Varanoidea | |
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Lace monitor (Varanus varius) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Squamata |
Infraorder: | Paleoanguimorpha |
Clade: | Goannasauria |
Superfamily: | Varanoidea Münster, 1834 |
Families | |
Varanoidea is a superfamily of lizards, including the well-known family Varanidae (the monitors and goannas). Also included in the Varanoidea are the Lanthanotidae (earless monitor lizards), and the extinct Palaeovaranidae.
Throughout their long evolutionary history, varanoids have exhibited great diversity, both in habitat and form. This superfamily includes the largest-known terrestrial lizard, Megalania (~5 meters), and the largest extant lizard, the Komodo dragon (Varanus komodoensis, ~3 meters).
Either synonymous with, or a subgroup of, the group Platynota, the varanoids first appear in the fossil record in the latter part of Early Cretaceous, but possible varanoid ancestors have been traced back as far as Early Jurassic times. Among the earliest known varanoids are the monitor-like necrosaurids Palaeosaniwa canadensis from the Campanian (roughly 71-82 mya) of North America and Estesia mongoliensis and Telmasaurus grangeri , both from the Campanian of Mongolia. Varanoids survived the Cretaceous–Paleogene extinction event and flourished worldwide during the Cenozoic Era. Carroll argued that the affinities of ancestral snakes suggested evolution from early aquatic or burrowing varanoid lineages, although recent evidence suggests a more distant common ancestor within Toxicofera. [1] [2] [3]
Carroll characterises the varanoids as "the most advanced of all lizards in achieving large size and an active, predaceous way of life". Some taxa, such as the extinct necrosaurids and the possibly varanoid Gila monsters, were armoured with osteoderms (bony deposits on the skin), and many forms have hinged jaws, allowing them to open their mouths very wide when feeding (though they cannot dislocate their jaws, contrary to popular belief). [1] [4]
According to Estes et al., 1988, which uses morphological characteristics, Varanoidea includes Helodermatidae, Lanthanotus , and Varanus . [5] Gauthier et al., 2012 also groups these three groups together, where Helodermatidae evolved earlier than Varanidae (which includes Lanthanotus and Varanus). [6]
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.
The earless monitor lizard is a semiaquatic, brown lizard native to the Southeast Asian island of Borneo. It is the only living species in the family Lanthanotidae and it is related to the true monitor lizards.
Monitor lizards are lizards in the genus Varanus, the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and one species is also found in the Americas as an invasive species. About 80 species are recognized.
Heloderma is a genus of toxicoferan lizards that contains five species, all of which are venomous. It is the only extant genus of the family Helodermatidae.
Megalania is an extinct species of giant monitor lizard, part of the megafaunal assemblage that inhabited Australia during the Pleistocene. It is the largest terrestrial lizard known to have existed, reaching an estimated length of 3.5 to 7 metres, and weighing between 97–1,940 kg (214–4,277 lb), but the fragmentary nature of known remains make estimates highly uncertain.
Mosasaurs comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764. They belong to the order Squamata, which includes lizards and snakes.
Toxicofera is a proposed clade of scaled reptiles (squamates) that includes the Serpentes (snakes), Anguimorpha and Iguania. Toxicofera contains about 4,600 species, of extant Squamata. It encompasses all venomous reptile species, as well as numerous related non-venomous species. There is little morphological evidence to support this grouping; however, it has been recovered by all molecular analyses as of 2012.
Palaeosaniwa canadensis is an extinct species of carnivorous lizard from the late Cretaceous of North America. The name, given by Charles Whitney Gilmore in 1928, means "ancient Saniwa from Canada".
Estesia is an extinct genus of Late Cretaceous anguimorph lizard found in the Gobi Desert in Mongolia.
The Anguimorpha is a suborder of squamates. The group was named by Fürbringer in 1900 to include all autarchoglossans closer to Varanus and Anguis than Scincus. These lizards, along with iguanians and snakes, constitute the proposed "venom clade" Toxicofera of all venomous reptiles.
Platynota is a polyphyletic 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.
Monstersauria is a clade of anguimorph lizards, defined as all taxa more closely related to Heloderma than Varanus. It includes Heloderma, as well as several extinct genera, such as Estesia, Primaderma and Gobiderma, but this group was found to be polyphyletic in the most recent and complete squamate phylogenetic analysis by Reeder et al. (2015).
Ovoo gurvel is an extinct varanid 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.
Telmasaurus is an extinct genus of varanoid lizard from the Late Cretaceous of Mongolia. Fossils have been found from the Djadokha and Barun Goyot Formations that date between the early and middle Campanian stage from approximately 80 to 75 million years ago. The type species Telmasaurus grangeri was named in 1943.
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.
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.
Varanus darevskii, also known as the Central Asian monitor, is an extinct species of monitor lizard from the Early Pliocene of Tajikistan, known from a partial skull. V. darevskii might have been a close relative of, and perhaps ancestral to, the modern desert monitor.