Sauria is the

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

diapsid Diapsids ("two arches") 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 earliest traditionally identified diapsids, the araeosc ...

s containing the

most recent common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...

of Archosauria (which includes

crocodilians Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchi ...

and

birds Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

) and

Lepidosauria The Lepidosauria (, from Greek meaning ''scaled lizards'') is a Order (biology), superorder or Class (biology), subclass of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata also includes Lizard, lizards and Snake, snakes. Sq ...

(which includes squamates and the

tuatara The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...

), and all its descendants. Since most molecular phylogenies recover turtles as more closely related to archosaurs than to lepidosaurs as part of

Archelosauria Archelosauria is a clade grouping turtles and archosaurs (birds and crocodilians) and their fossil relatives, to the exclusion of lepidosaurs (the clade containing lizards, snakes and the tuatara). The majority of phylogenetic analyses based on ...

, Sauria can be considered the

crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...

of diapsids, or

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

s in general. Depending on the systematics, Sauria includes all modern reptiles or most of them (including

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s, a type of archosaur) as well as various extinct groups. Sauria lies within the larger total group

Sauropsida Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...

, which also contains various

stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...

-reptiles which are more closely related to reptiles than to mammals. Prior to its modern usage, "Sauria" was used as a name for the

suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...

occupied by

lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...

s, which before 1800 were considered crocodilians.

Systematics

Sauria was historically used as a partial equivalent for

Squamata Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...

(which contains lizards and snakes). The redefinition to cover the last common ancestor of archosaurs and lepidosaurs was the result of papers by Jacques A. Gauthier and colleagues in the 1980s. Genomic studiesCrawford, Nicholas G., et al. "More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs." Biology letters 8.5 (2012): 783–786. and comprehensive studies in the fossil record suggest that

turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...

s are closely related to archosaurs as part of Sauria, and not to the non-saurian

parareptile Parareptilia ("near-reptiles") is an extinct group of Basal (phylogenetics), basal Sauropsida, sauropsids ("Reptile, reptiles"), traditionally considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds ...

s as previously thought.

Synapomorphies

The synapomorphies or characters that unite the clade Sauria also help them be distinguished from stem-saurians in Diapsida or stem-reptiles in clade

in the following categories based on the following regions of the body.Pough, F. H., Janis, C. M., & Heiser, J. B. (2005). Vertebrate life. Pearson/Prentice Hall. *Cephalad Region **Dorsal origin of temporal musculature **Loss of caniniform region in maxillary tooth row ** External nares close to the midline **

Postparietal Postparietals are cranial bones present in fish and many Tetrapod, tetrapods. Although initially a pair of bones, many lineages possess postparietals which were fused into a single bone. The postparietals were Dermal bone, dermal bones situated alo ...

absent **

Squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...

mainly restricted to top of skull **The occipital flange of the squamosal is little exposed on the occiput **Anterior process of squamosal narrow ** Quadrate exposed laterally **Unossified dorsal process of

stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other tetrapods which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the f ...

**Stapes slender *Trunk Region **Sacral ribs oriented laterally **

Ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...

fusion of caudal ribs **Trunk ribs mostly single headed *Pectoral Region **

Cleithrum The cleithrum (: cleithra) is a membrane bone which first appears as part of the skeleton in primitive Osteichthyes, bony fish, where it runs vertically along the scapula. Its name is derived from Greek κλειθρον = "key (lock)", by analogy ...

absent *Pelvic Region **Modified ilium *Limb Region **Tubular bone lost **Entepicondylar foramen absent **

Radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...

as long as

ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...

**Small proximal carpals and tarsal **Fifth distal tarsal absent **Short and stout fifth or hooked

metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...

**Perforating foramen of manus lost However, some of these characters might be lost or modified in several lineages, particularly among birds and turtles; it is best to see these characters as the ancestral features that were present in the ancestral saurian.

Phylogeny

The cladogram shown below follows the most likely result found by an analysis of turtle relationships using both fossil and genetic evidence by M.S. Lee, in 2013. This study found ''

Eunotosaurus ''Eunotosaurus'' (''Latin (language), Latin'': Stout-backed lizard) is an extinct genus of amniote, possibly a close relative of turtles. ''Eunotosaurus'' lived in the late Middle Permian (Capitanian stage) and fossils can be found in the Karoo ...

'', usually regarded as a turtle relative, to be only very distantly related to turtles in the clade

Parareptilia Parareptilia ("near-reptiles") is an extinct group of basal sauropsids (" reptiles"), traditionally considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds). Parareptiles first arose near the en ...

. The cladogram below follows the most likely result found by another analysis of turtle relationships, this one using only fossil evidence, published by Rainer Schoch and Hans-Dieter Sues in 2015. This study found ''Eunotosaurus'' to be an actual early stem-turtle, though other versions of the analysis found weak support for it as a parareptile. The cladogram below follows the analysis of Li ''et al''. (2018). It places turtles within Diapsida but outside of Sauria (the Lepidosauromorpha + Archosauromorpha clade). The following cladogram was found by Simões ''et al''. (2022):

References

External links

* * {{Taxonbar, from=Q2254408 Reptile taxonomy Tetrapod unranked clades Permian reptiles Extant Permian first appearances Guadalupian first appearances Taxa described in 1802