Squamata (original) (raw)

After Estes, de Queiroz & Gauthier, 1988, Rieppell, 1994, Evans & Barbadillo, 1998, and Caldwell, 1999

Alternative molecular phylogeny of (Recent) Squamata after Vidal & Hedges, 2005

Alternative phylogeny of Squamata after Lee, 2000

Alternative phylogeny of Squamata after Evans & Barbadillo, 1998

Alternative phylogeny of Squamata after Estes, de Queiroz & Gauthier, 1988

**<==**o Squamata Oppel, 1811 [non Merrem, 1820] (suomumatelijat; squamates) |?- †Megachirella wachtleri Renesto & Posenato, 2003; M.Trias. SEu. |--o †Bavarisauridae sensu Evans & Barbadillo, 1998 [have been treated as basal Gekkotans] | |-- †Bavarisaurus macrodactylus Hoffstetter, 1953; U.Jur. CEu. | -- †_Palaeolacerta bavarica_ Cocude-Michel, 1961; U.Jur. CEu. --+-- †Scandensia ciervensis Evans & Barbadillo, 1998 |--o †Eichstaettisauridae Kuhn, 1958 sensu Alifanov, 2000 [phylogenetic position after Evans & Barbadillo, 1998, has been treated as basal Scincomorphs] | |--o †Eichstaettisaurus Kuhn, 1958; U.Jur. Eu. [Broilisaurus Hoffstetter, 1953] | | |-- †E. schroederi (Broili, 1938) Hoffstetter, 1964 [Ardeosaurus schroederi Broili, 1938; Broilisaurus schroederi Hoffstetter, 1953] | | -- †_E. gouldi_ Evans et al., 2004 | -- †Globaura venusta Borsuk-Bialynicka, 1988; U.Cret. EAs. --+--o †_Ardeosaurus_ von Meyer, 1860 [_Eichstattisaurus_] U.Jur. Eu. [Ardeosauridae Camp, 1923] | |-- †_A. brevipes_ von Meyer, 1860 | -- †A. digitatellus (Grier, 1914) Hoffstetter, 1964 [Homeosaurus digitatellus Grier, 1914; Eichstaettisaurus digitatellus Cocude-Michel, 1963] --o “Neosquamata” [last common ancestor of all modern squamates] |?- †_Jucaraseps grandipes_ Bolet & Evans, 2012, L.Jur. SWEu. |-- [Iguania](iguania/iguania.html) Cuvier, 1817 [_non_? Cope, 1864] (iguaanimaiset liskot; _Iguania_-like lizards) | |-- [Pleurodonta](iguania/iguania.html) Cope, 1864 _sensu_ Conrad & Norell, 2007 | -- Acrodonta Cope, 1864 sensu Alifanov, 2000, Zonneveld, Gunnell & Bartels, 2000, and Evans, Prasad & Manhas, 2002 (agamat ja kameleontit; agamas & chamaeleonids) --o Scleroglossa Estes, de Queiroz & Gauthier, 1988 _sensu_ Estes, 1988 |?-o [Pythonomorpha](pythonomorpha/pythonomorpha%5F1.html) Cope, 1869 _sensu_ Caldwell, 1999 (snakes & sealizards; käärmeet & meriliskot) [for more common position see **[Platynota](anguimorpha/platynota%5F1.html)**] | |**==** †Mosasauroidea Camp, 1923 | -- Ophidia Linnaeus, 1758 sensu Lee & Caldwell, 2000; Martill, Tischlinger & Longrich, 2015 [Serpentes Linnaeus, 1758 sensu lato] (snakes; käärmeet) --o Scleroglossa _sensu_ Rieppel, 1996 |?- †_Eolacerta robusta_ Nöth, 1940 [_Iguanosaurus haupti_ Kuhn, 1944; _Iguanosauriscus haupti_ Kuhn, 1958; _Eolacerta robusta_ Estes, 1983] [problematic, see **[Anguimorpha](anguimorpha/anguimorpha.html)** or **[Lacertidae](scincomorpha/lacertidae%5F1.html)** for alternative positions] |?-o †Carusiidae Borsuk-Bialynicka, 1987 [Carolinidae Borsuk-Bialynicka, 1985, Carusioidea Borsuk-Bialynicka, 1987 _sensu_ Gao & Norell, 1998] [are scincomorps? (Alifanov, 2000) or anguimorphs? (Evans & Wang, 2005)] | |?- †_Dalinghosaurus longidigitus_ Ji, 1998; L.Cret. EAs. | |-- †_Carusia intermedia_ Borsuk-Bialynicka, 1987 [_Carolina intermedia_ Borsuk-Bialynicka, 1985 _non_ Thompson, 1880] | -- †Shinisauroides latipalatum Borsuk-Bialynicka, 1985 --o Autarchoglossa Wagler, 1830 [last common ancestor of _Scincus_ & _Varanus_] |--o [Anguimorpha](anguimorpha/anguimorpha.html) Fürbringer, 1900 [all autarchoglossans closer to _Varanus_ & _Anguis_ than _Scincus_] | |-- [Anguoidea](anguimorpha/anguimorpha.html#Anguoidea) Gray, 1825 [Anguioidea Fitzinger 1826] (vaskitsansukuiset liskot) | -- Platynota Camp, 1923 (varaanintapaiset liskot; Varanus-like lizards) --o Scincogekkonomorpha |?- †_Yabeinosaurus tenuis_ Endo & Shikama, 1942 [incl.? _Jeholacerta formosa_ Ji & Ren, 1999] L.Cret. EAs. ?U. Jur. SAs. [paraphyletic Ardeosauridae] |--o [Scincomorpha](scincomorpha/scincomorpha%5F1.html) Camp, 1923 (_Scincus_-like lizards; skinkkimäiset liskot) | |**==** †[Paramacellodidae](scincomorpha/scincomorpha%5F1.html#Paramacellodidae) Estes, 1983 [a paraphyletic form-family?] | |-- [Scincoidea](scincomorpha/scincomorpha%5F1.html#Scincoidea) Oppel, 1811 (skink-related lizards; skinkinsukuiset liskot) | --o Lacertoidea Camp, 1923 (sisiliskonsukuiset liskot) | |-- Xantusiomorpha | --o Lacertiformes Estes, de Queiroz & Gauthier, 1988 (sisiliskomaiset liskot) | |-- [Lacertidae](scincomorpha/lacertidae%5F1.html) Bonaparte, 1831 [_non_? Gray, 1827] (sisiliskot) | --+-- Teiioidea Estes, de Queiroz & Gauthier, 1988 (Teju-like lisards; teijunsukuiset liskot) | -- †[Boreoteiioidea](scincomorpha/scincomorpha%5F1.html#Boreoteiioidea) Nydam, Eaton & Sankey, 2007 --o Nyctisauria sensu Lee, 2000 |-- Amphisbaenia Gray, 1844 [Annulata] (wormlizards; matoliskot) --+-- [Didamidae](didamidae.html) -- Gekkota Cuvier, 1817 sensu stricto [Nyctisauria] (Gecko-like lizards; gekkomaiset liskot)

Squamata incertae sedis:

†_Huehuecuetzpalli mixtecus_ Reynoso, 1998
†_Conicodontosaurus_ Gilmore, 1943
    †_C. djadocbtaensis_ Gilmore, 1943
    †_C. kanhsienensis_ Young, 1973
    †_C. qinlingensis_ Li, Xue & Liu, 2004

| After Simões et al., 2018, and **<==**o Squamata |-- †Megachirella `--+--+-- †Marmoretta_ | `-- †_Huehuecuetzpalli `--+--+-- †Eichstaettisaurus_ | `--+-- †_Gobekko | `-- Gekkota `--+-- †Paramacellodus | -- †Ardeosaurus | --o Scincomorpha | | -- Xantusiidae | `--+-- Cordylidae | `-- Scincidae `--+--o Lacertiformes | | --+-- Lacertidae | | `-- Amphisbaenia | `--+-- †Globaura_ | | --+-- †Tepexisaurus | | `-- †_Slavoia | `--+-- †Meyasaurus_ | `--+-- †_Dalinghosaurus | `--+--+-- †Gobinatus_ | | `-- †_Gilmoreteius | `-- Teiioidea `--+--o Pythonomorpha | | -- †Mosasauroidea | `--+--+-- †Najash_ | | `-- †_pachyrhachis | `--+-- †Dinilysia_ | `-- Serpentes `--+-- Anguimorpha `--o Iguania | -- Iguanidae `--+-- †Pleurodontagama `--+-- †_Priscagama `-- Acrodonta Reference(s): Simões, T. R., Caldwell, M. W., Talanda, M., Bernardi, M., Palci, A., Vernygora, O., Bernardini, F., Mancini, L. & Nydam, R. L., 2018: The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps.–Nature: Vol. 557, #7707, pp. 706-709 [**doi: 10.1038/s41586-018-0093-3**] Talanda, M., 2018: An exceptionally preserved Jurassic skink suggests lizard diversification preceded fragmentation of Pangaea.–Palaeontology: Vol. 61, #5, Art.nr. pp. (659-677) [**doi: doi:10.1111/pala.12358**] | After Pyron, Burbrink & Wiens, 2013 **<==**o Squamata |-- **Dibamidae** `--+--o Gekkota | | --+-- Diplodactylidae | | `--+-- Carphodactylidae | | `-- Pygopodidae | `--+-- Eublepharidae | `--+-- Sphaerodactylidae | `--+-- Phyllodactylidae | `-- Gekkonidae `--+--o Scincoidea | | -- Scincidae | `--+-- Xantusiidae | `--+-- Gerrhosauridae | `-- Cordylidae `--o Episquamata | --o Lacertoidea | | --o Teiioidea | | | -- Gymnophthalmidae | | `-- Teiidae | `--+-- Lacertidae | `--o Amphisbaenia | | -- Bipedidae | `--+-- Blanidae | `--+-- Cadeidae | `--+-- Trogonophiidae | `-- Amphisbaenidae `--o Toxicofera | -- Serpentes `--+--o Anguimorpha | | --+-- Shinisauridae | | `--+-- Lanthanotidae | | `-- Varanidae | `--+-- Xenosauridae | `--+-- Helodermatidae | `--+-- Anniellidae | `-- Anguidae `--o Iguania | --o Acrodonta | | -- Agamidae | `-- Chamaeleonidae `--+-- Tropiduridae `--+-- Iguanidae `--+-- Leiocephalidae `--+--+-- Crotaphytidae | `-- Phrynosomatidae `--+--+-- Polychrotidae | `-- Hoplocercidae `--+--+-- Cotytophanidae | `-- Dactyloidae `--+-- Liolaemidae `--+-- Opluridae `-- Leiosauridae Reference(s): Pyron, R., Burbrink, F. & Wiens, J., 2013: A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes.–BMC Evolutionary Biology: Vol. 13, #1, Art. nr. 93, pp. 1-53 [**doi: 10.1186/1471-2148-13-93**] | After Hutchinson, Skinner & Lee, 2012 **<==**o Squamata |-- Dibamidae `--+-- Gekkota `--+-- Scincomorpha `--+--+--+-- Gymnophthalmidae | | `-- Teiidae | `--+-- Lacertidae | `--+-- †Polyglyphanodon | `--+-- †Macrocephalosaurus_ | `--+-- †_Sineoamphisbaena | `-- Amphisbaenia `--+-- Iguania `--+-- Serpentes `--+--+-- Lanthanotidae | `-- Varanidae `--+-- Shinisauridae `--+-- Xenosauridae `--+-- Anguidae `-- Helodermatidae Reference(s): Hutchinson, M. N., Skinner, A. & Lee, M. S. Y., 2012: Tikiguania_ and the antiquity of squamate reptiles (lizards and snakes).–Biology Letters: Vol. 8, #4, pp. 665-669 [**doi: 10.1098/rsbl.2011.1216**] | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | ---------------------------------------------------------------------------------------------------- | -------------------------------------------------------------- | ------------------ | ----------------------------------------------------------------------- | ----------------------------------------------------------------- | ----------------- | ------------------------------------------------------------------------------------ | --------------------------------------------------------------------- | -------------------------------------------------------- | -------------------------------------------------------------------------------- | ------------------------- | 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---------------------------------------------------------------------------------------- | ------------ | ---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- | | Conrad, 2008 **<==**o Squamata |-- †Huehuecuetzpalli | --o Iguanomorpha | | -- †Hoyalacerta | `-- Iguania `--o Scincogekkonomorpha | -- †Ardeosaurus | -- †Bavarisaurus `--+-- †_Eichstaettisaurus `--+-- †Scandensia_ | -- Gekkonomorpha `--o Evansauria | -- †_Tepexisaurus `--+--+-- †Globaura_ | `--+-- †_Bainguis | `--+-- †Myrmecodaptria_ | `-- †_Eoxanta `--+-- †Parmeosaurus_ `--o Autarchoglossa | --o Scincomorpha | | --o Scincoidea | | | -- †_Slavoia | | `--o Scinciformes | | | -- Scincidae | | `--+-- Scelotidae | | `--o Scincophidia | | | -- Feyliniidae | | `--+-- Acontidae | | `--+-- Didamidae | | `--+-- Amphisbaenia | | `-- Serpentes | `--+--o Cordyloidea | | | -- †Ornatocephalus | | | -- †Sakurasaurus | | `--o Cordyliformes | | | -- Cordylidae | | `-- Gerrhosauridae | `--o Lacertoidea | | -- Xantusiidae | `--o Lacertiformes | | -- †Chamops | | -- Lacertidae | `--+-- Gymnophthalmidae | `--+-- Teiidae | `-- †Polyglyphanodontidae `--o Anguimorpha | -- †Pseudosaurillus sp. `--+-- †Paramacellodus_ `--+-- †_Becklesius `--+-- †Yabeinosaurus_ | --+-- †Eolacerta | `--+-- †_Meyasaurus | `-- †Pseudosaurillus becklesi_ | --+-- Anguidae | `--o Carusioidea | | -- †_Carusia | `-- Xenosauridae `--o Platynota | -- †Dorsetisaurus `--+--o Shinisauria | | -- †Dalinghosaurus_ | `-- Shinisauridae `--o Varanoidea | -- †Parasaniwa `--+-- †_Parviderma `--+-- †Colpodontosaurus_ | --o Monstersauria | `-- Helodermatidae | --o Goannasauria Varanidae | `--o Varaniformes | `-- Varanidae `--o †Mosasauriformes `--o †Mosasauria `-- †Mosasauroidea Reference(s): Conrad, J. L., 2008: Phylogeny and systematics of Squamata (Reptilia) based on morphology.–Bulletin of the American Museum of Natural History: #310, pp. 1-182 | After Vidal & Hedges, 2005 **<==**o Squamata |--o Dibamia Vidal & Hedges, 2005 | `-- Dibamidae `--o Bifurcata Vidal & Hedges, 2005 | -- Gekkota `--o Unidentata Vidal & Hedges, 2005 | --o Scinciformata Vidal & Hedges, 2005 | | -- Scincidae | `--+-- Cordylidae | `-- Xantusiidae `--o Episquamata Vidal & Hedges, 2005 | --o Laterata Vidal & Hedges, 2005 | | --o Teiformata Vidal & Hedges, 2005 | | `--o Teiioidea | | | -- Teiidae | | `-- Gymnophthalmidae | `--o Lacertibaenia Vidal & Hedges, 2005 | | --o Lacertiformata Vidal & Hedges, 2005 | | `-- Lacertidae | `--o Amphisbaenia | | -- Rhineuridae | `--+-- Bipedidae | `--+-- Trogonophidae | `-- Amphisbaenidae `--o Toxicofera Vidal & Hedges, 2005 | -- Serpentes `--+--o Iguania | | -- Leiocephalus | `-- Iguaninae `--o Anguimorpha | -- Varanidae `--+-- Helodermatidae `-- Anguidae Reference(s): Fry, B. G., Vidal, N., Norman, J. A., Vonk, F. J., Scheib, H., Ramjan, S. F. R., Kuruppu, S., Fung, K., Hedges, S. B., Richardson, M. K., Hodgson, W. C., Ignjatovic, V., Summerhayes, R. & Kochva, E., 2006: Early evolution of the venom system in lizards and snakes.–Nature: Vol. 439, pp. 584-588 Vidal, N. & Hedges, S. B., 2005: The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes.–Comptes Rendus Biologies: Vol. 328, #10-11, pp. 1000-1008 | After Lee, 2000 **<==**o Squamata |-- †Scandensia `--+--o †Ardeosauridae | `-- †Ardeosaurus `--o “Neosquamata” | -- Iguania `--o Scleroglossa sensu Estes, 1988 | ?- Pythonomorpha `--o Scleroglossa | --o Nyctisauria | | -- Xantusiidae | `--+-- Gekkota | `--o Annulata | | ?- †_Sineoamphisbaena | `--+-- Didamidae | `-- Amphisbaenia `--+-- Lacertoidea `--o Diploglossa | -- Scincidae | -- Cordylidae `-- Anguimorpha Reference(s): Lee, M. S. Y., 2000: Soft anatomy, diffuse homoplasy, and the relationships of lizards and snakes.–Zoologica Scripta: Vol. 29, #1, pp. 101-130 | | After Evans & Barbadillo, 1998 **<==**o Squamata |--o †Bavarisauridae | | -- †Bavarisaurus_ | `-- †_Palaeolacerta `--+-- †Scandensia_ | --o †Eichstaettisauridae | | -- †Eichstaettisaurus | `-- †_Globaura `--+--o †Ardeosauridae | `-- †Ardeosaurus `--o “Neosquamata” | -- Iguania `--o Scleroglossa | ?- †Eolacerta | ?-o †Carusiidae | | -- †Carusia | `-- †Shinisauroides_ `--o Autarchoglossa | --o Anguimorpha | | -- Xenosauridae | `--+-- Anguidae | `--+--+-- Helodermatidae | | `-- Varanidae | `--+-- Gekkota | `--+-- Serpentes | `--+-- Didamidae | `-- Amphisbaenia `--o Scincomorpha | ?- †_Yabeinosaurus | --+-- †Meyasaurus | `--+-- Gymnophthalmidae | `-- Teiidae `--+-- Lacertidae `--+-- Xantusiidae `--+-- †Paramacellodus_ `--+-- Scincidae `--o Cordyliformes | -- Cordylidae `-- Gerrhosauridae Reference(s): Evans, S. E. & Barbadillo, L. J., 1998: An unusual lizard (Reptilia: Squamata) from the Early Cretaceous of Las Hoyas, Spain.–Zoological Journal of the Linnean Society: Vol. 124, pp. 235-265 | After Rieppell, 1994, and Caldwell, 1999 **<==**o Squamata |--o †Bavarisauridae | | -- †_Bavarisaurus | `-- †Palaeolacerta_ `--+-- †_Scandensia | --o †Eichstaettisauridae | | -- †Eichstaettisaurus | `-- †Globaura_ `--+-- †_Ardeosaurus `--o “Neosquamata” | -- Iguania `--o Scleroglossa | ?- Pythonomorpha sensu Caldwell, 1999 `--o Scleroglossa sensu_ Rieppel, 1996 | ?- †Eolacerta | ?-o †Carusiidae | | ?- †Dalinghosaurus | | -- †Carusia | `-- †_Shinisauroides `--o Autarchoglossa | -- Anguimorpha `--o Scincogekkonomorpha | ?- †Yabeinosaurus | -- Scincomorpha `--o Nyctisauria _sensu_ Lee, 2000 | -- Amphisbaenia `--+-- Didamidae `-- Gekkota Reference(s): Rieppel, O. 1994: The Lepidosauromorpha: an overview with special emphasis on the Squamata. 23-37. _in_ Fraser, N. C. & Sues, H-D. 1994: In the Shadow of the Dinosaurs.–Cambridge University Press, New York, 1994 Caldwell, M. W., 1999: Squamate phylogeny and the relationships of snakes and mosasauroids.–Zoological Journal of the Linnean Society; Vol. 125, pp. 115-147 | After Estes, de Queiroz & Gauthier, 1988 **<==**o Squamata |--o Iguania | | -- Iguanidae | `--o Acrodonta | | -- Agamidae | `-- Chamaeleontidae `--o Scleroglossa | --o Gekkota | | -- Pygopodidae | `-- Gekkonidae | ?- Didamidae | ?- Amphisbaenia | ?- Serpentes `--o Autarchoglossa | --o Scincomorpha | | --o Scincoidea | | | -- Cordylidae | | `-- Scincidae | `--o Lacertoidea | | -- Xantusiidae | `--o Lacertiformes | | -- Lacertidae | `--o Teiioidea | | -- Gymnophthalmidae | `-- Teiidae `--o Anguimorpha | -- Anguidae | -- Xenosauridae `--o Varanoidea | -- Helodermatidae `--+-- Lanthanotus `-- Varanus Reference(s): Estes, R., de Queiroz, K. & Gauthier, J., 1988: Phylogenetic Relationships within Squamata. pp. 119–281. in Estes, R. & Pregill, G. K., (eds.) 1988: Phylogenetic Relationships of the Lizard Families.–Stanford University Press, 1988, xii-631 | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | | |

Reference(s):