Platynota

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Platynota
Temporal range: Late Cretaceous - Holocene, 94–0  Ma
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Possible Valanginian record
Varanus gouldii Head.jpg
Sand goanna (Varanus gouldii)
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Anguimorpha
Clade: Platynota
Duméril and Bibron, 1839
Subgroups

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. [1] It is phylogenetically defined as a clade containing Varanidae (the monitor lizards). It also includes many extinct species.

Contents

Description

Many skeletal features support the grouping of monitor lizards, helodermatids, and several extinct species in Platynota. All platynotans have a hinged upper jaw with widely spaced teeth, each having a large base, that erupt from behind existing teeth. The teeth are plicidentine, meaning that they have highly folded layers of dentine in their centers. Many have fused frontal and nasal bones on the top of their skulls (in other lizards, these bones are separated into pairs). Some platynotan features that are not seen in the skeleton, and therefore only known from living species, include a deeply forked tongue and a venom gland called the gland of Gabe. [1]

History and classification

Platynota was first used as a superfamily of anguimorph lizards. In 1923, Charles Lewis Camp included the groups Varanoidea and Mosasauroidea, or monitor lizards and mosasaurs (a group of large marine reptiles from the Cretaceous). A close relationship between mosasaurs and snakes, which together formed the group Pythonomorpha, gained favor in the following years. Consequently, some researchers also included snakes within Platynota.

A more restricted definition was proposed by the herpetologists Samuel Booker McDowell and Charles Mitchill Bogert in 1954. Within Platynota, they included only monitor lizards, helodermatids, and their closest extinct relatives. [2] Their use of Platynota made it essentially equivalent to Varanoidea, a taxon that included monitor lizards and helodermatids and had been in use for many years. McDowell and Bogert's usage of Platynota gained favor in the following decades, although it was substituted by the name Varanoidea in some studies. [1]

Platynota was given a phylogenetic definition in 1997. It was erected as a stem-based clade that included Heloderma (the Gila monster and Beaded lizard), Lanthanotus (the Earless monitor lizard), and Varanus (monitor lizards), and all lizards that are more closely related to them than to other anguimorphs. [3] These other anguimorphs were traditionally called Diploglossa, and include Anguidae, Anniellidae (American legless lizards), and Xenosauridae (knob-scaled lizards). A node-based definition was first given in 1998 in which Platynota included the last common ancestor of Monstersauria (helodermatids) and Varanidae (monitor lizards) and all of its descendants. Under this definition, Platynota includes the same forms as Varanoidea in its traditional sense. Varanoidea, however, has been redefined as a node-based clade including the extinct Telmasaurus and varanids. [1]

Molecular evidence from DNA and other molecules conflicts with the commonly held classification of platynotans. A 2004 molecular study of living anguimorph lizards found a close relationship between helodermatids and xenosaurids, but not between helodermatids and varanids. [4] Similarly, a 2005 study found a close relationship between Anniella pulchra (the California legless lizard) and helodermatids. [5] A cladistic study synthesizing molecular and morphological data for squamates reinforced the placement of Helodermatidae as closer to Xenosauridae. [6]

Evolution

The oldest platynotans are from the Late Cretaceous of Mongolia, living around 80 million years ago. They include forms that are similar in appearance to living monitor lizards, such as Paravaranus , Proplatynotia , Gobiderma , Cherminotus , Telmasaurus , and Saniwides . Although they lived in close association with each other, they represent a diversity of different anguimorphs. The earliest and most generalized platynotans were once called necrosaurians after the genus Palaeovaranus . [1]

Related Research Articles

Lepidosauria Superorder of reptiles

The Lepidosauria is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes snakes, lizards, and amphisbaenians. Squamata contains over 9,000 species, making it by far the most species-rich and diverse order of reptiles in the present day. Rhynchocephalia was a formerly widespread and diverse group of reptiles in the Mesozoic Era. However, it is represented by only one living species: the tuatara, a superficially lizard-like reptile native to New Zealand.

Squamata 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. 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. Squamata is the most variably sized order of reptiles, ranging from the 16 mm (0.63 in) dwarf gecko to the 5.21 m (17.1 ft) green anaconda and the now-extinct mosasaurs, which reached lengths over 14 m (46 ft).

Varanidae Family of lizards

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.

<i>Heloderma</i> Genus of reptiles

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.

Ophidia Group of squamate reptiles

Ophidia is a group of squamate reptiles including modern snakes and all reptiles more closely related to snakes than to other living groups of lizards.

Mosasaur 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.

Toxicofera 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.

Autarchoglossa Clade of lizards

Autarchoglossa is a clade of squamates that includes skinks, anguimorphs, snakes, and relatives. Autarchoglossa is supported as a monophyletic grouping by morphological features in living and extinct lizards and snakes. Some phylogenetic analyses based on molecular features such as DNA sequences in living squamates do not support Autarchoglossa.

Pythonomorpha 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.

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".

Varanoidea Superfamily of reptiles

Varanoidea is a superfamily of lizards, including the well-known family Varanidae. Also included in the Varanoidea are the Lanthanotidae, and the extinct Palaeovaranidae.

<i>Sineoamphisbaena</i> Extinct genus of reptiles

Sineoamphisbaena is an extinct genus of squamate of uncertain phylogenetic placement. Its fossils are known from the Late Cretaceous deposits in Inner Mongolia, China. Wu et al. (1993), Wu et al. (1996) and Gao (1997) proposed and argued that it was the oldest known amphisbaenian; this, however, was challenged by other authors, such as Kearney (2003) and Conrad (2008), who instead assigned Sineoamphisbaena to the group of squamates variously known as Macrocephalosauridae, Polyglyphanodontidae or Polyglyphanodontia.

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.

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. longirostratia, was named in 1984.

Polyglyphanodontia 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.

Shinisauria Clade of lizards

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.

Neoanguimorpha 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.

Paleoanguimorpha 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.

References

  1. 1 2 3 4 5 Gao, K.; Norrel, M.A. (1998). "Taxonomic revision of Carusia (Reptilia: Squamata) from the Late Cretaceous of the Gobi Desert and phylogenetic relationships of anguimorphan lizards". American Museum Novitates (3230): 1–51. hdl:2246/3367.
  2. McDowell, S.B.; Bogert, C.M. (1954). "The systematic position of Lanthanotus and the affinities of the anguinomorphan lizards". Bulletin of the American Museum of Natural History. 105: 1–141.
  3. Lee, M.S.Y. (1997). "The phylogeny of varanoid lizards and the affinities of snakes". Philosophical Transactions: Biological Sciences. 352 (1349): 53–91. Bibcode:1997RSPTB.352...53L. doi:10.1098/rstb.1997.0005. PMC   1691912 .
  4. Townsend, T.M.; Larson, A.; Louis, E.; Macey, J.R. (2005). "Molecular phylogenetics of Squamata:the position of snakes, amphisbaenians, and dibamids, and the root of the squamate tree" (PDF). Systematic Biology. 53 (5): 735–757. doi: 10.1080/10635150490522340 . PMID   15545252.
  5. Vidal, N.; Hedges, S.B. (2005). "The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes" (PDF). Comptes Rendus Biologies. 328 (10–11): 1000–1008. doi:10.1016/j.crvi.2005.10.001. PMID   16286089. Archived from the original (PDF) on 2012-09-18.
  6. Reeder, Tod W.; Townsend, Ted M.; Mulcahy, Daniel G.; Noonan, Brice P.; Wood, Perry L.; Jr; Sites, Jack W.; Jr; Wiens, John J. (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.