Diapsid reptiles Temporal range: Pennsylvanian–Present, | |
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Skull diagram of the araeoscelidian Petrolacosaurus kansensis | |
Nile crocodile (Crocodylus niloticus) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Romeriida |
Clade: | Diapsida Osborn, 1903 |
Subgroups | |
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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 araeoscelidians, appeared about three hundred million years ago during the late Carboniferous period. [1] 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. [2] All modern reptiles and birds are placed within the neodiapsid subclade Sauria. 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 species of diapsid animals are extant: 9,159 birds, [3] and 7,925 snakes, lizards, tuatara, turtles, and crocodiles. [4]
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 jaw muscles, 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 (humerus).
Basal non-saurian neodiaspids were ancestrally lizard-like, but basal non-saurian neodiapsids include aquatic/amphibious taxa ( Claudiosaurus and some tangasaurids) [5] the gliding lizard-like Weigeltisauridae, [6] as well as the Triassic chameleon-like drepanosaurs. [7]
Diapsids were originally classified as one of four subclasses of the class Reptilia, all of which were based on the number and arrangement of openings in the skull. The other three subclasses were Synapsida (one opening low on the skull, for the "mammal-like reptiles"), Anapsida (no skull opening, including turtles and their relatives), and Euryapsida (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 Pappochelys 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 araeoscelidians. In 1991, Laurin defined Diapsida as a clade, "the most recent common ancestor of araeoscelidians, lepidosaurs, and archosaurs, and all its descendants". [8]
The clade Neodiapsida was given a phylogenetic definition by Laurin in 1991. He defined it as the branch-based clade containing all animals more closely related to "Younginiformes" (later, more specifically, emended to Youngina capensis ) than to Petrolacosaurus (representing Araeoscelidia). [9] The earliest known neodiapsids like Orovenator are known from the Early Permian, around 290 million years ago. [10]
All genetic studies have supported the hypothesis that turtles are diapsid reptiles; some have placed turtles within archosauromorpha, [11] [12] or, more commonly, as a sister group to extant archosaurs. [13] [14] [15] [16]
Modern reptiles and birds are placed within the neodiapsid subclade Sauria, 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). [17]
A cladistic analysis by Laurin and Piñeiro (2017) recovers Parareptilia as part of Diapsida, with pareiasaurs, turtles, millerettids, and procolophonoids recovered as more derived than the basal diapsid Younginia . [18] 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 Neodiapsida. They united this relationship between Parareptilia and Neodiapsida in the new clade Neoreptilia, defining it as the last common ancestor and all descendants of Procolophon trigoniceps and Youngina capensis . [19] However, this excludes mesosaurs, who were found to be basal among the sauropsids. [19] Other recent studies have found the more traditional arrangement of parareptiles being outside of Diapsida. [17]
The position of the highly derived Mesozoic marine reptile groups Thalattosauria, Ichthyosauromorpha and Sauropterygia within Neodiapsida is uncertain, and they may lie within Sauria. [17]
Below are cladograms showing the relations of the major groups of diapsids.
Cladogram after Bickelmann et al., 2009 [20] and Reisz et al., 2011: [21]
The cladogram of Lee (2013) below used a combination of genetic (molecular) and fossil (morphological) data. [22]
Diapsida |
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This second cladogram is based on the 2017 study by Pritchard and Nesbitt. [23]
Neodiapsida |
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The following cladogram was found by Simões et al. (2022): [17]
Neodiapsida |
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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.
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the Carboniferous period. Those of Amniota are defined as the smallest crown clade containing humans, the Greek tortoise, and the Nile crocodile.
Sauria is the clade of diapsids 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.
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.
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.
Eureptilia is one of the two major subgroups of the clade Sauropsida, the other one being Parareptilia. Eureptilia includes Diapsida, as well as a number of primitive Permo-Carboniferous forms previously classified under Anapsida, in the old order "Cotylosauria".
Youngina is an extinct genus of diapsid reptile from the Late Permian Beaufort Group of the Karoo Red Beds of South Africa. This, and a few related forms, make up the family Younginidae, within the order Eosuchia. Eosuchia, having become a wastebasket taxon for many probably distantly-related primitive diapsid reptiles ranging from the Late Carboniferous to the Eocene, Romer proposed that it be replaced by Younginiformes.
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 artificial or unnatural grouping. However, the clade was recovered again in 2021 following a redescription of Weigeltisaurus, raising the possibility that the clade may be valid after all, although subsequent analyses have not supported this result.
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".
Claudiosaurus is an extinct genus of diapsid reptiles from the Late Permian Sakamena Formation of the Morondava Basin, Madagascar. It has been suggested to be semi-aquatic.
Araeoscelidia or Araeoscelida is a clade of extinct amniotes superficially resembling lizards, extending from the Late Carboniferous to the Early Permian. The group contains the genera Araeoscelis, Petrolacosaurus, the possibly aquatic Spinoaequalis, and less well-known genera such as Kadaliosaurus and Zarcasaurus. This clade is usually considered to be the sister group to all later diapsids.
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.
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.
Orovenator is an extinct genus of diapsid from Lower Permian deposits of Oklahoma, United States. It is known from two partial skulls from the Richards Spur locality in Oklahoma. The holotype OMNH 74606 consists of a partial skull preserving snout and mandible, and the referred specimen, OMNH 74607, a partial skull preserving the skull roof, vertebrae and palatal elements. It was first named by Robert R. Reisz, Sean P. Modesto and Diane M. Scott in 2011 and the type species is Orovenator mayorum. The generic name means "mountain", oro, in Greek in reference to the Richards Spur locality, which was mountainous during the Permian period and "hunter", venator, in Latin. The specific name honours Bill and Julie May. Orovenator is the oldest and most basal neodiapsid to date.
Younginidae is an extinct family of diapsid reptiles from the Late Permian and Early Triassic. In a phylogenetic context, younginids are near the base of the clade Neodiapsida. Younginidae includes the species Youngina capensis from the Late Permian of South Africa and Thadeosaurus colcanapi from the Late Permian and Early Triassic of Madagascar. Heleosuchus griesbachi from the Late Permian of South Africa may also be a member of the family.
Pantestudines or Pan-Testudines is the proposed 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.