Research Article Figures and figure supplements The Jurassic rise of squamates as supported by lepidosaur disparity and evolutionary rates Arnau Bolet et al Bolet et al. eLife 2022;11:e66511. DOI: https://doi.org/10.7554/eLife.66511  1 of 57 Research article Evolutionary Biology Pamelina polonica Icarosaurus siefkeri Kuehneosaurus latus niensis Marmoretta oxoniensis Scandensia Scanden ciervensis Gephyrosaurus bridensis Planocephalosaurus robinsonae Diphydontosaurus avonis Kallimodon cerinensis Priosphenodon avelasi Pamizinsaurus tlayuaensis Palaeopleurosaurus posidoniae Pleurosaurus goldfussi Bharatagama rebbanensis Sphenodon punctatus Cynosphenodon huizachalensis Clevosaurus brasiliensis Polysphenodon mulleri Clevosaurus mcgilli Clevosaurus bairdi Ankylosphenodon pachyostosus Bavarisaurus macrodactylus Huehuecuetzpalli mixtecus Hoyalacerta sanzi Polrussia mongoliensis Sineoamphisbaena hexatabularis Dibamus novaeguineae Anelytropsis papillosus Norellius nyctisaurops Gobekko cretacicus Pygopus lepidopus Eublepharis macularius Aeluroscalabotes felinus Hoburogekko suchanovi Phelsuma lineata Gekko gecko Sphaerodactylus elegans Oedura tryoni Saltuarius cornutus Nephrurus levis Paramacellodus oweni Paramacellodid UTAH Tepexisaurus tepexii Xantusia vigilis Cricosaura typica Lepidophyma flavimaculatum Palaeoxantusia kyrentos Becklesius hoffstetteri Yabeinosaurus tenuis Ornatocephalus metzleri Eolacerta robusta Globaura venusta Eoxanta lacertifrons Hymenosaurus clarki Parmeosaurus scutatus Slavoia darevskii Pseudosaurillus becklesi Platysaurus intermedius Tracheloptychus petersi Eumeces algeriensis Chalcides ocellatus Scincus scincus Plestiodon fasciatus Plestiodon anthracinus s Lygosoma bowringii Mabuya mabouya Mochlus sundevallii Tiliqua scincoides es Proscelotes eggeligeli Acontias plumbeus beus Feylinia currori r ri Typhlacontiass gracilis gr Takydromus s septentrionalis septent p trionalis Lacerta viridis iridis r Dracaenosaurus croizeti Cryptolacerta hassiaca Rhineura floridana Bipes biporus Cadea blanoides Blanus Dracaena guianensis s Tupinambis teguixin Teius teyou Aspidoscelis tigris Ameiva ameiva Bachia bicolor Gymnophthalmus underwoodi di Colobosaura modesta Dryadosaura nordestina Potamites ecpleopus Alopoglossus festae Liushusaurus acanthocaudata Palaeolacerta bavarica Bainguis parvus Myrmecodaptria microphagosa Jucaraseps grandipes Portugalophis lignites Najash rionegrina Epictia goudotii Typhlops jamaicensis ensis Typhlophis squamosus uamosus Liotyphlops albirostris albirostr bi triis Anomochilus leonardi Rhinophis sanguineus guineus i Parviraptor estesi Dinilysia patagonica Anilius scytale Cylindrophis ruffus Boa constrictor Pachyrhachis problematicus Haasiophis terrasanctus Meyasaurus diazromerali Carusia intermedia Restes rugosus Exostinus serratus Xenosaurus grandis dis Parasaniwa wyomingensis Dalinghosaurus longidigitus Merkurosaurus ornatus Shinisaurus crocodilurus dilurus r Bahndwivici ammoskius Peltosaurus granulosus Elgaria coerulea Dorsetisaurus purbeckensis Eosaniwa koehni Proplatynotia longirostrata Adriosaurus microbrachis Adriosaurus suessi Paravaranus angustifrons Eonatator sternbergi Aigialosaurus Parviderma inexacta Telmasaurus grangeri Lanthanotus borneensis Saniwa feisti Saniwa ensidens Varanus varius Psammosaurus griseus Saniwides mongoliensis Gobiderma pulchrum Primaderma nessovi Paraderma bogerti Palaeosaniwa canadensis Heloderma horridum Estesia mongoliensis Tchingisaurus multivagus Gobinatus arenosus Cherminsaurus kozlowskii Adamisaurus magnidentatus Erdenetesaurus robinsonae naee Polyglyphanodon odon sternbergi ste terrnbergii Tianyusaurus zhengigi Gilmoreteius chulsanensis nensis Gilmoreteius ferrugenous rrugenou us Darchansaurus estesi sii Chometokadmon fitzingeri Eichstaettisaurus schroederi Eichstaettisaurus gouldi Schoenesmahl dyspepsia Unnamed gen PMUR58 Ardeosaurus brevipes Morunasaurus M rrunasau Mo annularis Hoplocercus H l spinosus Saichangurvel davidsoni Ctenomastax parva Temujinia ellisoni Anchaurosaurus gilmorei Zapsosaurus sceliphros Liolaemus Stenocercus Microlophus albermalensis Plica plica Tropidurus torquatus Chalarodon madagascariensis Oplurus cyclurus Uta stansburiana Sceloporus Callisaurus draconoides Urosaurus nigricaudus Phrynosoma asio Polychrus gutturosus Enyalius bilineatus Gambelia Crotaphytus collaris Sauromalus ater Brachylophus Leiosaurus catamarcensis Anolis carolinensis Afairiguana avius Laemanctus serratus Priscagama gobiensis Phrynosomimus asper Mimeosaurus crassus Flaviagama dzerzhinskii Gladidenagama semiplena Arretosaurus ornatus Rieppeleon brachyurus Brookesia brygooi Uromastyx aegyptia Physignathus cocincinus Agama agama Guadalupian Paleocene Oligocene Lopingian Pliocene Miocene Eocene Middle Middle Upper Upper Upper Lower Lower Lower Permian Triassic Jurassic Cretaceous Paleogene Neogene 260 250 240 230 220 210 200 190 180 170 160 150 140 130 120 110 100 90 80 70 60 50 40 30 20 10 0 Figure 1. Lepidosaur— phylogeny, morphospace, disparity, and evolutionary rates. Phylogeny represented by a single randomly selected tree among those most parsimonious trees (MPTs) of the constrained analysis, and temporarily calibrated with the ‘Hedman’ method. Fossil ranges for each lineage are indicated according to the temporal distribution of the sampled taxa. For complete phylogenies and alternative datings, see Figure 1— figure supplements 1–7. Bolet et al. eLife 2022;11:e66511. DOI: https://doi.org/10.7554/eLife.66511  2 of 57 Research article Evolutionary Biology Pamelina polonica Icarosaurus siefkeri Kuehneosaurus latus Marmoretta oxoniensis Scandensia ciervensis Gephyrosaurus bridensis Planocephalosaurus robinsonae Diphydontosaurus avonis Kallimodon cerinensis Priosphenodon avelasi Pamizinsaurus tlayuaensis Palaeopleurosaurus posidoniae Pleurosaurus goldfussi Bharatagama rebbanensis Sphenodon punctatus Cynosphenodon huizachalensis Clevosaurus bairdi Clevosaurus mcgilli Ankylosphenodon pachyostosus Clevosaurus brasiliensis Polysphenodon mulleri Bavarisaurus macrodactylus Huehuecuetzpalli mixtecus Hoyalacerta sanzi Dibamus novaeguineae Anelytropsis papillosus Norellius nyctisaurops Gobekko cretacicus Pygopus lepidopus Eublepharis macularius Aeluroscalabotes felinus Hoburogekko suchanovi Phelsuma lineata Gekko gecko Sphaerodactylus elegans Oedura tryoni Saltuarius cornutus Nephrurus levis Paramacellodus oweni Paramacellodid UTAH Tepexisaurus tepexii Xantusia vigilis Cricosaura typica Lepidophyma flavimaculatum Palaeoxantusia kyrentos Pseudosaurillus becklesi Bainguis parvus Globaura venusta Eoxanta lacertifrons Parmeosaurus scutatus Hymenosaurus clarki Myrmecodaptria microphagosa Slavoia darevskii Palaeolacerta bavarica Platysaurus intermedius Tracheloptychus petersi Proscelotes eggeli Acontias plumbeus Feylinia currori Typhlacontias gracilis Eumeces algeriensis Chalcides ocellatus Scincus scincus Plestiodon fasciatus Plestiodon anthracinus Tiliqua scincoides Mochlus sundevalli Mabuya mabouya Lygosoma bowringii Takydromus septentrionalis Lacerta viridis Dracaenosaurus croizeti Rhineura floridana Bipes biporus Cadea blanoides Blanus Liushusaurus acanthocaudata Dracaena guianensis Tupinambis teguixin Teius teyou Aspidoscelis tigris Ameiva ameiva Bachia bicolor Gymnophthalmus underwoodi Colobosaura modesta Dryadosaura nordestina Potamites ecpleopus Alopoglossus festae Sineoamphisbaena hexatabularis Cryptolacerta hassiaca Jucaraseps grandipes Epictia goudotii Typhlops jamaicensis Typhlophis squamosus Liotyphlops albirostris Portugalophis lignites Anomochilus leonardi Rhinophis sanguineus Parviraptor estesi Najash rionegrina Dinilysia patagonica Anilius scytale Cylindrophis ruffus Boa constrictor Pachyrhachis problematicus Haasiophis terrasanctus Unnamed gen PMUR58 Becklesius hoffstetteri Meyasaurus diazromerali Yabeinosaurus tenuis Ornatocephalus metzleri Eolacerta robusta Carusia intermedia Restes rugosus Exostinus serratus Xenosaurus grandis Parasaniwa wyomingensis Dalinghosaurus longidigitus Merkurosaurus ornatus Shinisaurus crocodilurus Bahndwivici ammoskius Peltosaurus granulosus Elgaria coerulea Dorsetisaurus purbeckensis Eosaniwa koehni Proplatynotia longirostrata Adriosaurus microbrachis Adriosaurus suessi Paravaranus angustifrons Eonatator sternbergi Aigialosaurus Parviderma inexacta Gobiderma pulchrum Primaderma nessovi Paraderma bogerti Palaeosaniwa canadensis Heloderma horridum Estesia mongoliensis Saniwides mongoliensis Telmasaurus grangeri Lanthanotus borneensis Saniwa feisti Saniwa ensidens Varanus varius Psammosaurus griseus Tchingisaurus multivagus Gobinatus arenosus Cherminsaurus kozlowskii Adamisaurus magnidentatus Erdenetesaurus robinsonae Polyglyphanodon sternbergi Tianyusaurus zhengi Gilmoreteius chulsanensis Gilmoreteius ferrugenous Darchansaurus estesi Chometokadmon fitzingeri Ardeosaurus brevipes Schoenesmahl dyspepsia Eichstaettisaurus schroederi Eichstaettisaurus gouldi Morunasaurus annularis Hoplocercus spinosus Saichangurvel davidsoni Ctenomastax parva Temujinia ellisoni Anchaurosaurus gilmorei Zapsosaurus sceliphros Liolaemus Stenocercus Gambelia Crotaphytus collaris Sauromalus ater Brachylophus Microlophus albermalensis Polrussia mongoliensis Plica plica Tropidurus torquatus Chalarodon madagascariensis Oplurus cyclurus Sceloporus Uta stansburiana Phrynosoma asio Urosaurus nigricaudus Callisaurus draconoides Leiosaurus catamarcensis Anolis carolinensis Afairiguana avius Polychrus gutturosus Enyalius bilineatus Laemanctus serratus Priscagama gobiensis Phrynosomimus asper Mimeosaurus crassus Flaviagama dzerzhinskii Gladidenagama semiplena Arretosaurus ornatus Rieppeleon brachyurus Brookesia brygooi Uromastyx aegyptia Physignathus cocincinus Agama agama Holocene Guadalupian Paleocene Oligocene Lopingian Miocene Pleistocene Eocene Middle Middle Upper Upper Upper Lower Lower Lower Pliocene Permian Triassic Jurassic Cretaceous Paleogene Neogene Quaternary Figure 1—figure supplement 1. Single, randomly selected, most parsimonious tree among those resulting from the constrained analysis of lepidosaurs, dated using the equal method, and showing the position and ranges of nodes times using all three methods (Hedman in green, equal in blue, and MBL in red). Bolet et al. eLife 2022;11:e66511. DOI: https://doi.org/10.7554/eLife.66511  3 of 57 Research article Evolutionary Biology Pamelina polonica 99 Icarosaurus siefkeri 100 Kuehneosaurus latus Marmoretta oxoniensis Scandensia ciervensis 100 100 Gephyrosaurus bridensis 97 99 Planocephalosaurus robinsonae Diphydontosaurus avonis 100 100 Kallimodon cerinensis 100 Priosphenodon avelasi 100 Pamizinsaurus tlayuaensis 100 Palaeopleurosaurus posidoniae 100 100 Pleurosaurus goldfussi Bharatagama rebbanensis 100 Sphenodon punctatus 100 100 Cynosphenodon huizachalensis 100 Clevosaurus brasiliensis 100 Polysphenodon mulleri 70 Clevosaurus mcgilli Clevosaurus bairdi Ankylosphenodon pachyostosus 97 Bavarisaurus macrodactylus Huehuecuetzpalli mixtecus Hoyalacerta sanzi 100 Dibamus novaeguineae 97 Anelytropsis papillosus 100 Norellius nyctisaurops 100 Gobekko cretacicus 97 Pygopus lepidopus 100 100 Eublepharis macularius 97 Aeluroscalabotes felinus Hoburogekko suchanovi 70 100 Phelsuma lineata Gekko gecko 64 Sphaerodactylus elegans 97 100 Oedura tryoni 100 Saltuarius cornutus Nephrurus levis 100 Paramacellodus oweni Paramacellodid UTAH 97 Tepexisaurus tepexii 100 Xantusia vigilis 63 100 Cricosaura typica 100 Lepidophyma flavimaculatum Palaeoxantusia kyrentos Pseudosaurillus becklesi 74 63 63 71 Bainguis parvus 78 Globaura venusta 63 Eoxanta lacertifrons 74 Parmeosaurus scutatus 63 Hymenosaurus clarki 63 63 Myrmecodaptria microphagosa Slavoia darevskii 68 Palaeolacerta bavarica 100 Platysaurus intermedius Tracheloptychus petersi 63 89 Proscelotes eggeli 89 Acontias plumbeus 100 Feylinia currori 89 Typhlacontias gracilis 89 Eumeces algeriensis 89 Chalcides ocellatus 67 Scincus scincus 100 Plestiodon fasciatus 78 Plestiodon anthracinus 74 78 Mochlus sundevalli Tiliqua scincoides 79 Mabuya mabouya Lygosoma bowringii 100 Takydromus septentrionalis 98 Lacerta viridis 74 Dracaenosaurus croizeti 100 Rhineura floridana 100 Bipes biporus 74 100 Cadea blanoides Blanus Liushusaurus acanthocaudata 100 Dracaena guianensis 74 100 Tupinambis teguixin 100 Teius teyou 100 100 Aspidoscelis tigris Ameiva ameiva 100 Bachia bicolor 100 Gymnophthalmus underwoodi 100 Colobosaura modesta 88 Dryadosaura nordestina 100 Potamites ecpleopus Alopoglossus festae 74 Sineoamphisbaena hexatabularis 63 Cryptolacerta hassiaca 74 Jucaraseps grandipes 95 100 Epictia goudotii 96 Typhlops jamaicensis 98 Typhlophis squamosus 100 Liotyphlops albirostris Portugalophis lignites 74 100 Anomochilus leonardi 74 Rhinophis sanguineus Parviraptor estesi 74 74 Najash rionegrina 74 Dinilysia patagonica 100 Anilius scytale 94 Cylindrophis ruffus 63 100 Boa constrictor 100 Pachyrhachis problematicus Haasiophis terrasanctus Unnamed gen PMUR58 Meyasaurus diazromerali 51 Becklesius hoffstetteri 62 Yabeinosaurus tenuis 63 85 Ornatocephalus metzleri Eolacerta robusta Carusia intermedia 100 Restes rugosus 92 100 Exostinus serratus Xenosaurus grandis 92 Parasaniwa wyomingensis 100 Dalinghosaurus longidigitus 63 100 Merkurosaurus ornatus Clades 92 100 Shinisaurus crocodilurus Bahndwivici ammoskius 100 Peltosaurus granulosus 92 Elgaria coerulea Dorsetisaurus purbeckensis STEM LEPIDOSAUR 92 Eosaniwa koehni 100 Paravaranus angustifrons 100 Eonatator sternbergi 100 83 Aigialosaurus RHYNCHOCEPHALIAN 60 Proplatynotia longirostrata 62 Adriosaurus microbrachis 70 Adriosaurus suessi 57 Parviderma inexacta STEM SQUAMATE 60 Telmasaurus grangeri 60 60 Saniwa feisti 57 Lanthanotus borneensis GEKKOTAN 60 100 100 Saniwa ensidens Varanus varius Psammosaurus griseus 59 Saniwides mongoliensis SCINCID 100 100 Gobiderma pulchrum Primaderma nessovi 100 Paraderma bogerti 100 Palaeosaniwa canadensis CORDYLIFORM 100 Heloderma horridum Estesia mongoliensis 97 Tchingisaurus multivagus 97 Gobinatus arenosus XANTUSIID 100 Cherminsaurus kozlowskii Adamisaurus magnidentatus 100 70 Erdenetesaurus robinsonae 100 Polyglyphanodon sternbergi TEIOID 74 100 Tianyusaurus zhengi 84 Gilmoreteius chulsanensis 60 Gilmoreteius ferrugenous Darchansaurus estesi LACERTID 74 Chometokadmon fitzingeri 74 Ardeosaurus brevipes Schoenesmahl dyspepsia 74 98 Eichstaettisaurus schroederi AMPHISBAENIAN Eichstaettisaurus gouldi 100 Morunasaurus annularis 74 Hoplocercus spinosus Saichangurvel davidsoni MOSASAUR 100 100 Ctenomastax parva 78 100 Temujinia ellisoni 100 Anchaurosaurus gilmorei SNAKE Zapsosaurus sceliphros 100 Liolaemus Stenocercus 74 78 Microlophus albermalensis ANGUIMORPH 61 100 Polrussia mongoliensis Plica plica Tropidurus torquatus 100 Chalarodon madagascariensis IGUANIAN 97 Oplurus cyclurus Uta stansburiana 74 100 60 Sceloporus 97 Callisaurus draconoides DIBAMID 60 Urosaurus nigricaudus Phrynosoma asio 100 Polychrus gutturosus 63 Enyalius bilineatus BORIOTEIIOID 100 100 Gambelia Crotaphytus collaris 100 Sauromalus ater 63 Brachylophus ARDEOSAUR 63 100 Leiosaurus catamarcensis Anolis carolinensis 63 Afairiguana avius Laemanctus serratus GLOBAURID 100 100 Priscagama gobiensis 100 Phrynosomimus asper 100 Mimeosaurus crassus 100 100 Flaviagama dzerzhinskii PARAMACELLODID Gladidenagama semiplena 100 Arretosaurus ornatus 100 Rieppeleon brachyurus 100 Brookesia brygooi OTHER 100 Uromastyx aegyptia 100 Physignathus cocincinus Agama agama Figure 1—figure supplement 2. Majority rule consensus tree for the constrained analysis of lepidosaurs. Bolet et al. eLife 2022;11:e66511. DOI: https://doi.org/10.7554/eLife.66511  4 of 57 Research article Evolutionary Biology Pamelina polonica Icarosaurus siefkeri Kuehneosaurus latus Marmoretta oxoniensis Scandensia ciervensis Planocephalosaurus robinsonae Diphydontosaurus avonis Gephyrosaurus bridensis Kallimodon cerinensis Priosphenodon avelasi Pamizinsaurus tlayuaensis Palaeopleurosaurus posidoniae Pleurosaurus goldfussi Bharatagama rebbanensis Sphenodon punctatus Cynosphenodon huizachalensis Clevosaurus bairdi Clevosaurus mcgilli Ankylosphenodon pachyostosus Clevosaurus brasiliensis Polysphenodon mulleri Schoenesmahl dyspepsia Bavarisaurus macrodactylus Huehuecuetzpalli mixtecus Ardeosaurus brevipes Unnamed gen PMUR58 Palaeolacerta bavarica Bainguis parvus Hoyalacerta sanzi Eichstaettisaurus schroederi Eichstaettisaurus gouldi Dibamus novaeguineae Anelytropsis papillosus Sineoamphisbaena hexatabularis Polrussia mongoliensis Globaura venusta Eoxanta lacertifrons Hymenosaurus clarki Norellius nyctisaurops Gobekko cretacicus Pygopus lepidopus Eublepharis macularius Aeluroscalabotes felinus Hoburogekko suchanovi Phelsuma lineata Gekko gecko Sphaerodactylus elegans Oedura tryoni Saltuarius cornutus Nephrurus levis Ornatocephalus metzleri Myrmecodaptria microphagosa Pseudosaurillus becklesi Becklesius hoffstetteri Yabeinosaurus tenuis Eolacerta robusta Paramacellodus oweni Paramacellodid UTAH Tepexisaurus tepexii Xantusia vigilis Cricosaura typica Lepidophyma flavimaculatum Palaeoxantusia kyrentos Parmeosaurus scutatus Slavoia darevskii Platysaurus intermedius Tracheloptychus petersi Chalcides ocellatus Scincus scincus Eumeces algeriensis Proscelotes eggeli Acontias plumbeus Feylinia currori Typhlacontias gracilis Plestiodon fasciatus Plestiodon anthracinus Mochlus sundevalli Tiliqua scincoides Mabuya mabouya Lygosoma bowringii Liushusaurus acanthocaudata Cryptolacerta hassiaca Lacerta viridis Dracaenosaurus croizeti Takydromus septentrionalis Rhineura floridana Bipes biporus Cadea blanoides Blanus Dracaena guianensis Tupinambis teguixin Teius teyou Aspidoscelis tigris Ameiva ameiva Bachia bicolor Dryadosaura nordestina Potamites ecpleopus Alopoglossus festae Gymnophthalmus underwoodi Colobosaura modesta Jucaraseps grandipes Portugalophis lignites Najash rionegrina Epictia goudotii Typhlops jamaicensis Typhlophis squamosus Liotyphlops albirostris Anomochilus leonardi Rhinophis sanguineus Parviraptor estesi Dinilysia patagonica Anilius scytale Cylindrophis ruffus Boa constrictor Pachyrhachis problematicus Haasiophis terrasanctus Meyasaurus diazromerali Clades Carusia intermedia Restes rugosus Exostinus serratus Xenosaurus grandis Parasaniwa wyomingensis STEM LEPIDOSAUR Dalinghosaurus longidigitus Merkurosaurus ornatus Shinisaurus crocodilurus Bahndwivici ammoskius RHYNCHOCEPHALIAN Peltosaurus granulosus Elgaria coerulea Dorsetisaurus purbeckensis Saniwa feisti STEM SQUAMATE Eosaniwa koehni Proplatynotia longirostrata Adriosaurus microbrachis Adriosaurus suessi GEKKOTAN Paravaranus angustifrons Eonatator sternbergi Aigialosaurus Parviderma inexacta SCINCID Saniwides mongoliensis Telmasaurus grangeri Lanthanotus borneensis CORDYLIFORM Saniwa ensidens Varanus varius Psammosaurus griseus Gobiderma pulchrum XANTUSIID Primaderma nessovi Paraderma bogerti Palaeosaniwa canadensis Heloderma horridum TEIOID Estesia mongoliensis Chometokadmon fitzingeri Tchingisaurus multivagus Gobinatus arenosus LACERTID Cherminsaurus kozlowskii Adamisaurus magnidentatus Erdenetesaurus robinsonae Polyglyphanodon sternbergi AMPHISBAENIAN Gilmoreteius ferrugenous Tianyusaurus zhengi Gilmoreteius chulsanensis Darchansaurus estesi MOSASAUR Morunasaurus annularis Hoplocercus spinosus Uta stansburiana Sceloporus SNAKE Phrynosoma asio Urosaurus nigricaudus Callisaurus draconoides Chalarodon madagascariensis ANGUIMORPH Oplurus cyclurus Saichangurvel davidsoni Ctenomastax parva Temujinia ellisoni IGUANIAN Anchaurosaurus gilmorei Zapsosaurus sceliphros Liolaemus DIBAMID Stenocercus Microlophus albermalensis Plica plica Tropidurus torquatus BORIOTEIIOID Gambelia Crotaphytus collaris Sauromalus ater Brachylophus ARDEOSAUR Polychrus gutturosus Enyalius bilineatus Leiosaurus catamarcensis Anolis carolinensis GLOBAURID Afairiguana avius Laemanctus serratus Priscagama gobiensis Phrynosomimus asper PARAMACELLODID Mimeosaurus crassus Flaviagama dzerzhinskii Gladidenagama semiplena Arretosaurus ornatus OTHER Rieppeleon brachyurus Brookesia brygooi Uromastyx aegyptia Physignathus cocincinus Agama agama Figure 1—figure supplement 3. Adams consensus tree for the constrained analysis of lepidosaurs. Bolet et al. eLife 2022;11:e66511. DOI: https://doi.org/10.7554/eLife.66511  5 of 57 Research article Evolutionary Biology Pamelina polonica Icarosaurus siefkeri Kuehneosaurus latus Marmoretta oxoniensis Scandensia ciervensis Hoyalacerta sanzi Liushusaurus acanthocaudata Sineoamphisbaena hexatabularis Cryptolacerta hassiaca Jucaraseps grandipes Tchingisaurus multivagus Gobinatus arenosus Microlophus albermalensis Polrussia mongoliensis Leiosaurus catamarcensis Eichstaettisaurus schroederi Eichstaettisaurus gouldi Chometokadmon fitzingeri Ardeosaurus brevipes Schoenesmahl dyspepsia Unnamed gen PMUR58 Meyasaurus diazromerali Carusia intermedia Dorsetisaurus purbeckensis Becklesius hoffstetteri Yabeinosaurus tenuis Ornatocephalus metzleri Eolacerta robusta Chalcides ocellatus Scincus scincus Eumeces algeriensis Mochlus sundevalli Tiliqua scincoides Mabuya mabouya Lygosoma bowringii Proscelotes eggeli Acontias plumbeus Palaeolacerta bavarica Globaura venusta Eoxanta lacertifrons Bainguis parvus Parmeosaurus scutatus Hymenosaurus clarki Myrmecodaptria microphagosa Slavoia darevskii Pseudosaurillus becklesi Tepexisaurus tepexii Bavarisaurus macrodactylus Huehuecuetzpalli mixtecus Dibamus novaeguineae Anelytropsis papillosus Morunasaurus annularis Hoplocercus spinosus Liolaemus Stenocercus Plica plica Tropidurus torquatus Chalarodon madagascariensis Oplurus cyclurus Polychrus gutturosus Enyalius bilineatus Anolis carolinensis Afairiguana avius Peltosaurus granulosus Elgaria coerulea Paramacellodus oweni Paramacellodid UTAH Plestiodon fasciatus Plestiodon anthracinus Feylinia currori Typhlacontias gracilis Platysaurus intermedius Tracheloptychus petersi Lacerta viridis Dracaenosaurus croizeti Takydromus septentrionalis Restes rugosus Exostinus serratus Xenosaurus grandis Rhineura floridana Bipes biporus Cadea blanoides Blanus Gambelia Crotaphytus collaris Sauromalus ater Brachylophus Xantusia vigilis Cricosaura typica Lepidophyma flavimaculatum Palaeoxantusia kyrentos Uta stansburiana Sceloporus Urosaurus nigricaudus Phrynosoma asio Callisaurus draconoides Saichangurvel davidsoni Ctenomastax parva Temujinia ellisoni Anchaurosaurus gilmorei Zapsosaurus sceliphros Parasaniwa wyomingensis Dalinghosaurus longidigitus Merkurosaurus ornatus Shinisaurus crocodilurus Bahndwivici ammoskius Cherminsaurus kozlowskii Adamisaurus magnidentatus Erdenetesaurus robinsonae Polyglyphanodon sternbergi Gilmoreteius ferrugenous Gilmoreteius chulsanensis Darchansaurus estesi Tianyusaurus zhengi Dracaena guianensis Tupinambis teguixin Teius teyou Aspidoscelis tigris Ameiva ameiva Bachia bicolor Clades Dryadosaura nordestina Potamites ecpleopus Alopoglossus festae Gymnophthalmus underwoodi STEM LEPIDOSAUR Colobosaura modesta Laemanctus serratus Priscagama gobiensis Phrynosomimus asper RHYNCHOCEPHALIAN Mimeosaurus crassus Flaviagama dzerzhinskii Gladidenagama semiplena Arretosaurus ornatus STEM SQUAMATE Rieppeleon brachyurus Brookesia brygooi Uromastyx aegyptia Physignathus cocincinus GEKKOTAN Agama agama Norellius nyctisaurops Gobekko cretacicus Pygopus lepidopus SCINCID Hoburogekko suchanovi Sphaerodactylus elegans Phelsuma lineata Gekko gecko CORDYLIFORM Eublepharis macularius Aeluroscalabotes felinus Oedura tryoni XANTUSIID Saltuarius cornutus Nephrurus levis Portugalophis lignites Parviraptor estesi TEIOID Najash rionegrina Dinilysia patagonica Anomochilus leonardi Rhinophis sanguineus LACERTID Anilius scytale Cylindrophis ruffus Boa constrictor Pachyrhachis problematicus AMPHISBAENIAN Haasiophis terrasanctus Epictia goudotii Typhlops jamaicensis Typhlophis squamosus MOSASAUR Liotyphlops albirostris Planocephalosaurus robinsonae Diphydontosaurus avonis Gephyrosaurus bridensis SNAKE Kallimodon cerinensis Priosphenodon avelasi Pamizinsaurus tlayuaensis Palaeopleurosaurus posidoniae ANGUIMORPH Pleurosaurus goldfussi Bharatagama rebbanensis Sphenodon punctatus Cynosphenodon huizachalensis IGUANIAN Clevosaurus mcgilli Clevosaurus bairdi Ankylosphenodon pachyostosus Clevosaurus brasiliensis DIBAMID Polysphenodon mulleri Saniwa feisti Parviderma inexacta BORIOTEIIOID Lanthanotus borneensis Telmasaurus grangeri Saniwides mongoliensis Adriosaurus microbrachis ARDEOSAUR Adriosaurus suessi Proplatynotia longirostrata Eosaniwa koehni Saniwa ensidens GLOBAURID Varanus varius Psammosaurus griseus Paravaranus angustifrons Eonatator sternbergi PARAMACELLODID Aigialosaurus Gobiderma pulchrum Primaderma nessovi Paraderma bogerti OTHER Palaeosaniwa canadensis Heloderma horridum Estesia mongoliensis Figure 1—figure supplement 4. Strict consensus tree for the constrained analysis of lepidosaurs. Bolet et al. eLife 2022;11:e66511. 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