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Sauropsida

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220px-Tortoise and the duck
Sauropsids

Temporal range: Pennsylvanian–Present, 312–0 MaPreЄ

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Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Amniota
Clade: Sauropsida

Watson, 1956

Subgroups
  • †Parareptilia
  • Eureptilia
    • Brouffia
    • Coelostegus
    • Thuringothyris
    • †Captorhinidae
    • Romeriida

Sauropsida ("lizard faces") is a group ofamniotes that includes all existing birdsand reptiles as well as their fossil ancestors and other extinct relatives. Large land animals are either in this group or in its sister group, Synapsida, which includes mammals and their fossil relatives. This clade includesParareptilia and other extinct clades. All living sauropsids are members of the sub-group Diapsida, the Parareptilia clade having died out 200 million years ago. The term originated in 1864 withThomas Henry Huxley,[1] who grouped birds with reptiles based on fossil evidence.

History of classification Edit

220px-Crystal Palace Dicynodon

Benjamin Waterhouse Hawkins' 1855 reconstruction of a Dicynodon as aturtle-like creature.

The term Sauropsida ("lizard faces") has a long history, and hails back toThomas Henry Huxley, and his opinion that birds had risen from the dinosaurs. He based this chiefly on the fossils ofHesperornis and Archaeopteryx, that were starting to become known at the time.[2] In the Hunterian lecturesdelivered at the Royal College of Surgeons in 1863, Huxley grouped the vertebrate classes informally intomammals, sauroids, and ichthyoids (the latter containing the anamniotes), based on the gaps in physiological traits and lack of transitional fossils that seem to exist between the three groups. Early in the following year he proposed the names Sauropsida and Ichthyopsida for the two latter.[1] Huxley did however include groups on the mammalian line (synapsids) like Dicynodon among the sauropsids. Thus, under the original definition, Sauropsida contained not only the groups usually associated with it today, but also several groups that today are known to be in the mammalian side of the tree.[3]

Sauropsids redefined Edit

By the early 20th century, the fossils ofPermian synapsids from South Africahad become well known, allowing palaeontologists to trace synapsid evolution in much greater detail. The term Sauropsida was taken up by E.S. Goodrich in 1916 much like Huxley's, to include lizards, birds and their relatives. He distinguished them from mammalsand their extinct relatives, which he included in the sister group Theropsida (now usually replaced with the nameSynapsida). Goodrich's classification thus differs somewhat from Huxley's, in which the non-mammalian synapsids (or at least the dicynodontians) fell under the sauropsids. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, the Protosauria ("first lizards"), which included somePaleozoic amphibians as well as earlyreptiles predating the sauropsid/synapsid split (and thus not true sauropsids).[3]

Detailing the reptile family tree Edit

In 1956, D.M.S. Watson observed that sauropsids and synapsids diverged very early in the reptilian evolutionary history, and so he divided Goodrich's Protosauria between the two groups. He also reinterpreted the Sauropsida and Theropsida to exclude birds and mammals respectively, making themparaphyletic, unlike Goodrich's definition. Thus his Sauropsida includedProcolophonia, Eosuchia, Millerosauria,Chelonia (turtles), Squamata[4] (lizards and snakes), Rhynchocephalia,Crocodilia, "thecodonts" (paraphyleticbasal Archosauria), non-aviandinosaurs, pterosaurs, ichthyosaurs, and sauropyterygians.[5]

This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer's classic Vertebrate Paleontology[6]) into four subclasses according to the positioning of temporal fenestrae, openings in the sides of the skull behind the eyes. Since the advent of phylogenetic nomenclature, the termReptilia has fallen out of favor with many taxonomists, who have used Sauropsida in its place to include amonophyletic group containing the traditional reptiles and the birds.

Cladistics and the Sauropsida Edit

220px-Reptilia-Sauropsida

Sauropsida and the traditional classReptilia superimposed on a cladogram of Tetrapods, showing the difference in coverage

The class Reptilia has been known to be an evolutionary grade rather than a clade for as long as evolution has been recognised. Reclassifying reptiles has been among the key aims ofphylogenetic nomenclature.[7] The term Sauropsida had from the mid 20th century been used to denote all species not on the synapsid side after the synapsid/sauropsid split, a branch-based clade. This group encompasses all now-living reptiles as well as birds, and as such is comparable to Goodrich's classification, the difference being that better resolution of the early amniote tree has split up most of the Goodrich's "Protosauria", though definitions of Sauropsida essentially identical to Huxley's (i.e. including the mammal-like reptiles) are also forwarded.[8][9] Some later cladistic work has used Sauropsida more restrictively, to signify the crown group, i.e. all descendants of the last common ancestor of extant reptiles and birds. A number of phylogenetic stem, node and crown definitions have been published, anchored in a variety of fossil and extant organisms, thus there is currently no consensus of the actual definition (and thus content) of Sauropsida as a phylogenetic unit.[10]

Some taxonomists, such as Benton (2004), have co-opted the term to fit into traditional rank-based classifications, making Sauropsida and Synapsida class-level taxa to replace the traditional Class Reptilia, while Modesto and Anderson (2004), using the PhyloCodestandard, have suggested replacing the name Sauropsida with their redefinition of Reptilia, arguing that the latter is by far better known and should have priority.[10]

Evolutionary history Edit

220px-Europasaurus holgeri Scene 2

Mesozoic sauropsids: the dinosaursEuropasaurus and Iguanodon, and the early bird Archaeopteryx perched on the foreground tree stump.

Sauropsids evolved from basal amniotes stock approximately 320 million years ago in the Paleozoic Era. In the Mesozoic Era (from about 250 million years ago to about 66 million years ago), sauropsids were the largest animals on land, in the water, and in the air. The Mesozoic is sometimes called the Age of Reptiles. Sixty-six million years ago, the large-bodied sauropsids died out in the global extinction event at the end of the Mesozoic era. With the exception of a few species of birds, the entire dinosaur lineage became extinct; in the following era, the Cenozoic, the remaining birds diversified so extensively that, today, nearly one out of every three species of land vertebrate is a bird species.

Phylogen Edit

The cladogram presented here illustrates the "family tree" of sauropsids, and follows a simplified version of the relationships found by M.S. Lee, in 2013.[11] All genetic studies have supported the hypothesis that turtles are diapsid reptiles; some have placed turtles within archosauriformes,[12][13][14][15][16][11]though a few have recovered turtles as lepidosauriformes instead.[17] The cladogram below used a combination of genetic (molecular) and fossil (morphological) data to obtain its results.[11]

Sauropsida
unnamed
†Parareptilia

†Millerettidae 

unnamed

Eunotosaurus

†Hallucicrania

†Lanthanosuchidae 

†Procolophonia

†Procolophonoidea 


†Pareiasauromorpha 





Eureptilia

†Captorhinidae 

Romeriida

Paleothyris

Diapsida

†Araeoscelidia 

Neodiapsida

Claudiosaurus



†Younginiformes 

Sauria
Lepidosauromorpha

†Kuehneosauridae

Lepidosauria

Rhynchocephalia (tuatara and their extinct relatives) 


Squamata (lizards and snakes) 



Archosauromorpha


†Choristodera 



†Prolacertiformes




Trilophosaurus 


†Rhynchosauria 



Archosauriformes(crocodiles, birds, and their extinct relatives) 




 Pantestudines 

†Eosauropterygia 



†Placodontia 



Sinosaurosphargis



Odontochelys

Testudinata

Proganochelys


Testudines(turtles) 






220px-Europasaurus holgeri Scene 2
220px-Crystal Palace Dicynodon

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