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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 extinct stem-group relatives of modern reptiles. The most popular definition states that Sauropsida is the sister 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.
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.
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco, lepospondyls lived from the Early Carboniferous (Mississippian) to the Early Permian and were geographically restricted to what is now Europe and North America. Five major groups of lepospondyls are known: Adelospondyli; Aïstopoda; Lysorophia; Microsauria; and Nectridea. Lepospondyls have a diverse range of body forms and include species with newt-like, eel- or snake-like, and lizard-like forms. Various species were aquatic, semiaquatic, or terrestrial. None were large, and they are assumed to have lived in specialized ecological niches not taken by the more numerous temnospondyl amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by Karl Alfred von Zittel, who coined the name to include some tetrapods from the Paleozoic that shared some specific characteristics in the notochord and teeth. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota. It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive 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 at the end of the Permian during the Permian-Triassic extinction event.
Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an Archaeothyris-like synapsid in the Late Carboniferous. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.
Ophiacodontidae is an extinct family of early eupelycosaurs from the Carboniferous and Permian. Archaeothyris, and Clepsydrops were among the earliest ophiacodontids, appearing in the Late Carboniferous. Ophiacodontids are among the most basal synapsids, an offshoot of the lineage which includes therapsids and their descendants, the mammals. The group became extinct by the Middle Permian, replaced by anomodonts, theriodonts, and the diapsid reptiles.
Parareptilia ("near-reptiles") is a subclass or clade of basal sauropsids, 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.
The Procolophonia are a suborder 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.
Araeoscelidia or Araeoscelida is a clade of extinct diapsid reptiles 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 considered to be the sister group to all later diapsids.
Robert Rafael Reisz is a Canadian paleontologist and specialist in the study of early amniote and tetrapod evolution.
Acheloma is an extinct genus of temnospondyl that lived during the Early Permian. The type species is A. cumminsi.
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.
The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives, but excluding the crown group Tetrapoda. They are thus paraphyletic, though acceptable in phylogenetic nomenclature as the group is defined by strict reference to phylogeny rather than to traits as in traditional systematics. Thus, some finned sarcopterygians are considered to be stem tetrapods.
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.
Apsisaurus is an extinct genus of Early Permian varanopid synapsids known from Texas of the United States. It was first named by Michel Laurin in 1991 and the type species is Apsisaurus witteri. Apsisaurus witteri is known from the holotype MCZ 1474, a three-dimensionally preserved partial skeleton including an incomplete skull and mandibles. The skull roof of Archeria is also articulated to the postcranial skeleton. It was collected in the Archer City Bonebed 1 site, from the Archer City Formation of the Wichita Group, dating to the Early Permian epoch. Apsisaurus was formerly assigned as an "eosuchian" diapsid. In 2010, it was redescribed by Robert R. Reisz, Michel Laurin and David Marjanović; their phylogenetic analysis found it to be a basal varanopid synapsid.
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.
References
- 1 2 "Michel Laurin". on the website of the Muséum national d'Histoire naturelle . Retrieved January 8, 2013.
- ↑ Laurin, Michel (1991). "The osteology of a Lower Permian eosuchian from Texas and a review of diapsid phylogeny". Zoological Journal of the Linnean Society. 101: 59–95. doi:10.1111/j.1096-3642.1991.tb00886.x.
- ↑ Modesto, Sean P.; Reisz, Robert R. (2003). "An enigmatic new diapsid reptile from the Upper Permian of eastern Europe". Journal of Vertebrate Paleontology. 22 (4): 851–855. doi:10.1671/0272-4634(2002)022[0851:AENDRF]2.0.CO;2. S2CID 131566543.
- ↑ Laurin, Michel; Reisz, Robert R. (1995). "A reevaluation of early amniote phylogeny" (PDF). Zoological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x. Archived from the original (PDF) on 2019-06-08. Retrieved 2012-12-06.
- ↑ Laurin, M. (1996) Phylogeny of Stegocephalians, from the Tree of Life Web Project
- ↑ "Comptes Rendus Palevol". Elsevier. Retrieved January 9, 2013.
- ↑ "Journal of Evolutionary Biology". Journal of Evolutionary Biology. CiteSeerX 10.1.1.523.9731 . doi:10.1111/(ISSN)1420-9101 . Retrieved January 8, 2013.
- ↑ ISPN council, official list from International Society for Phylogenetic Nomenclature
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