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


Characteristics
The name Diapsida means "two arches", and diapsids are traditionally classified based on their two ancestral skull openings (temporal fenestrae) posteriorly above and below the eye. This arrangement allows for the attachment of larger, stronger , and enables the jaw to open more widely. A more obscure ancestral characteristic is a relatively long lower arm bone (the radius) compared to the upper arm bone ().

Basal non-saurian neodiapsids were ancestrally lizard-like, but basal non-saurian neodiapsids include aquatic/amphibious taxa ( and some ) the -like , as well as the Triassic chameleon-like .


Classification
Diapsids were originally classified as one of four subclasses of the class , all of which were based on the number and arrangement of openings in the skull. The other three subclasses were (one opening low on the skull, for the "mammal-like reptiles"), (no skull opening, including turtles and their relatives), and (one opening high on the skull, including many prehistoric marine reptiles). With the advent of phylogenetic nomenclature, this system of classification was heavily modified. Today, the synapsids are often not considered true reptiles, while Euryapsida were found to be an unnatural assemblage of diapsids that had lost one of their skull openings. Genetic studies and the discovery of the Triassic have shown that this is also the case in turtles, which are actually heavily modified diapsids. In phylogenetic systems, birds (descendants of traditional diapsid reptiles) are also considered to be members of this group.

Some modern studies of reptile relationships have preferred to use the name "diapsid" to refer to the crown group of all modern diapsid reptiles but not their extinct relatives. However, many researchers have also favored a more traditional definition that includes the prehistoric . In 1991, Laurin defined Diapsida as a , "the most recent common ancestor of araeoscelidians, , and , and all its descendants".Benton, M. J., Donoghue, P. C., Asher, R. J., Friedman, M., Near, T. J., & Vinther, J. (2015). "Constraints on the timescale of animal evolutionary history." Palaeontologia Electronica, 18.1.1FC; 1-106; palaeo-electronica.org/content/fc-1

The clade Neodiapsida was given a definition by Laurin in 1991. He defined it as the branch-based containing all animals more closely related to "" (later, more specifically, emended to Youngina capensis) than to (representing ). The earliest known neodiapsids like are known from the Early , around 290 million years ago.

All genetic studies have supported the hypothesis that turtles are cladistically diapsid reptiles despite being morphologically anapsid, most placing them as more closely related to living archosaurs (including crocodiles and birds) than to lepidosaurs (lizards, snakes, etc).

Modern reptiles and birds are placed within the neodiapsid subclade , defined as the last common ancestor of Lepidosauria (which includes lizards, snakes and the tuatara), and Archosauria (which includes crocodilians and dinosaurs, including birds, among others).

A cladistic analysis by Laurin and Piñeiro (2017) recovers as part of Diapsida, with pareiasaurs, turtles, millerettids, and procolophonoids recovered as more derived than the basal diapsid . A 2020 study by David P. Ford and Roger B. J. Benson also recovered Parareptilia as deeply nested within Diapsida as the sister group to . They united this relationship between Parareptilia and Neodiapsida in the new clade , defining it as the last common ancestor and all descendants of Procolophon trigoniceps and Youngina capensis. However, this excludes , who were found to be basal among the sauropsids. Other recent studies have found the more traditional arrangement of parareptiles being outside of Diapsida.

The position of the highly derived Mesozoic marine reptile groups , Ichthyosauromorpha and within Neodiapsida is uncertain, and they may lie within Sauria.

In the 2022 and 2023 studies, Araeoscelidia was found to have no close relationship with Neodiapsida and was not even part of Sauropsida.


Relationships
Below are showing the relations of the major groups of diapsids.

Cladogram after Bickelmann et al., 2009 and Reisz et al., 2011:

The cladogram of Lee (2013) below used a combination of genetic (molecular) and fossil (morphological) data.

This second cladogram is based on the 2017 study by Pritchard and Nesbitt.

The following cladogram was found by Simões et al. (2022):


See also


External links
  • Diapsida. Michel Laurin and Jacques A. Gauthier. Tree of Life Web Project. June 22, 2000.

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