An anapsid is an amniote whose skull lacks one or more skull openings near the temples. Traditionally, the Anapsida are considered 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.
Sauria is the clade containing the most recent common ancestor of Archosauria and Lepidosauria, and all its descendants. 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. Depending on the systematics, Sauria includes all modern reptiles or most of them as well as various extinct groups.
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 earliest traditionally identified diapsids, the araeoscelidians, 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 often 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. All modern reptiles and birds are placed within the neodiapsid subclade Sauria. 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 also include extinct stem-group relatives of modern reptiles and 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 recognized as a subgroup of archosaurian reptiles despite originally being named as a separate class in Linnaean taxonomy.
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. It is possible that sauropterygians are a distant relatives of turtles, uniting them under the group Pantestudines, although this is still debatable as sauropterygians might be archosauromorphs or completely unrelated to both.
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.
Pareiasaurs are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with osteoderms which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, before becoming globally distributed during the Late Permian. Pareiasaurs were the largest reptiles of the Permian, reaching sizes equivalent to those of contemporary therapsids. Pareiasaurs became extinct in the Permian–Triassic extinction event.
Euryapsida is a polyphyletic group of sauropsids that are distinguished by a single temporal fenestra, an opening behind the orbit, under which the post-orbital and squamosal bones articulate. They are different from Synapsida, which also have a single opening behind the orbit, by the placement of the fenestra. In synapsids, this opening is below the articulation of the post-orbital and squamosal bones. It is now commonly believed that euryapsids are in fact diapsids that lost the lower temporal fenestra. Euryapsids are usually considered entirely extinct, although turtles might be part of the sauropterygian clade while other authors disagree. Euryapsida may also be a synonym of Sauropterygia sensu lato.
Parareptilia ("near-reptiles") is an extinct 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.
Procolophonia is an extinct suborder (clade) of herbivorous reptiles that lived from the Middle Permian till the end of the Triassic period. They were originally included as a suborder of the Cotylosauria but are now considered a clade of Parareptilia. They are closely related to other generally lizard-like Permian reptiles such as the Millerettidae, Bolosauridae, Acleistorhinidae, and Lanthanosuchidae, all of which are included under the Anapsida or "Parareptiles".
Procolophonomorpha is an order or clade containing most parareptiles. Many papers have applied various definitions to the name, though most of these definitions have since been considered synonymous with modern parareptile clades such as Ankyramorpha and Procolophonia. The current definition of Procolophonomorpha, as defined by Modesto, Scott, & Reisz (2009), is that of as a stem-based group containing Procolophon and all taxa more closely related to it than to Milleretta. It constitutes a diverse assemblage that includes a number of lizard-like forms, as well as more diverse types such as the pareiasaurs. Lee 1995, 1996, 1997 argues that turtles evolved from pareiasaurs, but this view is no longer considered likely. Rieppel and deBraga 1996 and deBraga and Rieppel, 1997 argue that turtles evolved from sauropterygians, and there is both molecular and fossil (Pappochelys) evidence for the origin of turtles among diapsid reptiles.
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 and Malawi. Eunotosaurus resided in the swamps of what is now 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.
Testudinata is the group of all tetrapods with a true turtle shell. It includes both modern turtles (Testudines) and many of their extinct, shelled relatives (stem-turtles), though excluding Odontochelys and Eorhynchochelys, which are placed in the more inclusive Pantestudines.
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.
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.
Perichelydia is a clade within Pantestudines known from the Middle Jurassic to Holocene. Alongside crown group Testudines, it also contains Helochelydridae, which is known from the Cretaceous of Europe and North America, Sichuanchelyidae from the Middle Jurassic to Paleocene of Asia and Europe, Meiolaniformes, which is known from the Cretaceous to Holocene of South America, Australia and Oceania, and Spoochelys, known from the Mid-Cretaceous Griman Creek Formation of Australia. Kallokibotion from the Late Cretaceous of Europe is also considered part of this group. Several other groups, including the proposed clade Angolachelonia, Paracryptodira, Macrobaenidae, Sinemydidae and Xinjiangchelyidae, which are sometimes considered members of Cryptodira, have also been found outside crown Testudines in several analyses. These groups are usually considered to be closer to the crown group than the other members of Perichelydia.
