 KB
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302_Takydromus_sexlineatus.avi
11.34 MB
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303_Anolis_equestris.tiff
2.13 MB
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306_Gekko_gecko.avi
14.61 MB
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307_Physignathus_coccincinus.avi
48.42 MB
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31_Uroplatus_henkeli.avi
1.51 MB
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310_Ameiva_ameiva.avi
13.37 MB
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311_Praoedura_pictus_b.bmp
2.63 MB
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312_Gallotia_galloti.tif
346.53 KB
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314_Correlophus_ciliatus_reslice.avi
60.31 MB
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314_Correlophus_ciliatus.avi
67.74 MB
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315_Algyroides_nigropunctatus.avi
17 MB
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316_Basiliscus_plumifrons.tiff
2.26 MB
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317_Trioceros_melleri.tiff
2.59 MB
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32_uroplatus_henkeli.avi
1.32 MB
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33_Uroplatus_henkeli.avi
1.44 MB
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1.41 MB
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357_Laemanctus_longipes.tif
421.41 KB
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358_Ptychozoon_kuhli_b.bmp
2.63 MB
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361_Calumma_malthe.tif
363.72 KB
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362_Hemitheconyx_caudicinctus.tif
372.12 KB
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365_Trioceros_ellioti.tif
321.33 KB
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366_Furcifer_verrucosus.tif
482.14 KB
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367_Brookesia_thieli.tif
272.05 KB
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368_Calumma_brevicorne.avi
15.11 MB
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369_Calumma_malthe.tif
351.97 KB
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372_Takydromus_sexlineatus.avi
17.99 MB
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373_Correlophus_ciliatus_reslice.avi
84.03 MB
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373_Correlophus_ciliatus.avi
85.81 MB
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374_Aeluroscalabotes_felinus.avi
16.16 MB
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375_Aeluroscalabotes_felinus_b.bmp
2.63 MB
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376_Eurydactylodes_agricolae.tif
215.31 KB
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378_Anolis_equestris.avi
13.80 MB
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379_Cordylus_tropidosternum_dist.bmp
2.63 MB
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383_Gerrhosaurus_flavigularis.bmp
2.63 MB
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385_Leiocephalus_personatus.avi
7.41 MB
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386_Dipsosaurus_dorsalis.avi
7.13 MB
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387_Tropidurus_hispidus.avi
16.58 MB
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389_Pristurus_somalicus_b.bmp
2.63 MB
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39_Uroplatus_lineatus.avi
1.33 MB
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391_Broadleysaurus_major.bmp
2.63 MB
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392_Uromastyx_ornata_reslice.avi
39.56 MB
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392_Uromastyx_ornatus.avi
40.29 MB
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4_Anolis_allisoni.avi
2.18 MB
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40_Uroplatus_lineatus.avi
1.17 MB
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41_Uroplatus_henkeli.avi
1.33 MB
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410_Zonosaurus_quadrilineatus.avi
13.79 MB
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414_Zonosaurus_quadrilineatus.bmp
2.63 MB
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416_Zonosaurus_quadrilineatus.bmp
2.63 MB
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417_Zonosaurus_karsteni.bmp
2.63 MB
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418_Zonosaurus_karsteni_reslice.avi
108.51 MB
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418_Zonosaurus_karsteni.avi
110.73 MB
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42_Phelsuma_cepediana.avi
17.32 MB
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420_Agama_somalica.avi
15.02 MB
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421_Crotaphytus_collaris.avi
8.94 MB
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422_Crotaphytus_collaris.avi
8.95 MB
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423_Paralaudakia_caucasia.avi
8.47 MB
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43_Timon_tangitanus.avi
2.16 MB
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432_Corytophanes_cristatus.avi
8.43 MB
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434_Podarcis_melissellensis.avi
33.92 MB
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435_Anolis_roquet_distal.avi
11.70 MB
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436_Scincus_scincus.avi
15.59 MB
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438_Tiliqua_scincoides.avi
13.35 MB
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439_Tiliqua_scincoides.avi
12.91 MB
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44_Varanus_boehmei.avi
2.58 MB
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440_Tiliqua_gigas.avi
13.80 MB
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441_Heloderma_exaspera_prox.bmp
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449_Timon_lepidus.avi
13.91 MB
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45_Varanus_macrei.avi
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451_Phelsuma_standingi.avi
14.73 MB
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452_Phelsuma_standingi_prox.bmp
2.63 MB
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48_Uromastyx_acanthinura_nigropunctata.avi
2.75 MB
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1.34 MB
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2.18 MB
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523_Varanus_pilbarensis.bmp
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53_Varanus_glauerti_uCT_frontal.AVI
105.99 MB
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53_Varanus_glauerti_uCT_sagital.AVI
106.45 MB
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53_Varanus_glauerti_uCT_transversal.AVI
738.89 MB
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53_Varanus_glauerti_uRTG.AVI
26.18 MB
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534_Eumeces_schneideri.avi
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537_Nephurus_levis.avi
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548_Podarcis_erhadii.avi
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565_Iguana_iguana_prox.bmp
2.63 MB
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571_Anolis_baracoae.avi
62.33 MB
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58_Leiocephalus_carinatus.avi
2.25 MB
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580_Leiocephalus_carinatus.avi
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581_Tropidurus_hispidus.avi
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587_Gekko_gecko_prox.bmp
2.63 MB
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588_Ptychozoon_kuhli.avi
15.95 MB
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16.10 MB
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597_Nephrurus_levis.avi
15.18 MB
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598_Hemidactylus_imbricatus.avi
17.19 MB
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599_Coleonyx_elegans.avi
16 MB
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1.13 MB
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2.22 MB
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600_Cyrtodactylus_pulchellus.avi
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607_Hemidactylus_imbricatus.avi
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608_Hemidactylus_imbricatus.bmp
2.63 MB
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2.63 MB
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612_Teratoscincus_keyserlingi.bmp
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15.32 MB
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629_Bellatorias_frerei.bmp
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2.12 MB
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632_Zonosaurus_quadrilineatus.bmp
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633_Tiliqua_gigas.avi
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16.25 MB
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637_Hemidactylus_platyurys.avi
18.02 MB
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639_Hemidactylus_platyurys.avi
17.04 MB
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64_Hemitheconyx_caudicintus.avi
2.05 MB
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642_Darevskia_unisexualis.bmp
2.63 MB
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644_Darevskia_unisexualis.avi
11.11 MB
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645_Darevskia_raddei_dist.bmp
2.63 MB
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646_Darevskia_raddei.bmp
2.63 MB
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648_Leiocephalus_carinatus.bmp
2.63 MB
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2.11 MB
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652_Eublepharis_macularius_prox.bmp
2.63 MB
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2.63 MB
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656_Tropidurus_hispidus.rar
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657_Eublepharis_macularius.bmp
2.63 MB
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659_Furcifer_pardalis_proxb.bmp
2.63 MB
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660_Pogona_barbata_prox.bmp
2.63 MB
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2.36 MB
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2.45 MB
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2.34 MB
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3.17 MB
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71_Varanus_salvadorii.avi
1.10 MB
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73_Furcifer_oustaleti.avi
1.31 MB
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List_of_specimens_dryad.xlsx
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Abstract
Body growth is typically thought to be indeterminate in ectothermic vertebrates. Indeed, until recently, this growth pattern was considered to be ubiquitous in ectotherms. Our recent observations of a complete growth plate cartilage (GPC) resorption, a reliable indicator of arrested skeletal growth, in many species of lizards clearly reject the ubiquity of indeterminate growth in reptiles and raise the question about the ancestral state of the growth pattern. Using X-ray micro-computed tomography (µCT), here we examined GPCs of long bones in three basally branching clades of squamate reptiles, namely in Gekkota, Scincoidea, and Lacertoidea. A complete loss of GPC, indicating skeletal growth arrest, was the predominant finding. Using a dataset of 164 species representing all major clades of lizards and the tuataras, we traced the evolution of determinate growth on the phylogenetic tree of Lepidosauria. The reconstruction of character states suggests that determinate growth is ancestral for the squamate reptiles (Squamata) and remains common in the majority of lizard lineages, while extended (potentially indeterminate) adult growth evolved several times within squamates. Although traditionally associated with endotherms, determinate growth is coupled with ectothermy in this lineage. These findings combined with existing literature suggest that determinate growth predominates in both extant and extinct amniotes.
We analyzed 194 femoral bones from lizards belonging to three clades of Squamata (Gekkota, Scincoidea, and Lacertoidea). We analyzed the growth plate cartilage (GPC) in the proximal part of the femur. We employed X-ray micro-computed tomography (µCT) and micro-radiography (µRTG) for vizualization of GPC. We scored the GPC state in a binary fashion, as either present (1) or absent (0). Ancestral state reconstruction was performed in R using the Hidden State Speciation and Extinction (HiSSE) model implemented in the R package hisse. Additionally, we performed the ancestral states reconstruction using maximum parsimony, as implemented in the R package castor. The reconstructed states were plotted using the R package phytools.