Anapsid

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Anapsids
Temporal range: 312–0  Ma
Skull anapsida 1.png
Anapsid skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Informal group: Anapsida
Williston, 1917
Subgroups

An anapsid is an amniote whose skull lacks one or more skull openings (fenestra, or fossae) near the temples. [1] 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.

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Anapsids and the turtles

Anapsid skull of Caretta caretta (Loggerhead sea turtle), a Testudine Caretta carettaZZ.jpg
Anapsid skull of Caretta caretta (Loggerhead sea turtle), a Testudine

While "anapsid reptiles" or "anapsida" were traditionally spoken of as if they were a monophyletic group, it has been suggested that several groups of reptiles that had anapsid skulls might be only distantly related. Scientists still debate the exact relationship between the basal (original) reptiles that first appeared in the late Carboniferous, the various Permian reptiles that had anapsid skulls, and the Testudines (turtles, tortoises, and terrapins). However, it was later suggested that the anapsid-like turtle skull is due to reversion rather than to anapsid descent. The majority of modern paleontologists believe that the Testudines are descended from diapsid reptiles that lost their temporal fenestrae. More recent morphological phylogenetic studies with this in mind placed turtles firmly within diapsids, [2] [3] [4] [5] [6] or, more commonly, within Archelosauria. [7]

Phylogenetic position of turtles

All molecular studies have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria, [8] or, more commonly, as a sister group to extant archosaurs. [9] [10] [11] [12] [13] One of the most recent molecular studies, published on 23 February 2012, suggests that turtles are lepidosauromorph diapsids, most closely related to the lepidosaurs (lizards, snakes, and tuataras). [14] However, in a later paper from the same authors, published in 2014, based on more extensive data, the archosauromorph hypothesis is supported. [7]

Reanalysis of prior phylogenies suggests that they classified turtles as anapsids both because they assumed this classification (most of them were studying what sort of anapsid turtles are) and because they did not sample fossil and extant taxa broadly enough for constructing the cladogram. Testudines is suggested to have diverged from other diapsids between 200 and 279 million years ago, though the debate is far from settled. [15] [9] [16] Although procolophonids managed to survive into the Triassic, most of the other reptiles with anapsid skulls, including the millerettids, nycteroleterids, and pareiasaurs, became extinct in the Late Permian period by the Permian-Triassic extinction event.

Despite the molecular studies, there is evidence that contradicts their classification as diapsids. All known diapsids excrete uric acid as nitrogenous waste (uricotelic), and there is no known case of a diapsid reverting to the excretion of urea (ureotelism), even when they return to semi-aquatic lifestyles. Crocodilians, for example, are still uricotelic, although they are also partly ammonotelic, meaning they excrete some of their waste as ammonia. Ureotelism appears to be the ancestral condition among primitive amniotes, and it is retained by mammals, which likely inherited ureotelism from their synapsid and therapsid ancestors. Ureotelism therefore would suggest that turtles were more likely anapsids than diapsids. The only known uricotelic chelonian is the desert tortoise, which likely evolved it recently as adaptation to desert habitats. Some desert mammals are also uricotelic, so since practically all known mammals are ureotelic, uricotelic adaptation is a likely result of convergence among desert species. Therefore, turtles would have to be the only known case of a uricotelic reptile reverting to ureotelism.[ citation needed ]

Anapsida in modern taxonomy

Anapsida is still sporadically recognized as a valid group, but this is not favoured by current workers. [17] [18] Anapsids in the traditional meaning of the word are not a clade, but rather a paraphyletic group composed of all the early reptiles retaining the primitive skull morphology, grouped together by the absence of temporal openings. [17] [18] Gauthier, Kluge and Rowe (1988) attempted to redefine Anapsida so it would be monophyletic, defining it as the clade containing "extant turtles and all other extinct taxa that are more closely related to them than they are to other reptiles". [19]

This definition explicitly includes turtles in Anapsida; because the phylogenetic placement of turtles within Amniota is very uncertain, it is unclear what taxa, other than turtles themselves, would be included in such defined Anapsida, and whether its content would be similar to the Anapsida of tradition. Indeed, Gauthier, Kluge and Rowe (1988) themselves included only turtles and Captorhinidae in their Anapsida, while excluding the majority of anapsids in the traditional sense of the word from it. [19]

Temporal openings in traditional anapsids

Tsuji and Müller (2009) noted that the name Anapsida implies a morphology (lack of temporal openings) that is in fact absent in the skeletons of a number of taxa traditionally included in the group. [18] A temporal opening in the skull roof behind each eye, similar to that present in the skulls of synapsids, has been discovered in the skulls of a number of members of Parareptilia (the group containing most of reptiles traditionally referred to as anapsids), including lanthanosuchoids, millerettids, bolosaurids, some nycteroleterids, some procolophonoids and at least some mesosaurs. [18] [20] [21] The presence of temporal openings in the skulls of these taxa makes it uncertain whether the ancestral reptiles had an anapsid-like skull as traditionally assumed or a synapsid-like skull instead. [21]

See also

Related Research Articles

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

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 amphibian ancestors during the Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids. They are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration.

<span class="mw-page-title-main">Sauria</span> Clade of reptiles

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.

<span class="mw-page-title-main">Diapsid</span> Clade of amniote tetrapods with two holes in each side of their skulls

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.

<span class="mw-page-title-main">Sauropsida</span> Taxonomic clade

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.

<span class="mw-page-title-main">Mesosaur</span> Extinct family of reptiles

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.

<span class="mw-page-title-main">Archosauromorpha</span> Infraclass of reptiles

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.

<span class="mw-page-title-main">Neodiapsida</span> Clade of reptiles

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.

<span class="mw-page-title-main">Eureptilia</span> Clade of reptiles

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".

<span class="mw-page-title-main">Pareiasauria</span> Extinct clade of reptiles

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.

<span class="mw-page-title-main">Euryapsida</span>

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.

<span class="mw-page-title-main">Parareptilia</span> Subclass of reptiles

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.

<span class="mw-page-title-main">Procolophonia</span> Extinct suborder of reptiles

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".

<span class="mw-page-title-main">Procolophonomorpha</span> Order of reptiles (fossil)

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.

<span class="mw-page-title-main">Temporal fenestra</span> Opening in the skull behind the orbit in some animals

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.

<i>Eunotosaurus</i> Extinct genus of reptiles

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.

<i>Acleistorhinus</i> Extinct genus of reptiles

Acleistorhinus (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian of Oklahoma. It is notable for being the earliest known anapsid reptile yet discovered. The morphology of the lower temporal fenestra of the skull of Acleistorhinus bears a superficial resemblance to that seen in early synapsids, a result of convergent evolution. Only a single species, A. pteroticus, is known, and it is classified in the Family Acleistorhinidae, along with Colobomycter.

<i>Owenetta</i> Extinct genus of reptiles

Owenetta is an extinct genus of owenettid procolophonian parareptile. Fossils have been found from the Beaufort Group in the Karoo Basin of South Africa. Although most procolophonians lived during the Triassic, Owenetta existed during the Wuchiapingian and Changhsingian stages of the Late Permian as well as the early Induan stage of the Early Triassic. It is the type genus of the family Owenettidae, and can be distinguished from other related taxa in that the posterior portion of the supratemporal bears a lateral notch and that the pineal foramen is surrounded by a depressed parietal surface on the skull table.

<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

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.

<span class="mw-page-title-main">Pantestudines</span> Clade of reptiles

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.

<i>Pappochelys</i> Extinct genus of reptiles

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.

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