Anguidae

Slowworm Anguis fragilis, copyright Hans Hillewaert.

Belongs within: Anguimorpha.

The Anguidae, glass lizards and allies, are a group of lizards found in Eurasia, northern Africa and the Americas that are characterised by rectangular scales, osteoderms in the ventral skin, a reduced supratemporal arch and striations on the medial face of the tooth crowns (Animal Diversity Web).

<==Anguidae
    |--Diploglossus [Diploglosseae, Diploglossinae] H81
    |    |--D. bilobatus MH11
    |    |--D. occiduus (Shaw 1802) H81
    |    `--D. pleei MH11
    |--Glyptosaurinae SH96
    |    |--Helodermoides Douglass 1903 [Glyptosaurini] SH96
    |    |    `--H. tuberculatus Douglass 1903 [incl. Glyptosaurus giganteus Gilmore 1928, G. montanus Douglass 1908] SH96
    |    `--Melanosaurini SH96
    |         |--Melanosaurus LBG12
    |         `--Peltosaurus Cope 1873 SH96
    |              `--*P. granulosus Cope 1873 (see below for synonymy) SH96
    |--Ophisaurus Daudin 1803 [incl. Dopasia Gray 1853; Ophisaureae, Ophisaurinae] RB05
    |    |--O. attenuatus JP79
    |    |--O. canadensis SH96
    |    |--O. compressus DO99
    |    |--O. formosensis Kishida 1930 TYM08
    |    |--O. harti [=Dopasia harti] SH96
    |    |--O. roqueprunensis Augé 1992 SH96
    |    `--O. ventralis AM99
    `--Anguinae SH96
         |--Parophisaurus Sullivan 1987 SH96
         |    `--*P. pawneensis (Gilmore 1928) (see below for synonymy) SH96
         `--Anguis Linnaeus 1758 [Anguieae] L58
              |--A. bipes Linnaeus 1758 L58
              |--A. cephallonica [incl. A. fragilis peloponnesiacus] N10
              |--A. cerastes Linnaeus 1758 L58
              |--A. colchica [=A. fragilis colchica, A. fragilis colchicus] N10
              |--A. colubrina Linnaeus 1758 L58
              |--A. eryx Linnaeus 1758 L58
              |--A. fragilis Linnaeus 1758 BB03
              |--A. graeca N10
              |--A. jaculus Linnaeus 1758 L58
              |--A. laticauda Linnaeus 1758 L58
              |--A. lumbricalis Linnaeus 1758 L58
              |--A. maculata Linnaeus 1758 L58
              |--A. meleagris Linnaeus 1758 L58
              |--A. reticulata Linnaeus 1758 L58
              `--A. scytale Linnaeus 1758 L58

Anguidae incertae sedis:
  Odaxosaurus piger Gilmore 1928 B93 [=Pancelosaurus piger G88]
  Proxestops LBG12
  Parodaxosaurus LBG12
  Paraglyptosaurus LBG12
  Apodosauriscus LBG12
  Eodiploglossus LBG12
  Gaultia LBG12
  Gerrhonotus [Gerrhonoteae] LBG12
    |--G. liocephalus MH11
    `--G. multicarinatus GWN71
  Abronia DD61
    |--A. deppi DD61
    `--A. graminea MH11
  Barisia DD61
    |--B. gadovi DD61
    `--B. imbricata MH11
  Celestus LBG12
    |--C. enneagrammus MH11
    |--C. occiduus FS01
    `--C. sagrae F15
  Pseudopus Merrem 1820 RB05
    |--P. apodus (Pallas 1775) RB05 [=Ophisaurus apodus B89]
    |--P. laurillardi (Lartet 1851) [=Anguis laurillardi, Ophisaurus laurillardi] RB05
    |--P. moguntinus (Boettger 1875) RB05 (see below for synonymy)
    `--P. pannonicus (Kormos 1911) RB05 [=Ophisaurus pannonicus B89]
  Elgaria AS09
    |--E. coerulea MH11
    |--E. kingii MH11
    |--E. multicarinata MH11
    |--E. panamintina MH11
    `--E. paucicarinata MH11
  Ophisauriscus quadrupes SH96
  Placosaurus SH96

*Parophisaurus pawneensis (Gilmore 1928) [=Xestops pawneensis, Machaerosaurus pawneensis, Pancelosaurus pawneensis] SH96

*Peltosaurus granulosus Cope 1873 [incl. P. abbotti Gilmore 1928, P. abbottii (l. c.), P. floridanus Vanzolini 1952] SH96

Pseudopus moguntinus (Boettger 1875) RB05 [=Ophisaurus moguntinus B89; incl. Propseudopus fraasi Hilgendorf 1883 RB05]

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

[AM99] Anderson, J. F., & L. A. Magnarelli. 1999. Enzootiology of Borrelia burgdorferi in the northeastern and northcentral United States. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 385–389. Ohio Biological Survey: Columbus (Ohio).

[B89] Bailon, S. 1989. Les amphibiens et les reptiles du Pliocene superieur de Balaruc II (Herault, France). Palaeovertebrata 19 (1): 7–28.

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[BB03] Blain, H.-A., & S. Bailon. 2003. Les amphibiens et les reptiles des couches du Pléistocène supérieur ancien du gisement d’Artenac (Charente, France). Quaternaire 14 (1): 85–95.

[DD61] Davis, W. B., & J. R. Dixon. 1961. Reptiles (exclusive of snakes) of the Chilpancingo Region, Mexico. Proceedings of the Biological Society of Washington 74: 37–56.

[DO99] Durden, L. A., & J. H. Oliver Jr. 1999. Ecology of Ixodes scapularis and Lyme disease in coastal Georgia, USA. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 379–383. Ohio Biological Survey: Columbus (Ohio).

[FS01] Flannery, T., & P. Schouten. 2001. A Gap in Nature: Discovering the World’s Extinct Animals. Text Publishing: Melbourne.

[F15] Fowler, H. W. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 244–269.

[GWN71] Gorman, G. C., A. C. Wilson & M. Nakanishi. 1971. A biochemical approach towards the study of reptilian phylogeny: evolution of serum albumin and lactic dehydrogenase. Systematic Zoology 20 (2): 167–185.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H81] Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141–158.

[JP79] Jackson, J. F., & J. A. Pounds. 1979. Comments on assessing the dedifferentiating effect of gene flow. Systematic Zoology 28 (1): 78–85.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[LBG12] Longrich, N. R., B.-A. S. Bhullar & J. A. Gauthier. 2012. Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 109 (52): 21396–21401.

[MH11] Müller, J., C. A. Hipsley, J. J. Head, N. Kardjilov, A. Hilger, M. Wuttke & R. R. Reisz. 2011. Eocene lizard from Germany reveals amphisbaenian origins. Nature 473: 364–367.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

[SH96] Sullivan, R. M., & J. A. Holman. 1996. Squamata. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 354–372. Cambridge University Press.

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

Colubridae

Collared reed snake Calamaria pavimentata, copyright Rushenb.

Belongs within: Caenophidia.
Contains: Thamnophis, Lampropeltis, Pseudorabdion, Natricinae, Xenodontinae, Colubrinae.

The Colubridae are a cosmopolitan group of snakes, including both non-venomous and rear-fanged venomous species, characterised by the presence of solid teeth on the maxilla, palatine, pterygoid and dentary, but never on the premaxilla.

Colubridae
    |--+--Natricinae VH02
    |  `--+--Pseudoxenodon karlschmidti VH02
    |     `--Xenodontinae VH02
    `--+--Grayia ornata VH02
       `--+--Colubrinae VH02
          `--Calamariinae VH02
               |--Oreocalamus hanitschi VH02
               `--Calamaria VH02
                    |--C. babia S60
                    |--C. favae de Filippi 1840 (n. d.) [=Atractus favae] S60
                    |--C. pavimentata VH02
                    |--C. pfefferi Stejneger 1901 I92
                    `--C. stahlknechti M89

Colubridae incertae sedis:
  Thamnophis VH02
  Dendrophidion dendrophis DS86
  Drymarchon corais DS86
  Drymobius rhombifer DS86
  Drymoluber dichrous DS86
  Leptophis DS86
    |--L. abaetulla DS86
    |    |--L. a. abaetulla DS86
    |    `--L. a. nigromarginatus DS86
    |--L. cupreus DS86
    `--L. mexicanus M86
  Pseustes DS86
    |--P. poecilonotus DS86
    |    |--P. p. poecilonotus DS86
    |    `--P. p. polylepis DS86
    |--P. sexcarinatus DS86
    `--P. sulphureus DS86
  Spilotes pullatus DS86
  Xenopholis scalaris DS86
  Psomophis C07
  Coronella austriaca Laurenti 1768 BB03 [=Coluber austriacus K08]
  Texasophis Holman 1977 RB05
    |--*T. fossilis Holman 1977 SH96
    |--T. bohemiacus RB05
    |--T. galbreathi Holman 1984 SH96
    `--T. meini Rage & Holman 1984 RB05
  Dromicus H81
    |--D. cursor (Lacépède 1789) [=Leimadophis cursor] H81
    `--D. ornatus Garman 1887 [=Leimadophis ornatus] H81
  Phalotris JK08
    |--P. cuyanus (Cei 1984) JK08
    |--P. lemniscatus (Duméril, Bibron & Duméril 1854) JK08
    |--P. matogrossensis Lema, D’Agostini & Cappellari 2005 JK08
    |--P. mertensi (Hoge 1955) JK08
    |--P. nasutus (Gomes 1915) JK08
    |--P. sansebastiani Jansen & Köhler 2008 JK08
    `--P. tricolor (Duméril, Bibron & Duméril 1854) [incl. P. punctatus (Lema 1979)] JK08
  Elapomorphus JK08
    |--E. mertensi Hoge 1955 H55
    `--E. tricolor H55
  Dendrophis R13
    |--D. calligaster R13
    |--D. caudolineata M89
    `--D. picta M89
  Trimorphodon biscutatus N10
  Pituophis GWN71
    |--P. catenifer OB11
    `--P. melanoleucus GWN71
  Apostolepis G11
  Lytorhynchus diadema [incl. L. kennedyi Schmidt 1939] A-O00
  Dendrelaphis Boulenger 1890 C18
    |--D. calligastra (Günther 1867) C18
    |--D. caudolineatus VH02
    |--D. modestus B28
    |--D. pictus VH02
    `--D. punctulatus (Gray 1826) C18
  Lampropeltis G88
  Aspidura trachyprocta CD88
  Afronatrix anoscopus B06
  Stegonotus Duméril, Bibron & Duméril 1854 C18
    |--S. cucullatus (Duméril, Bibron & Duméril 1854) C18
    `--S. parvus (Meyer 1874) C18
  Tropidonophis Jan 1863 C18
    `--T. mairii (Gray 1841) C18
         |--T. m. mairii C18
         `--T. m. brongersmai C18
  Pantherophis guttata C18
  Zamenis SBM01
    |--Z. andreana SBM01
    `--Z. dipsas B28
  Ninia sebae M86
  Pseudorabdion IL61

*Type species of generic name indicated

REFERENCES

[A-O00] Al-Oran, R. M. 2000. Notable herpetological records from central and southern Jordan. Zoology in the Middle East 21: 31–36.

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[BB03] Blain, H.-A., & S. Bailon. 2003. Les amphibiens et les reptiles des couches du Pléistocène supérieur ancien du gisement d'Artenac (Charente, France). Quaternaire 14 (1): 85–95.

[B06] Boruchowitz, D. E. 2006. Guide to Cichlids. T.F.H. Publications: Neptune City (New Jersey).

[C07] Cadle, J. E. 2007. The snake genus Sibynomorphus (Colubridae: Dipsadinae: Dipsadini) in Peru and Ecuador, with comments on the systematics of Dipsadini. Bulletin of the Museum of Comparative Zoology 158 (5): 183–284.

[C18] Cogger, H. G. 2018. Reptiles and Amphibians of Australia updated 7^th ed. CSIRO Publishing: Collingwood.

[CD88] Crusz, H., & D. M. A. K. Daundasekera. 1988. Parasites of the relict fauna of Ceylon. IX. Helminths from burrowing scincine lizards. Annales de Parasitologie Humaine et Comparee 63: 439–447.

[DS86] Dixon, J. R., & P. Soini. 1986. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum: Milwaukee.

[GWN71] Gorman, G. C., A. C. Wilson & M. Nakanishi. 1971. A biochemical approach towards the study of reptilian phylogeny: evolution of serum albumin and lactic dehydrogenase. Systematic Zoology 20 (2): 167–185.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[G11] Guadanucci, J. P. L. 2011. Cladistic analysis and biogeography of the genus Oligoxystre Vellard 1924 (Araneae: Mygalomorphae: Theraphosidae). Journal of Arachnology 39 (2): 320–326.

[H55] Hoge, A. R. 1955. Eine neue Schlange der Gattung Elapomorphus aus Brasilien. Senckenbergiana Biologica 36 (5-6): 301–304.

[H81] Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141–158.

[IL61] Inger, R. F., & A. E. Leviton. 1961. A new colubrid snake the genus Pseudorabdion from Sumatra. Fieldiana: Zoology 44 (5): 45–47.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[JK08] Jansen, M., & G. Köhler. 2008. A new species of Phalotris from the eastern lowlands of Bolivia (Reptilia, Squamata, Colubridae). Senckenbergiana Biologica 88 (1): 103–110.

[K08] Korsós, Z. 2008. History of the herpetological collection of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 37–93.

[M86] McCoy, C. J. 1986. Results of the Carnegie Museum of Natural History expeditions to Belize. I. Systematic status and geographic distribution of Sibon neilli (Reptilia, Serpentes). Annals of Carnegie Museum 55 (6): 117–123.

[M89] Modigliani, E. 1889. Materiali per la fauna erpetologica dell'isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 113–124.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[OB11] Orenstein, R. I., & D. Brewer. 2011. Family Cardinalidae (cardinals). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 330–427. Lynx Edicions: Barcelona.

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

[R13] Roux, J. 1913. Les reptiles de la Nouvelle-Calédonie et des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 2 pp. 79–160. C. W. Kreidels Verlag: Wiesbaden.

[S60] Savage, J. M. 1960. A revision of the Ecuadorian snakes of the colubrid genus Atractus. Miscellaneous Publications Museum of Zoology, University of Michigan 112: 1–86.

[SBM01] Schätti, B., I. Baran & P. Maunoir. 2001. Taxonomie, Morphologie und Verbreitung der Masken-Schlanknatter Coluber (s. l.) collaris (Müller, 1878). Revue Suisse de Zoologie 108 (1): 11–30.

[SH96] Sullivan, R. M., & J. A. Holman. 1996. Squamata. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 354–372. Cambridge University Press.

[VH02] Vidal, N., & S. B. Hedges. 2002. Higher-level relationships of caenophidian snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus Biologies 325: 987–995.

Natricinae

Barred grass snake Natrix natrix helvetica, copyright Benny Trapp.

Belongs within: Colubridae.

The Natricinae, watersnakes, are a group of snakes characterised by the presence of hypapophyses on all dorsal vertebrae, and minutely spinose hemipenes bearing one or more enlarged basal hooks. Members of the subfamily commonly hunt around water, feeding on small aquatic vertebrates.

<==Natricinae
    |  i. s.: Palaeonatrix Szyndlar 1982 RB05
    |           |--P. lehmani (Rage & Roček 1983) [=Dolniceophis lehmani] RB05
    |           `--P. silesiaca Syndlar 1982 RB05
    |         Neonatrix Holman 1973 RB05
    |           |--N. europaea Rage & Holman 1984 RB05
    |           |--N. natricoides Augé & Rage 2000 RB05
    |           `--N. nova RB05
    |         Mionatrix G88
    |--+--Natriciteres olivacea VH02
    |  `--+--Rhabdophis subminiatus VH02
    |     `--Xenochrophis VH02
    |          |--X. flavipunctatus VH02
    |          `--X. trianguligerus VH02
    `--+--Sinonatrix annularis VH02
       |--Nerodia VH02
       |    |--N. harteri JP79
       |    |--N. rhombifera VK00
       |    |--N. sipedon BK03 [=Natrix sipedon FM80]
       |    |    |--N. s. sipedon BK03
       |    |    `--N. insularum BK03
       |    `--N. taxispilota VH02
       `--Natrix Laurenti 1768 VH02, RB05
            |--N. chrysarga D56
            |--N. fasciatus F15
            |--N. longivertebrata Szyndlar 1984 RB05
            |--N. merkurensis Ivanov 2002 RB05
            |--N. mlynarskii Rage 1988 B93
            |--N. natrix (Linnaeus 1758) VK00 [=Coluber natrix K08, Tropidonotus natrix M01]
            |    |--N. n. natrix M55a
            |    |--N. n. cetti M55a
            |    `--N. n. helvetica M55a
            |--N. piscator D56
            |--N. sansaniensis (Lartet 1851) [=Coluber sansaniensis, Pilemophis sansaniensis, Pylmophis sansaniensis] RB05
            |--N. septemvittata R56
            |--N. stolata CS77
            `--N. tesselata M55b [=N. tessellata (l. c.) M55b, Tropidonotus tessellatus K08]

*Type species of generic name indicated

REFERENCES

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[BK03] Bittner, T. D., & R. B. King. 2003. Gene flow and melanism in garter snakes revisited: a comparison of molecular markers and island vs. coalescent models. Biological Journal of the Linnean Society 79: 389–399.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F15] Fowler, H. W. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 244–269.

[FM80] Futuyma, D. J., & G. C. Mayer. 1980. Non-allopatric speciation in animals. Systematic Zoology 29 (3): 254–271.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[JP79] Jackson, J. F. & J. A. Pounds. 1979. Comments on assessing the dedifferentiating effect of gene flow. Systematic Zoology 28 (1): 78–85.

[K08] Korsós, Z. 2008. History of the herpetological collection of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 37–93.

[M01] Méhely, L. 1901. Csúszómászók és kétéltűek [Reptilien und Amphibien]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 41–68. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[M55a] Mertens, R. 1955a. Die Amphibien und Reptilien der Insel Elba. Senckenbergiana Biologica 36: 287–296.

[M55b] Mertens, R. 1955b. Der Typus von Vipera lebetina schweizeri. Senckenbergiana Biologica 36: 297–299.

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

[R56] Reed, C. F. 1956. The herpetofauna of Harford County, Maryland. Journal of the Washington Academy of Sciences 46 (2): 58–60.

[VH02] Vidal, N., & S. B. Hedges. 2002. Higher-level relationships of caenophidian snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus Biologies 325: 987–995.

[VK00] Vidal, N., S. G. Kindl, A. Wong & S. B. Hedges. 2000. Phylogenetic relationships of xenodontine snakes inferred from 12S and 16S ribosomal RNA sequences. Molecular Phylogenetics and Evolution 14: 389–402.

Last updated: 29 January 2022.

Anguimorpha

Reconstruction of Coniophis, copyright N. Tamura.

Belongs within: Lepidosauria.
Contains: Anguidae, Amphisbaenia, Serpentes, Varanidae.

The Anguimorpha are a clade of lizards including the glass lizards, monitors, snakes and related taxa. As shown below, phylogenetic analysis of the squamates by Longrich et al. (2012b) also included the Amphisbaenia within this clade but this is almost certainly an artefact of the analysis drawing legless taxa together. Supporting characters of the Anguimorpha include the ability to retract the foretongue. Members include the beaded lizards Heloderma, strikingly coloured venomous lizards from southern North America.

Synapomorphies (from Rieppel 1988): Foretongue retractible within hindtongue at a transverse zone of invagination; inferior labial glands differentiated; type 2 fundic glands present; genital tubercle lateral or ventral to sinus genitalis.

<==Anguimorpha [Anguinomorpha, Anguoidea, Varanoidea]
    |  i. s.: Carolinidae B93
    |           |--Carusia intermedia (Borsuk-Bialynicka 1985) B93
    |           `--Shinisauroides latipalatum Borsuk-Bialynicka 1985 B93
    |         Dorsetisaurus [Dorsetisauridae] B93
    |           |--D. hebetidens Hoffstetter 1967 B93
    |           `--D. purbeckensis Hoffstetter 1967 [incl. Introrsisaurus pollicidens Seiffert 1973] B93
    |--Xenosauridae LBG12b
    |    |--Xenosaurus LBG12b
    |    |    |--X. grandis LBG12b
    |    |    `--X. platyceps LBG12b
    |    |--Restes LBG12b
    |    `--Exostinus Cope 1873 SH96
    |         |--*E. serratus Cope 1873 SH96
    |         `--E. lancensis Gilmore 1928 B93
    `--+--Anguidae LBG12b
       `--+--+--Sineoamphisbaena Wu, Brinkman et al. 1993 LBG12b, D07
          |  |    `--S. hexatubularis D07
          |  `--+--Anniella [Anniellidae] LBG12b
          |     |    `--A. pulchra LBG12b
          |     `--+--+--Cryptolacerta Müller, Hipsley et al. 2011 MH11
          |        |  |    `--*C. hassiaca Müller, Hipsley et al. 2011 MH11
          |        |  `--Amphisbaenia LBG12b
          |        `--Ophidia LBG12b
          |             |  i. s.: Rottophis atavus (von Meyer 1860) S05
          |             |         Cheilophis LBG12b
          |             |         Sanjuanophis LBG12b
          |             |--+--Serpentes LBG12a
          |             |  `--Najash Apesteguía & Zaher 2006 LBG12a, AZ06
          |             |       `--*N. rionegrina Apesteguía & Zaher 2006 AZ06
          |             `--Coniophis Marsh 1892 [Coniophidae] LBG12a
          |                  |--C. carinatus McGrew et al. 1959 LBG12a
          |                  |--C. cosgriffi Armstrong-Ziegler 1978 LBG12a
          |                  |--C. dabiebus Rage & Werner 1999 LBG12a
          |                  |--C. platycarinatus McGrew et al. 1959 LBG12a
          |                  `--C. precedens Marsh 1892 LBG12a
          `--+--Shinisaurus LBG12b [Shinisauridae MH11]
             |    `--S. crocodilurus LBG12b
             `--Platynota LBG12b
                  |  i. s.: Labrodioctes LBG12b
                  |         Provaranosaurus acutus LBG12b
                  |         Primaderma Nydam 2000 AS09, D07
                  |           `--P. nessovi D07
                  |--Litakis gilmorei Estes 1964 LBG12b, B93
                  |--Colpodontosaurus cracens LBG12b
                  `--+--Cemeterius Longrich, Bhullar & Gauthier 2012 LBG12b
                     |    `--*C. monstrosus Longrich, Bhullar & Gauthier 2012 LBG12b
                     `--+--Palaeosaniwa canadensis LBG12b
                        |--Paraderma bogerti Estes 1964 LBG12b, B93
                        |--Necrosauridae G88
                        |    |--Parasaniwa wyomingensis LBG12b
                        |    |--Parviderma inexacta Borsuk-Bialynicka 1984 B93
                        |    `--Necrosaurus eucarinatus (Kuhn 1940) B93
                        |--Gobiderma pulchrum LBG12b
                        `--+--Estesia LBG12b
                           `--+--Aiolosaurus oriens LBG12b
                              `--+--Helodermatidae MH11
                                 |    |--Eurheloderma SH96
                                 |    |--Lowesaurus Pregill, Gauthier & Greene 1986 SH96
                                 |    |    `--*L. matthewi (Gilmore 1928) [=Heloderma matthewi] SH96
                                 |    `--Heloderma LBG12b
                                 |         |--H. horridum LBG12a
                                 |         `--H. suspectum MH11
                                 `--+--+--Saniwa Leidy 1870 LBG12b, SH96
                                    |  |    |--*S. ensidens Leidy 1870 SH96
                                    |  |    `--S. feisti MH11
                                    |  `--Varanidae LBG12b
                                    `--Lanthanotus [Lanthanotidae] LBG12b
                                         `--L. borneensis LBG12a

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

[AZ06] Apesteguía, S. & H. Zaher. 2006. A Cretaceous terrestrial snake with robust hindlimbs and a sacrum. Nature 440: 1037–1040.

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[LBG12a] Longrich, N. R., B.-A. S. Bhullar & J. A. Gauthier. 2012. A transitional snake from the Late Cretaceous period of North America. Nature 488: 205–208.

[LBG12b] Longrich, N. R., B.-A. S. Bhullar & J. A. Gauthier. 2012b. Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 109 (52): 21396–21401.

[MH11] Müller, J., C. A. Hipsley, J. J. Head, N. Kardjilov, A. Hilger, M. Wuttke & R. R. Reisz. 2011. Eocene lizard from Germany reveals amphisbaenian origins. Nature 473: 364–367.

[S05] Scanlon, J. D. 2005. Cranial morphology of the Plio-Pleistocene giant madtsoiid snake Wonambi naracoortensis. Acta Palaeontologica Polonica 50: 139–180.

[SH96] Sullivan, R. M., & J. A. Holman. 1996. Squamata. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 354–372. Cambridge University Press.

Last updated: 30 January 2022.

Serpentes

Western blind snake Leptotyphlops humilis, copyright Ashley Wahlberg.

Belongs within: Anguimorpha.
Contains: Palaeophiidae, Typhlopidae, Madtsoiidae, Tropidophiidae, Anilioidea, Macrostomata.

The Serpentes are the crown group of the snakes, a distinctive group of legless squamates (though remnant spurs are present in some species). Within the Serpentes, there is general agreement that the blind snakes or worm snakes of the Scolecophidia represent the sister group to the remainder (the Alethinophidia). The Scolecophidia have a limited gape and feed mostly on ants and termites whereas the Alethinophidia possess independent mandibles allowing a much larger gape (Vidal & Hedges 2002a). Scolecophidia are divided between three families. Members of Typhlopidae and Anomalopedidae share a transverse, movable, toothed maxilla whereas in Leptotyphlops the maxilla is immobile, longitudunal and toothless. Anomalopedidae differ from the other two scolecophidian families in possessing teeth in the lower jaw.

The Anilioidea within the Alethinophidia are fossorial like the Scolecophidia and such habits have been suggested to be basal within the snakes. However, support for this premise may be affected by the positions of extinct non-fossorial groups such as the Madtsoiidae and Similiophiidae. The latter were marine snakes from the Late Cretaceous, at least some species of which retained distinct hind limbs.

Synapomorphies (from Rieppel 1988): Intracranial course of ophthalmic division of trigeminal nerve; exoccipitals meeting dorsal to foramen magnum; opening of Jacobson's organ enclosed between septomaxilla and vomer; crista circumfenestralis enclosing juxtastapedial recess and perilymphatic sinus; epiphyses absent.

<==Serpentes [Booidea]
    |--Scolecophidia LBG12a
    |    |--+--Typhlopidae HB09
    |    |  `--Anomalopedidae HB09
    |    |       |--Liotyphlops albirostris HB09, LBG12a
    |    |       `--Helminthophis petersi HB09
    |    `--Leptotyphlops LBG12a [Leptotyphlopidae HB09]
    |         |--L. carlae HB09
    |         |--L. columbi VH02a
    |         |--L. dulcis LBG12a
    |         |--L. goudotii SP-H86
    |         |    |--L. g. goudotii SP-H86
    |         |    `--L. g. phenops (Cope 1876) [=Stenostoma phenops] SP-H86
    |         |--L. humilis HB09
    |         `--L. macrorhynus LBG12a
    `--+--Dinilysia [Dinilysiidae] LBG12a
       |    `--D. patagonica Woodward 1901 S05
       `--+--Madtsoiidae LBG12a
          `--Alethinophidia LBG12a
               |  i. s.: Cerberophis Longrich, Bhullar & Gauthier 2012 LBG12b
               |           `--*C. robustus Longrich, Bhullar & Gauthier 2012 LBG12b
               |         Coprophis Parris & Holman 1978 SH96
               |           `--C. dakotaensis Parris & Holman 1978 SH96
               |--Tropidophiidae HB09
               |--Anilioidea LBG12a
               `--+--Macrostomata LBG12a
                  `--Simoliopheoidea B93
                       |--Lapparantophiidae B93
                       |    |--Lapparantophis defrennei Hoffstetter 1960 B93
                       |    `--Pouitella Rage 1988 B93, RB03
                       |         `--P. pervetus Rage 1988 B93
                       `--Similiophiidae LBG12a
                            |  i. s.: Simoliophis rochebrunei Sauvage 1880 B93
                            |--Haasiophis terrasanctus Tchernov et al. 2000 LBG12a, S05
                            `--+--Eupodophis Rage & Escuillié 2000 LBG12a, D07
                               |    `--E. descouensi (Rage & Escuillié 2000) S05
                               `--Pachyrhachis Haas 1979 LGB12a, D07
                                    `--P. problematicus Haas 1979 S05

Serpentes incertae sedis:
  Macropisthodon rudis VH02b
  Oligodon VH02b
    |--O. octolineatus VH02b
    `--O. trilineatus M89
  Storeria VH02b
    |--S. dekayi BK03
    `--S. occipitomaculata VH02b
  Opisthotropis kikuzatoi (Okada & Takara 1958) I92
  Calliophis I92
    |--C. japonicus Günther 1868 I92
    `--C. macclellandii I92
         |--C. m. macclellandii I92
         `--C. m. iwasakii Maki 1935 I92
  Opheodrys I92
    |--O. aestivus F15
    |--O. herminae (Boettger 1895) I92
    `--O. vernalis W-CL06
  Cemophora lineri RM05
  Homalocranium C07
    |--H. marcapatae [=Oxyrhopus marcapatae] C07
    `--H. virgatum F15
  Anoplodipsas viridis R13
  Dipsadoboa unicolor R13
  Tropidocloneum lineatum B75
    |--T. l. lineatum F68
    `--T. l. annectens F68
  Diemenia textilis H98
  Haldea valeriae R56
  Palaeophiidae B93
  Ahaetulla nasutus S72
  Dendrelapsis ahaetulla S72

*Type species of generic name indicated

REFERENCES

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[BK03] Bittner, T. D., & R. B. King. 2003. Gene flow and melanism in garter snakes revisited: a comparison of molecular markers and island vs. coalescent models. Biological Journal of the Linnean Society 79: 389–399.

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[C07] Cadle, J. E. 2007. The snake genus Sibynomorphus (Colubridae: Dipsadinae: Dipsadini) in Peru and Ecuador, with comments on the systematics of Dipsadini. Bulletin of the Museum of Comparative Zoology 158 (5): 183–284.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[F68] Fisher, D. R. 1968. A study of faunal resemblance using numerical taxonomy and factor analysis. Systematic Zoology 18 (4): 48–63.

[F15] Fowler, H. W. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 244–269.

[H98] Halliday, R. B. 1998. Mites of Australia: A checklist and bibliography. CSIRO Publishing: Collingwood.

[HB09] Head, J. J., J. I. Bloch, A. K. Hastings, J. R. Bourque, E. A. Cadena, F. A. Herrera, P. D. Polly & C. A. Jaramillo. 2009. Giant boid snake from the Palaeocene neotropics reveals hotter past equatorial temperatures. Nature 457: 715–717.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[LBG12a] Longrich, N. R., B.-A. S. Bhullar & J. A. Gauthier. 2012a. A transitional snake from the Late Cretaceous period of North America. Nature 488: 205–208.

[LBG12b] Longrich, N. R., B.-A. S. Bhullar & J. A. Gauthier. 2012b. Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 109 (52): 21396–21401.

[M89] Modigliani, E. 1889. Materiali per la fauna erpetologica dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 113–124.

[RB03] Rage, J.-C., S. Bajpai, J. G. M. Thewissen & B. N. Tiwari. 2003. Early Eocene snakes from Kutch, western India, with a review of the Palaeophiidae. Geodiversitas 25 (4): 695–716.

[R56] Reed, C. F. 1956. The herpetofauna of Harford County, Maryland. Journal of the Washington Academy of Sciences 46 (2): 58–60.

[RM05] Reilly, S. M., R. W. Manning, C. C. Nice & M. R. J. Forstner. 2005. Systematics of isolated populations of short-tailed shrews (Soricidae: Blarina) in Texas. Journal of Mammalogy 86 (5): 887–894.

[R13] Roux, J. 1913. Les reptiles de la Nouvelle-Calédonie et des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 2 pp. 79-160. C. W. Kreidels Verlag: Wiesbaden.

[S05] Scanlon, J. D. 2005. Cranial morphology of the Plio-Pleistocene giant madtsoiid snake Wonambi naracoortensis. Acta Palaeontologica Polonica 50: 139–180.

[SP-H86] Smith, H. M., & G. Pérez-Higareda. 1986. Nomenclatural name-forms. Systematic Zoology 35 (3): 421–422.

[S72] Soderberg, P. S. 1972. Cannibalism and feeding in two snakes, Trimeresurus albolabris and Ahaetulla nasutus. Journal of the Bombay Natural History Society 69 (1): 192–193.

[SH96] Sullivan, R. M., & J. A. Holman. 1996. Squamata. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 354–372. Cambridge University Press.

[VH02a] Vidal, N., & S. B. Hedges. 2002a. Higher-level relationships of snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus Biologies 325: 977–985.

[VH02b] Vidal, N., & S. B. Hedges. 2002b. Higher-level relationships of caenophidian snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus Biologies 325: 987–995.

[W-CL06] Watkins-Colwell, G. J., T. A. Leenders, B. T. Roach, D. J. Drew, G. Dancho & J. Yuckienuz. 2006. New distribution records for amphibians and reptiles in Connecticut, with notes on the status of an introduced species. Bulletin of the Peabody Museum of Natural History 47 (1–2): 47–62.

Last updated: 17 June 2022.