Saurians | |
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clockwise from top left: Agkistrodon contortrix (the copperhead, a snake), Dinemellia dinemelli (the white-faced buffalo-weaver, a bird), various extinct ornithischian dinosaurs, Chelonia mydas (the green sea turtle), Anurognathus (an extinct pterosaur), and Alligator mississippiensis (the american alligator, a crocodilian) Contents | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Neodiapsida |
Clade: | Sauria Macartney, 1802 |
Groups | |
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Sauria is the clade containing the most recent common ancestor of Archosauria (which includes crocodilians and birds) and Lepidosauria (which includes squamates and the tuatara), and all its descendants. [1] Since most molecular phylogenies recover turtles as more closely related to archosaurs than to lepidosaurs as part of Archelosauria, Sauria can be considered the crown group of diapsids, or reptiles in general. [2] Depending on the systematics, Sauria includes all modern reptiles [3] or most of them (including birds, a type of archosaur) as well as various extinct groups. [4]
Sauria lies within the larger total group Sauropsida, which also contains various stem-reptiles which are more closely related to reptiles than to mammals. [3] Prior to its modern usage, "Sauria" was used as a name for the suborder occupied by lizards, which before 1800 were considered crocodilians.
Genomic studies [5] [6] [7] and comprehensive studies in the fossil record [8] suggest that turtles are closely related to archosaurs, not to the pre-Saurian parareptiles as previously thought. In a 2018 cladistic analysis, Pantestudines (turtles and close relatives) were placed within Diapsida but outside of Sauria. [4]
The synapomorphies or characters that unite the clade Sauria also help them be distinguished from stem-saurians in Diapsida or stem-reptiles in clade Sauropsida in the following categories based on the following regions of the body. [9] [10] [11]
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. [9]
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 , usually regarded as a turtle relative, to be only very distantly related to turtles in the clade Parareptilia. [8]
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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. [12]
Sauria |
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The cladogram below follows the analysis of Li et al. (2018). It places turtles within Diapsida but outside of Sauria (the Lepidosauromorpha + Archosauromorpha clade). [4]
Eureptilia |
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Reptiles, as commonly defined, are a group of tetrapods with an ectothermic ('cold-blooded') metabolism and amniotic development. Living reptiles comprise four orders: Testudines (turtles), Crocodilia (crocodilians), Squamata, and Rhynchocephalia. As of May 2023, about 12,000 living species of reptiles are listed in the Reptile Database. The study of the traditional reptile orders, customarily in combination with the study of modern amphibians, is called herpetology.
An anapsid is an amniote whose skull lacks one or more skull openings near the temples. Traditionally, the Anapsida are the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolved, making anapsids paraphyletic. It is however doubtful that all anapsids lack temporal fenestra as a primitive trait, and that all the groups traditionally seen as anapsids truly lacked fenestra.
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.
Sauropsida is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to include both extinct stem-group relatives of modern reptiles, as well as birds. The most popular definition states that Sauropsida is the sibling 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.
The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.
Mesosaurs were a group of small aquatic reptiles that lived during the early Permian period (Cisuralian), roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to an aquatic lifestyle from more terrestrial ancestors. It is uncertain which and how many terrestrial traits these ancestors displayed; recent research cannot establish with confidence if the first amniotes were fully terrestrial, or only amphibious. Most authors consider mesosaurs to have been aquatic, although adult animals may have been amphibious, rather than completely aquatic, as indicated by their moderate skeletal adaptations to a semiaquatic lifestyle. Similarly, their affinities are uncertain; they may have been among the most basal sauropsids or among the most basal parareptiles.
Sauropterygia is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria became extinct at the end of that period. The plesiosaurs would continue to diversify until the end of the Mesozoic. Sauropterygians are united by a radical adaptation of their pectoral girdle, adapted to support powerful flipper strokes. Some later sauropterygians, such as the pliosaurs, developed a similar mechanism in their pelvis.
Archosauromorpha is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs rather than lepidosaurs. Archosauromorphs first appeared during the late Middle Permian or Late Permian, though they became much more common and diverse during the Triassic period.
Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade Sauria.
Avicephala is a potentially polyphyletic grouping of extinct diapsid reptiles that lived during the Late Permian and Triassic periods characterised by superficially bird-like skulls and arboreal lifestyles. As a clade, Avicephala is defined as including the gliding weigeltisaurids and the arboreal drepanosaurs to the exclusion of other major diapsid groups. This relationship is not recovered in the majority of phylogenetic analyses of early diapsids and so Avicephala is typically regarded as an unnatural grouping. However, the clade was recovered again in 2021 in a redescription of Weigeltisaurus, raising the possibility that the clade may be valid after all.
Parareptilia ("near-reptiles") is a subclass or clade of basal sauropsids/reptiles, typically considered the sister taxon to Eureptilia. Parareptiles first arose near the end of the Carboniferous period and achieved their highest diversity during the Permian period. Several ecological innovations were first accomplished by parareptiles among reptiles. These include the first reptiles to return to marine ecosystems (mesosaurs), the first bipedal reptiles, the first reptiles with advanced hearing systems, and the first large herbivorous reptiles. The only parareptiles to survive into the Triassic period were the procolophonoids, a group of small generalists, omnivores, and herbivores. The largest family of procolophonoids, the procolophonids, rediversified in the Triassic, but subsequently declined and became extinct by the end of the period.
Drepanosaurs are a group of extinct reptiles that lived between the Carnian and Rhaetian stages of the late Triassic Period, approximately between 230 and 210 million years ago. The various species of drepanosaurid were characterized by specialized grasping limbs and often prehensile tails, adaptions for arboreal (tree-dwelling) and fossorial (digging) lifestyles, with some having also been suggested to be aquatic. Fossils of drepanosaurs have been found in Arizona, New Mexico, New Jersey, Utah, England, and northern Italy. The name is taken from the family's namesake genus Drepanosaurus, which means "sickle lizard," a reference to their strongly curved claws.
Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit. These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly seen in the fossilized skulls of dinosaurs and other sauropsids. The major reptile group Diapsida, for example, is defined by the presence of two temporal fenestrae on each side of the skull. The infratemporal fenestra, also called the lateral temporal fenestra or lower temporal fenestra, is the lower of the two and is exposed primarily in lateral (side) view.
Acerosodontosaurus is an extinct genus of neodiapsid reptiles that lived during the Late Permian of Madagascar. The only species of Acerosodontosaurus, A. piveteaui, is known from a natural mold of a single partial skeleton including a crushed skull and part of the body and limbs. The fossil was discovered in deposits of the Lower Sakamena Formation. Based on skeletal characteristics, it has been suggested that Acerosodontosaurus individuals were at least partially aquatic.
Eunotosaurus is an extinct genus of amniote, possibly a close relative of turtles. Eunotosaurus lived in the late Middle Permian and fossils can be found in the Karoo Supergroup of South Africa. Eunotosaurus resided in the swamps of southern Africa. Its ribs were wide and flat, forming broad plates similar to a primitive turtle shell, and the vertebrae were nearly identical to those of some turtles. Accordingly, it is often considered as a possible transitional fossil between turtles and their prehistoric ancestors. However, it is possible that these turtle-like features evolved independently of the same features in turtles, since other anatomical studies and phylogenetic analyses suggest that Eunotosaurus may instead have been a parareptile, an early-diverging neodiapsid unrelated to turtles, or a synapsid.
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.
Archelosauria is a clade grouping turtles and archosaurs and their fossil relatives, to the exclusion of lepidosaurs. The majority of phylogenetic analyses based on molecular data have supported a sister-group relationship between turtles and archosaurs. On the other hand, Archelosauria had not been historically supported by most morphological analyses, which have instead found turtles to either be descendants of parareptiles, early-diverging diapsids outside of Sauria, or close relatives of lepidosaurs within the clade Ankylopoda. Some recent morphological analyses have also found support for Archelosauria.
Pantestudines or Pan-Testudines is the group of all reptiles more closely related to turtles than to any other living animal. It includes both modern turtles and all of their extinct relatives. Pantestudines with a complete shell are placed in the clade Testudinata.
Pappochelys is an extinct genus of diapsid reptile possibly related to turtles. The genus contains only one species, Pappochelys rosinae, from the Middle Triassic of Germany, which was named by paleontologists Rainer Schoch and Hans-Dieter Sues in 2015. The discovery of Pappochelys provides strong support for the placement of turtles within Diapsida, a hypothesis that has long been suggested by molecular data, but never previously by the fossil record. It is morphologically intermediate between the definite stem-turtle Odontochelys from the Late Triassic of China and Eunotosaurus, a reptile from the Middle Permian of South Africa.
Ankylopoda was a proposed clade that hypothetically contains turtles and lepidosaurs and their fossil relatives. This clade was historically supported based on microRNA analysis as well as some cladistic analyses. However, it was strongly contradicted by molecular evidence which supports Archelosauria, and other recent cladistic analyses have supported Archelosauria over Ankylopoda.