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 Cuvier, 1817 [non? Cope, 1864] (iguaanimaiset liskot; Iguania-like lizards)
            |  |-- Pleurodonta 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 Cope, 1869 sensu Caldwell, 1999 (snakes & sealizards; käärmeet & meriliskot) [for more common position see Platynota]
               |  |==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 or Lacertidae 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 Fürbringer, 1900 [all autarchoglossans closer to Varanus & Anguis than Scincus]
                     |  |-- 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 Camp, 1923 (Scincus-like lizards; skinkkimäiset liskot)
                        |  |==Paramacellodidae Estes, 1983 [a paraphyletic form-family?]
                        |  |-- 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 Bonaparte, 1831 [non? Gray, 1827] (sisiliskot)
                        |        `--+-- Teiioidea Estes, de Queiroz & Gauthier, 1988 (Teju-like lisards; teijunsukuiset liskot)
                        |           `-- †Boreoteiioidea Nydam, Eaton & Sankey, 2007
                        `--o Nyctisauria sensu Lee, 2000
                           |-- Amphisbaenia Gray, 1844 [Annulata] (wormlizards; matoliskot)
                           `--+-- Didamidae
                              `-- 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]

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):