Monstersauria

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Monstersauria
Temporal range: Albian–Recent
Heloderma suspectum.jpg
Gila monster, Heloderma suspectum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Infraorder: Neoanguimorpha
Clade: Monstersauria
Norell and Gao, 1997 [2]
Subgroups

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). [3]

Classification

Traditionally, Monstersauria was thought to include the modern Helodermatidae along with fossil genera such as Gobiderma and Estesia on the finding that it was a sister to Varanidae. But in more recent years, such as 2004 and 2008, more precise molecular studies have shown that the extant Heloderma is closer to Anguidae & kin than to Varanoidea. A large-scale integrated analysis on squamate phylogeny incorporating 737 characters of morphological and molecular data in 2015 analyzed the traditionally-monstersaurian fossil taxa along with the rest of the dataset, and what it found was a well-supported separation of the extinct monstersaurians from the extant Heloderma . In total, three different possibilities exist: either Heloderma is sister to the rest of Neoanguimorpha and fossil monstersaurians nest within Varanoidea (based on molecular and combined data; optimal arrangement); they both nest with each other in Neoanguimorpha (unlikely possibility based on parsimony analysis of combined data), or they both nest with each other in Varanoidea (based on morphology only). The most likely tree chosen by the authors, based on the combined dataset of 691 morphological characters and 46 molecular characters across 210 operational taxonomic units, is as shown, focusing on Anguimorpha: [4]

Squamata

~ Huehuecuetzpalli

~ Eichstaetisaurus

Dibamidae

Gekkota

Scincomorpha

Lacertoidea

Amphisbaenia

Teiidae

Toxicofera

Mosasauria

Serpentes

Polyglyphanodontia

Iguania

Anguimorpha
Paleoanguimorpha

Shinisauridae

Gobiderma

Aiolosaurus

Estesia

Varanoidea

Lanthanotus

Varanidae

Saniwa

Varanus

sensu Estes 1998
sensu Lee 1997
Neoanguimorpha
Monstersauria

Heloderma

Diploglossa

Xenosauridae

Anguioidea

Glyptosauridae

Diploglossidae

Anniellidae

Anguidae

Related Research Articles

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

Amniotes belong to the clade Amniota, a clade of tetrapod vertebrates that comprises sauropsids and synapsids. They are distinguished from the other living tetrapod clade—the lissamphibians—by the development of three extraembryonic membranes, thicker and more keratinized skin, and costal respiration.

<span class="mw-page-title-main">Squamata</span> Order of reptiles

Squamata is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians, which are collectively known as squamates or scaled reptiles. With over 10,900 species, it is also the second-largest order of extant (living) vertebrates, after the perciform fish. Members of the order are distinguished by their skins, which bear horny scales or shields, and must periodically engage in molting. They also possess movable quadrate bones, making possible movement of the upper jaw relative to the neurocranium. This is particularly visible in snakes, which are able to open their mouths very wide to accommodate comparatively large prey. Squamates are the most variably sized living reptiles, ranging from the 16 mm (0.63 in) dwarf gecko to the 6.5 m (21 ft) Reticulated python. The now-extinct mosasaurs reached lengths over 14 m (46 ft).

<span class="mw-page-title-main">Diapsid</span> Clade of amniote tetrapods with two holes in each side of their skulls

Diapsids are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago during the late Carboniferous period. All diapsids other than the most primitive ones in the clade Araeoscelidia are sometimes placed into the clade Neodiapsida. The diapsids are extremely diverse, and include birds and all modern reptile groups, including turtles, which were historically thought to lie outside the group. Although some diapsids have lost either one hole (lizards), or both holes, or have a heavily restructured skull, they are still classified as diapsids based on their ancestry. At least 17,084 species of diapsid animals are extant: 9,159 birds, and 7,925 snakes, lizards, tuatara, turtles, and crocodiles.

<span class="mw-page-title-main">Crotaphytidae</span> Family of lizards

The Crotaphytidae, or collared lizards, are a family of desert-dwelling reptiles native to the Southwestern United States and northern Mexico. Alternatively they are recognized as a subfamily, Crotaphytinae, within the clade Pleurodonta. They are very fast-moving animals, with long limbs and tails; some species are capable of achieving bipedal running at top speed. This species is carnivorous, feeding mainly on insects and smaller lizards. The two genera contain 12 species.

<span class="mw-page-title-main">Mexican blind lizard</span> Species of lizard

The Mexican blind lizard is a species of legless lizard in the family Dibamidae, and the only species in the genus Anelytropsis. It is endemic to Mexico. They look like Amphisbaenia, but are in fact, only distantly related.

<span class="mw-page-title-main">Sauropsida</span> Taxonomic clade

Sauropsida is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early synapsids have historically been referred to as "mammal-like reptiles", all synapsids are more closely related to mammals than to any modern reptile. Sauropsids, on the other hand, include all amniotes more closely related to modern reptiles than to mammals. This includes Aves (birds), which are now recognized as a subgroup of archosaurian reptiles despite originally being named as a separate class in Linnaean taxonomy.

<span class="mw-page-title-main">Mosasaur</span> Extinct marine lizards of the Late Cretaceous

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.

<span class="mw-page-title-main">Toxicofera</span> Proposed clade of scaled reptiles

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.

<span class="mw-page-title-main">Iguanomorpha</span> Infraorder of lizards

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.

<span class="mw-page-title-main">Pythonomorpha</span> Clade of lizards

Pythonomorpha was originally proposed by paleontologist Edward Drinker Cope (1869) as a reptilian order comprising mosasaurs, which he believed to be close relatives of Ophidia (snakes). The etymology of the term Pythonomorpha comes from the Greek Python and morphe ("form"), and refers to the generally serpentine body plan of members of the group. Cope wrote, "In the mosasauroids, we almost realize the fictions of snake-like dragons and sea-serpents, in which men have been ever prone to indulge. On account of the ophidian part of their affinities, I have called this order Pythonomorpha." Cope incorporated two families, the Clidastidae and the Mosasauridae.

<span class="mw-page-title-main">Aigialosauridae</span> Extinct family of lizards

Aigialosauridae is a family of Late Cretaceous semiaquatic pythonomorph lizards closely related to the mosasaurs. Regarded by some paleontologists as a distinct monophyletic group and by others as an adaptive grade within the basal mosasauroids, recent molecular and morphological data suggests that they are the oldest known members of the lineage leading to the mosasaurs.

<span class="mw-page-title-main">Anguimorpha</span> Order of lizards

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.

<span class="mw-page-title-main">Platynota</span> Clade of lizards

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.

<i>Aigialosaurus</i> Extinct genus of lizards

Aigialosaurus is an extinct genus of Late Cretaceous marine or semiaquatic lizard classified as part of the family Aigialosauridae within the Mosasauroidea. Exclusively found in deposits of Cenomanian age near Hvar, Croatia, the genus contains one valid species, A. dalmaticus. According to recent molecular and morphological data, Aigialosaurus is the oldest known member of the lineage leading to large Cretaceous marine reptiles called mosasaurs, a group most closely related to snakes among living squamates. It was a relatively small reptile with a complete specimen measuring 65 cm (2.13 ft) long.

<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.

<span class="mw-page-title-main">Polyglyphanodontia</span> Extinct clade of lizards

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 three 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. Macrocephalosaurus, from the Gobi Desert, was a specialized herbivore; it grew to roughly a meter long and had multicusped, leaf-shaped teeth like those of modern iguanas. 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.

<span class="mw-page-title-main">Lacertoidea</span> Superfamily of lizards

The Lacertoidea is a group of squamate reptiles that includes the Lacertidae, Teiidae, Gymnophthalmidae, and Amphisbaenia. The finding from molecular phylogenetic studies that the burrowing Amphisbaenia were nested in a clade with the lizard forms led Vidal & Hedges (2005) to propose a new name for the group based on shared morphogical characters, Laterata, "referring to the presence of tile-like scales that form the rings in Amphisbaenia, and are also present ventrally in Lacertiformata and Teiformata".

<span class="mw-page-title-main">Neoanguimorpha</span> Clade of lizards

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.

<span class="mw-page-title-main">Paleoanguimorpha</span> Clade of lizards

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.

<span class="mw-page-title-main">Diploglossa</span> Clade of lizards

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.

References

  1. Tadahiro Ikeda, Hidetoshi Ota, Tomonori Tanaka, Kenji Ikuno, Katsuhiro Kubota, Kohei Tanaka and Haruo Saegusa. 2022. A Fossil Monstersauria (Squamata: Anguimorpha) from the Lower Cretaceous Ohyamashimo Formation of the Sasayama Group in Tamba City, Hyogo Prefecture, Japan. Cretaceous Research. 130, 105063. DOI: 10.1016/j.cretres.2021.105063
  2. M. A. Norell and K. Gao. (1997). Braincase and phylogenetic relationships of Estesia mongoliensis from the Late Cretaceous of the Gobi Desert and the recognition of a new clade of lizards. American Museum Novitates 3211:1-25
  3. Tod W. Reeder; Ted M. Townsend; Daniel G. Mulcahy; Brice P. Noonan; Perry L. Wood, Jr.; Jack W. Sites, Jr.; John J. Wiens (2015). "Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa". PLOS ONE. 10 (3): e0118199. Bibcode:2015PLoSO..1018199R. doi: 10.1371/journal.pone.0118199 . PMC   4372529 . PMID   25803280.
  4. Tod W. Reeder; Ted M. Townsend; Daniel G. Mulcahy; Brice P. Noonan; Perry L. Wood, Jr.; Jack W. Sites, Jr.; John J. Wiens (2015). "Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa://S8 Fig Estimated phylogeny of squamates based on likelihood analysis of the combined morphological and molecular data, including all taxa (-lnL = 979677.56)". PLOS ONE. 10 (3): e0118199. Bibcode:2015PLoSO..1018199R. doi: 10.1371/journal.pone.0118199 . PMC   4372529 . PMID   25803280.