The Varanidae are a family of lizards in the superfamily Varanoidea and order Anguimorpha. The family, a group of carnivorous and frugivorous lizards, includes the living genus Varanus and a number of extinct genera more closely related to Varanus than to the earless monitor lizard (Lanthanotus). 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 earless monitor lizard and chinese crocodile lizard. The oldest members of the family are known from the Late Cretaceous of Mongolia.
Iguania is an infraorder of squamate reptiles that includes iguanas, chameleons, agamids, and New World lizards like anoles and phrynosomatids. Using morphological features as a guide to evolutionary relationships, the Iguania are believed to form the sister group to the remainder of the Squamata, which comprise nearly 11,000 named species, roughly 2000 of which are iguanians. However, molecular information has placed Iguania well within the Squamata as sister taxa to the Anguimorpha and closely related to snakes. The order has been under debate and revisions after being classified by Charles Lewis Camp in 1923 due to difficulties finding adequate synapomorphic morphological characteristics. Most Iguanias are arboreal but there are several terrestrial groups. They usually have primitive fleshy, non-prehensile tongues, although the tongue is highly modified in chameleons. The group has a fossil record that extends back to the Early Jurassic. Today they are scattered occurring in Madagascar, the Fiji and Friendly Islands and Western Hemisphere.
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. longirostrata, was named in 1984.
Polyglyphanodontia, also known as the Borioteiioidea, is an extinct clade of lizards from the Cretaceous that includes around a dozen genera. Polyglyphanodontians were the dominant group of lizards in North America and Asia during the Late Cretaceous. Most polyglyphanodontians are Late Cretaceous in age, though the oldest one, Kuwajimalla kagaensis, is known from the Early Cretaceous Kuwajima Formation (Japan). Early Cretaceous South American taxon Tijubina, and possibly also Olindalacerta, might also fall within Polyglyphanodontia or be closely allied to the group, but if so, they would be two of only four Gondwanan examples of an otherwise Laurasian clade. They produced a remarkable range of forms. Chamopsiids, including Chamops, were characterized by large, blunt, crushing teeth, and were most likely omnivores. Polyglyphanodon, from the Maastrichtian of Utah, was another herbivore, but its teeth formed a series of transverse blades, similar to those of Trilophosaurus. Peneteius had remarkable, multicusped teeth, similar to those of mammals. The polyglyphanodontids first appear in the latter part of the Early Cretaceous in North America, and became extinct during the Cretaceous-Paleogene extinction event. Polyglyphanodontians closely resembled the teiid lizards, and purported teiid lizards from the Late Cretaceous appear to be polyglyphanodontians. The only species known to have survived the Cretaceous was Chamops, which survived until the very early Ypresian.
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:
Chamaeleontiformes is a hypothesized clade of iguanian lizards defined as all taxa sharing a more recent common ancestor with Chamaeleo chamaeleon than with Hoplocercus spinosus, Polychrus marmoratus, or Iguana iguana. It was named by paleontologist Jack Conrad in 2008 to describe a clade recovered in his phylogenetic analysis that included the extinct genus Isodontosaurus, the extinct family Priscagamidae, and the living clade Acrodonta, which includes agamids and chameleons. It is a stem-based taxon and one of two major clades within Iguania, the other being Pleurodonta. Below is a cladogram from Daza et al. (2012) showing this phylogeny:
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.
Heterodontagama is an extinct genus of iguanian lizard from the Early Eocene of India. It belongs to the extinct family Priscagamidae, which is otherwise only known from the Late Cretaceous of Mongolia. The type species Heterodontagama borsukae was named in 2013 from several isolated upper and lower jaws found in an exposure of the Cambay Shale in an open-pit coal mine in Gujarat.
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.
Pleurodontagama is an extinct genus of iguanian lizard from the Late Cretaceous of Mongolia. The type species, Pleurodontagama aenigmatodes, was named in 1984 on the basis of a mostly complete skull and isolated lower jaw from a fossil locality called Khermeen Tsav. It has a wide skull with a flat snout, large eye sockets, and small bumps on the surfaces of the bones. Pleurodontagama was initially classified in the family Priscagamidae, which is usually grouped in a large clade of iguanians called Acrodonta, members of which are characterized by an "acrodont" dentition in which the teeth grow from the margins of the jaws. However, Pleurodontagama is unusual in that it has a sub-pleurodont dentition, meaning that some of its teeth grow from the inner surfaces of the jaw. There is also evidence to suggest that its teeth may have been continuously replaced throughout life, as opposed to the permanent teeth of acrodontans. Pleurodontagama may have been a transitional form between the derived acrodont type and the pleurodont type inferred for the ancestors of Acrodonta.
Mimeosaurus is an extinct genus of iguanian lizard from the Late Cretaceous of Mongolia. It is part of an extinct family of iguanians called Priscagamidae, and was the first priscagamid to have been described, having been named by American paleontologist Charles W. Gilmore in 1943. Currently only one species, the type species Mimeosaurus crassus, belongs within the genus. A second species, M. tugrikinensis, was named in 1989, but later studies argued that the specimens on which the new name is based are not sufficiently different from M. crassus specimens to warrant being classified as a separate species. Mimeosaurus is unique among iguanians in having premaxilla bones at the tip of the snout that are reduced in size, as well as having two pairs of enlarged canine-like teeth in the maxilla.
Priscagama is an extinct genus of iguanian lizard from the Late Cretaceous of Mongolia and China. It belongs to an extinct family of iguanians called Priscagamidae. Several incomplete skulls have been found in the Barun Goyot and Djadochta formations, and were originally referred to the genus Mimeosaurus; the type species Priscagama gobiensis was named in 1984 when it was recognized that these skulls belonged to a distinct species. Priscagama differs from most other priscagamids in having a more elongate, lightly built skull. It is very similar in appearance to another priscagamid called Pleurodontagama, as the two can only be distinguished by the shape of their teeth.
Phrynosomimus is an extinct genus of iguanian lizard from the Late Cretaceous of Mongolia belonging to the extinct family Priscagamidae. The type species Phrynosomimus asper was named in 1996. Fossils have been found in the Barun Goyot and Djadochta formations and include several complete skulls. Phrynosomimus has a short, triangular skull with bony spikes projecting from the back, stemming from the squamosal and parietal bones. These spikes give it a similar appearance to the modern horned lizard Phrynosoma and inspire its name, which means "Phrynosoma mimic." Like other priscagamids it has an acrodont dentition, meaning that its teeth grow from the margins of the jaws rather than their inner surfaces, as is the case for the pleurodont dentitions of most lizards.
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:
Xihaina is an extinct genus of iguanian lizard from the Late Cretaceous of Inner Mongolia, China. The type species Xihaina aquilonia was named in 1995 from the Djadochta Formation and is known from a partial skeleton that preserves parts of the skull, most of the vertebral column, the pelvis, and the right hind limb. The incomplete nature of this specimen makes the classification of Xihaina difficult; it has never been incorporated into a phylogenetic analysis, but it shares similarities with a group of Late Cretaceous Mongolian lizards called Gobiguania, particularly the gobiguanian genera Anchaurosaurus and Polrussia. The fact that many skeletal elements are missing yet the rest of the skeleton is articulated suggests the individual may have been partially eaten by a predator or scavenger and then rapidly buried soon after.
Polrussia is an extinct genus of iguanian lizards dating to the Late Cretaceous epoch, found in what is now Mongolia. It belongs to a group of extinct iguanians called Gobiguania that was endemic to the Gobi Desert during the Late Cretaceous. The type species Polrussia mongoliensis was named in 1991 on the basis of a skull found in the Barun Goyot Formation. The genus name refers to the Polish and Russian paleontologists who worked together to find and describe the material. Polrussia has a short skull, slightly pointed and flattened snout, and large eye sockets. The teeth each have one cusp, as opposed to the multiple cusps seen in some other gobiguanians. The skull is only 1.2 centimetres (0.47 in) long, making Polrussia one of the smallest gobiguanians.
Igua is an extinct genus of iguanian lizards belonging to a group called Gobiguania that was endemic to the Gobi Desert during the Late Cretaceous. The type species Igua minuta was named in 1991 on the basis of a skull from the Barun Goyot Formation in Mongolia. The skull itself is very small, only 14 millimetres (0.55 in) long, and may have belonged to a juvenile given that it possesses a large fontanelle and that many of the bones are unfused. The snout-vent length of the individual is estimated to have been 55 to 65 millimetres. Igua differs from related gobiguanians like Polrussia in having a more rounded skull. It is similar in appearance to the living genera Liolaemus and Tropidurus. The teeth are tricuspid and pleurodont, meaning they are attached to inner surfaces of the jaws.
Acrodonta are a subclade of iguanian squamates consisting almost entirely of Old World taxa. Extant representation include the families Chamaeleonidae (chameleons) and Agamidae, with at least over 500 species described. A fossil genus, Gueragama, was found in Brazil, making it the only known American representative of the group.
Magnuviator is a genus of extinct iguanomorph lizard from the Late Cretaceous of Montana, US. It contains one species, M. ovimonsensis, described in 2017 by DeMar et al. from two specimens that were discovered in the Egg Mountain nesting site. Magnuviator is closest related to the Asian Saichangurvel and Temujinia, which form the group Temujiniidae. Unlike other members of the Iguanomorpha, however, Magnuviator bears a distinct articulating notch on its tibia for the ankle bones, which has traditionally been considered a characteristic of non-iguanomorph lizards. The morphology of its teeth suggests that its diet would have mainly consisted of wasps, like the modern phyrnosomatid iguanians Callisaurus and Urosaurus, although it also shows some adaptations to herbivory.
