Carusioidea Temporal range: Late Cretaceous - present | |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Squamata |
Infraorder: | Neoanguimorpha |
Clade: | Carusioidea Gao and Norell, 1998 |
Subgroups | |
Carusioidea is a clade (evolutionary grouping) of lizards that includes the family Xenosauridae (knob-scaled lizards) from Central America and the extinct genus Carusia from the Late Cretaceous of Mongolia. It was named in 1998 after a sister-group relationship was found between Carusia and Xenosauridae. Phylogenetic analysis indicates that Carusioidea is the most basal clade within Anguimorpha. Features that help define Carusioidea include closely spaced orbits or eye sockets separated by fused frontal bones, a connection between the jugal and squamosal bones below the supratemporal arch, and a covering of bony osteoderms over the skull roof. Below is a cladogram showing the phylogenetic relationships of carusioids from Gao and Norell (1998): [1]
Squamata (lizards) |
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Xenosauridae is a family of anguimorph lizards whose only living representative is the genus Xenosaurus, which is native to Central America. Xenosauridae also includes the extinct genera Exostinus and Restes. Also known as knob-scaled lizards, they have rounded, bumpy scales and osteoderms. Most living species prefer humid, rocky habitats, although they are widespread within their native regions, with some inhabiting semi-arid scrub environments. They are carnivorous or insectivorous, and give birth to live young.
The Varanidae are a family of lizards in the superfamily Varanoidea within the Anguimorpha group. The family, a group of carnivorous and frugivorous lizards, includes the living genus Varanus and a number of extinct genera. Varanus includes the Komodo dragon, crocodile monitor, savannah monitor, the goannas of Australia and Southeast Asia, and various other species with a similarly distinctive appearance. Their closest living relatives are the anguid and helodermatid lizards.
Heloderma is a genus of toxicoferid lizards that contains five species, all of which are venomous. It is the only extant genus of the family Helodermatidae.
Goronyosaurus is an extinct genus of marine lizard belonging to the mosasaur family. Fossils of Goronyosaurus are exclusively known from the Dukamaje Formation of Niger and Nigeria and also the Rima Formation of Nigeria, and are both Maastrichtian in age. The holotype was first described in 1930 as Mosasaurus nigeriensis, but subsequent remains revealed a highly unique set of adaptations that prompted the species to be reclassified as the only species of the new genus Goronyosaurus in 1972. These unique adaptations have made Goronyosaurus notoriously difficult to classify within the Mosasauridae and it is often left out of phylogenetic analyses, although most authors agree that Goronyosaurus belonged to Mosasauridae.
Estesia is an extinct genus of Late Cretaceous helodermatoid lizard found in the Gobi Desert in Mongolia. It was discovered in June 1990 by a joint expedition made up of Mongolian and American palaeontologists, and described in 1992 by Mark Norell, Malcolm McKenna and Michael Novacek. This animal is of interest to palaeontologists, not only because it is close to the lineage of modern Gila monsters (Heloderma), but also because its dentition shows evidence that it was venomous.
Platynota is a group of anguimorph lizards and thus belongs to the order Squamata of the class Reptilia. Since it was named in 1839, it has included several groups, including monitor lizards, snakes, mosasaurs, and helodermatids. Its taxonomic use still varies, as it is sometimes considered equivalent to the group Varanoidea and other times viewed as a distinct group. It is phylogenetically defined as a clade containing Varanidae. It also includes many extinct species.
Ovoo gurvel is an extinct monitor lizard from the Late Cretaceous of Mongolia. It is one of the smallest and earliest monitor lizards. It was described in 2008. Ovoo possesses a pair of two small bones in its skull that are not seen in any other lizard.
Aiolosaurus is an extinct genus of monitor lizard from the Late Cretaceous of Mongolia. The type and only species, Aiolosaurus oriens, was named in 2000 from Ukhaa Tolgod, a rich fossil site in the Campanian-age Djadochta Formation.
Proplatynotia is an extinct genus of varanoid lizard from the Late Cretaceous of Mongolia. Fossils have been found in the Barun Goyot Formation, which is mid-Campanian in age. The type and only species, P. longirostratia, was named in 1984.
Gekkonomorpha is a clade of lizards that includes geckos and their closest relatives. Although it was first named in 1900, Gekkonomorpha was not widely used as a formal taxon until it was given a phylogenetic definition in the 1990s. Under this definition, Gekkonomorpha is a stem-based taxon containing the node-based taxon Gekkota, the group that includes the last common ancestor of all living geckos and its descendants. The extent of Gekkonomorpha beyond gekkotans differs between studies. For example, Lee (1998) defined Gekkonomorpha in such a way that it included not only Gekkota but the legless amphisbaenian and dibamid lizards as well. The phylogenetic analysis of Conrad (2008), which did not support a close relationship between geckos and legless lizards, used Gekkonomorpha in a much more restrictive sense so that it included only Gekkota and a few extinct lizards more closely related to Gekkota than to any other living group of lizards. Some of the most recent phylogenetic analyses suggest that the extinct lizards Gobekko and Parviraptor may be stem gekkotans, although other analyses find that Gobekko may instead be within Gekkota and Parviraptor outside Gekkonomorpha.
Glyptosaurinae is an extinct subfamily of anguid lizards that lived in the Northern Hemisphere during the Late Cretaceous and the Paleogene.
Shinisauria is a clade or evolutionary grouping of anguimorph lizards that includes the living Chinese crocodile lizard Shinisaurus and several of its closest extinct relatives. Shinisauria was named in 2008 as a stem-based taxon to include all anguimorphs more closely related to Shinisaurus than to any other lizard. Several recent phylogenetic analyses of lizard evolutionary relationships place Shinisauria in a basal position within the clade Platynota, which also includes monitor lizards, helodermatids, and the extinct mosasaurs. Shinisaurians were once thought to be closely related to the genus Xenosaurus, but they are now considered distant relatives within Anguimorpha. The fossil record of shinisaurians extends back to the Early Cretaceous with Dalinghosaurus, which is from the Aptian aged Yixian Formation of China. Two other extinct shinisaurians are currently known: Bahndwivici from the Eocene of Wyoming and Merkurosaurus from the Late Oligocene of Germany and the Early Miocene of the Czech Republic. An indeterminate shinisaurian is known from an isolated tail found in the Eocene aged Messel pit in Germany.
Gobiguania is an extinct clade of iguanian lizards from the Late Cretaceous. All known gobiguanians are endemic to the Gobi Desert of Mongolia. Gobiguania was given a phylogenetic definition by Jack Conrad and Mark Norell in 2007 as all taxa more closely related to Anchaurosaurus gilmorei than to Iguana iguana, Crotaphytus collaris, or Agama agama. According to Conrad and Norell's phylogenetic analysis, Gobiguania includes Anchaurosaurus as well as several other Late Cretaceous lizards such as Ctenomastax, Temujinia, Saichangurvel, and Zapsosaurus. A phylogenetic analysis published in 2012 indicated that three other lizard genera — Igua, Isodontosaurus, and Polrussia, all from Mongolia and all from the Late Cretaceous — are also gobiguanians. Below is a cladogram from the analysis:
Myrmecodaptria is an extinct genus of scleroglossan lizard from the Late Cretaceous Djadokhta Formation in Ukhaa Tolgod, Mongolia. The type and only species, Myrmecodaptria microphagosa, was named in 2000 by paleontologists Gao Keqin and Mark Norell. Myrmecodaptria is known from a single holotype skull and lower jaws. It is distinguished from all other lizards by its extremely elongated skull. The eyes are placed close to the snout, which is short and rounded. The top of the skull is covered in bony knobs called osteoderms. The parietal bone at the back of the skull is elongated and about as long as the frontal bones, which are the usually the longest bones along the top of the skull in lizards. The squamosal bone at the back of the skull reaches forward to connect with the jugal bone behind the eye, forming a thin arch between the temporal fenestrae. Myrmecodaptria also has fewer and more widely spaced teeth in its jaws than do most other lizards.
Carusia is an extinct genus of lizards from the Late Cretaceous of Mongolia. It is a close relative of the family Xenosauridae, which includes living knob-scaled lizards. Fossils of the type and only species Carusia intermedia come from the late-Campanian age Barun Goyot Formation and have been found in the Flaming Cliffs, Ukhaa Tolgod, and Kheerman Tsav fossil localities. Carusia was first described in 1985 under the name Carolina intermedia, but since the name Carolina was preoccupied by a genus of scarab beetles that had been named in 1880, it was renamed Carusia intermedia. Carusia had initially been known from fragmentary skull material, complicating efforts to determine its evolutionary relationships with other lizards; it had variously been described as an indeterminate scincomorph, a xenosaurid, or some other type of autarchoglossan lizard convergent with xenosaurids. However, the discovery of 35 complete skulls in the 1990s, three of which were described in a detailed 1998 monograph, revealed that Carusia was the sister taxon of Xenosauridae, compelling the authors of the monograph to create a new clade called Carusioidea to include both taxa.
Saichangurvel is an extinct genus of iguanian lizards from the Late Cretaceous of Mongolia. It is a member of a clade called Gobiguania, an exclusively Late Cretaceous group of iguanian lizards that was likely endemic to the Gobi Desert. The type species, Saichangurvel davidsoni, was named by paleontologists Jack Conrad and Mark Norell of the American Museum of Natural History in 2007. It is known from a single nearly complete and fully articulated skeleton called IGM 3/858, which was found eroding from a block of sandstone during a thunderstorm at a fossil locality called Ukhaa Tolgod. IGM 3/858 comes from the Djadochta Formation, which is between 75 and 71 million years in age. Just as it is today, the Gobi was a desert during the Cretaceous. IGM 3/858 may have died in a collapsing sand dune, the rapid burial preserving its skeleton in pristine condition.
Anchaurosaurus is an extinct genus of iguanian lizard from the Late Cretaceous of Inner Mongolia, China. It belongs to an extinct clade of iguanians called Gobiguania that was endemic to the Gobi Desert during the Late Cretaceous. The type species, Anchaurosaurus gilmorei, was named in 1995 on the basis of a well-preserved skull and incomplete skeleton from the Djadochta Formation. Compared to other iguanians, Anchaurosaurus has a relatively elongated skull, large eye sockets, and higher tooth crowns. Phylogenetic analysis indicates that among gobiguanians, Anchaurosaurus is most closely related to Zapsosaurus from Mongolia. Below is a cladogram from Daza et al. (2012) showing the phylogenetic relationships of Anchaurosaurus:
Neoanguimorpha is a clade of anguimorphs comprising Monstersauria and Diploglossa. Morphological studies in the past had classified helodermatids with the varanoids in the clade Platynota, while the Chinese crocodile lizard was classified as a xenosaurid. However molecular work found no support in these groupings and instead has found the helodermatids more related to Diploglossa, while the Chinese crocodile lizard and varanoids to form the clade Paleoanguimorpha.
Paleoanguimorpha is a clade of anguimorphs comprising Shinisauria and Goannasauria. Morphological studies in the past also classified helodermatids and pythonomorphs with the varanoids in the clade Platynota, while the Chinese crocodile lizard was classified as a xenosaurid. Current molecular work finds no support in these groupings and instead has found the helodermatids more related to Diploglossa in the sister clade Neoanguimorpha, while the Chinese crocodile lizard is the closet living relative to varanoids. Pythonomorphs represented by snakes today are not closely related to varanoids and are instead a sister lineage to Anguimorpha and Iguania in the clade Toxicofera.
Diploglossa is a clade of neoanguimorphs represented by the families Xenosauridae, Diploglossidae, Anniellidae and Anguidae, the latter three placed in the superfamily Anguioidea. In the past the Chinese crocodile lizard was classified as a xenosaurid; current molecular work has shown evidence the species related to varanoids in the clade Paleoanguimorpha.