Nyctiphruretidae Temporal range: Early-Middle Permian, | |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | † Parareptilia |
Order: | † Procolophonomorpha |
Node: | † Ankyramorpha |
Suborder: | † Procolophonia |
Clade: | † Nyctiphruretia Lee, 1997 |
Family: | † Nyctiphruretidae Efremov, 1938 |
Genera | |
Nyctiphruretidae is an extinct family of hallucicranian parareptiles known from the late Early to the late Middle Permian of European Russia and south-central United States. [1]
Nyctiphruretidae was named by Efremov in 1938. The type genus is Nyctiphruretus known from several Middle Permian locations in European Russia. Michael S. Y. Lee in 1997 placed the family in the larger clade Nyctiphruretia. The order was defined to include both nyctiphruretids and nycteroleterids. [2] However, recent cladistic analyses suggest that nycteroleterids are more closely related to the Pareiasauria instead, thus making Nyctiphruretus the only genus of its order. [3] [4] In 2014, MacDougall & Reisz described and named a second genus of Nyctiphruretidae, Abyssomedon , from the middle Leonardian stage of the late Early Permian of Comanche County, Oklahoma, south-central United States. It contains a single species, A. williamsi, which represents the first nyctiphruretid known from North America, and the oldest species of the family. [1]
Before the discovery of Abyssomedon, all phylogenetic analyses including Säilä (2010) and Tsuji et al. (2012) recovered Nyctiphruretidae, which was represented solely by N. acudens, as the sister taxon of Procolophonia. [3] [4] However, the addition of Abyssomedon by MacDougall and Reisz (2014) to an updated version of the Tsuji et al. (2012) data-set, recovered a monophyletic Nyctiphruretidae, nested within Procolophonia as the sister taxon of Procolophonoidea - a clade formed by owenettids and procolophonids. The following cladogram is simplified after the phylogenetic analysis of MacDougall and Reisz (2014) and shows the placement of Nyctiphruretidae within Parareptilia. Relationships within bolded terminal clades are not shown. [1]
Parareptilia |
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Mesosaurs were a group of small aquatic reptiles that lived during the early Permian period, 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.
Pareiasaurs are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with scutes 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.
Parareptilia 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.
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.
Nyctiphruretus is an extinct genus of nyctiphruretid parareptile known from the Guadalupian series of European Russia.
Rhipaeosaurus is an extinct genus of nycteroleterid parareptile known from an articulated skeleton from the mid Middle Permian of European Russia. It contained a single species, Rhipaeosaurus tricuspidens. A bayesian analysis suggests that it is more closely related to pareiasaurs than to the other nycteroleterids, due to skull and tooth features. For this reason, "Nycteroleteridae" may be a grade rather than a clade, unless redefined to exclude Rhipaeosaurus.
Colobomycter is an extinct genus of lanthanosuchoid parareptile known from the Early Permian of Oklahoma.
Delorhynchus is an extinct genus of lanthanosuchoid parareptile known from the late Early Permian Garber Formation of Comanche County, Oklahoma. It contains three species: the type species D. priscus is based on a series of maxillae. The second species to be described, D. cifellii, is known from a larger number of well-preserved skulls and skeletal material. The third species, D. multidentatus, is based on a fragmentary skull with several rows of teeth on its jaw.
Acleistorhinidae is an extinct family of Late Carboniferous and Early Permian-aged parareptiles. Acleistorhinids are most diverse from the Richards Spur locality of the Early Permian of Oklahoma. Richards Spur acleistorhinids include Acleistorhinus, Colobomycter, and possibly Delorhynchus and Feeserpeton. Other taxa include Carbonodraco from the Late Carboniferous of Ohio and Karutia from the Early Permian of Brazil. Acleistorhinidae is commonly considered a subgroup of lanthanosuchoids, related to taxa such as Chalcosaurus, Lanthaniscus and Lanthanosuchus. However, a re-examination of parareptile phylogeny conducted by Cisneros et al. (2021) argued that lanthanosuchids were not closely related to acleistorhinids. The phylogenetic analysis conducted by these authors recovered acleistorhinids as the sister group of the clade Procolophonia, while lanthanosuchids were recovered within the procolophonian subgroup Pareiasauromorpha.
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.
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.
Microleter is an extinct genus of basal procolophonomorph parareptiles which lived in Oklahoma during the Early Permian period. The type and only known species is Microleter mckinzieorum. Microleter is one of several parareptile taxa described from the Richards Spur fissure fills, and can be characterized from its high tooth count, lacrimal/narial contact, short postfrontal, and slit-like temporal emargination edged by the postorbital, jugal, squamosal, and quadratojugal. Contrary to Australothyris, which had a similar phylogenetic position as a basal procolophonomorph, Microleter suggests that early parareptile evolution occurred in Laurasia and that multiple lineages developed openings or emarginations in the temporal region.
Ankyramorpha is an extinct clade of procolophonomorph parareptiles which lived between the early Cisuralian epoch to the latest Triassic period of Africa, Antarctica, Asia, Australia, Europe, North America and South America.
Lanthanosuchoidea is an extinct superfamily of ankyramorph parareptiles from the middle Pennsylvanian to the middle Guadalupian epoch of Europe, North America and Asia. It was named by the Russian paleontologist Ivachnenko in 1980, and it contains two families Acleistorhinidae and Lanthanosuchidae.
Lanthaniscus is an extinct genus of lanthanosuchoid ankyramorph parareptile known from the Guadalupian epoch of Eastern Europe, Russia. Lanthaniscus was first named by M. F. Ivakhnenko in 1980 and the type species is Lanthaniscus efremovi. L. efremovi was originally described on the basis of the holotype PIN 3706/9 from Peza-1 locality, Krasnoshchel' Formation, of Arkhangelsk. Various authors had assigned it to the family Lanthanosuchidae; however, Ivakhnenko, who described an additional specimen of L. efremovi in 2008, assigned Lanthaniscus to its own family, the Lanthaniscidae. The additional specimen PIN 4543/2, was collected from the same formation as the holotype, from the Nisogora locality, which is slightly younger in age.
Pareiasauromorpha is a group of parareptilian amniotes from the Permian. It includes genera found all over the world, with many genera from Asia and South Africa. The clade was first used as a group by Linda A. Tsuji in 2011, in order to group the family Nycteroleteridae (nycteroleters) and the superfamily Pareiasauroidea (pareiasaurs). Pareiasauromorpha is considered to be a monophyletic node, the sister group to procolophonoids.
Pappochelys is an extinct genus of diapsid reptile closely 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.
Abyssomedon is an extinct genus of a nyctiphruretid parareptile known from Early Permian fissure fills at Richards Spur in Comanche County, Oklahoma, south-central United States. It contains a single species, Abyssomedon williamsi, which represents oldest known nyctiphrureptid species and the first to be discovered in North America.
Richards Spur is a Permian fossil locality located at the Dolese Brothers Limestone Quarry north of Lawton, Oklahoma. The locality preserves clay and mudstone fissure fills of a karst system eroded out of Ordovician limestone and dolomite, with the infilling dating to the Artinskian stage of the early Permian (Cisuralian), around 289 to 286 million years ago. Fossils of terrestrial animals are abundant and well-preserved, representing one of the most diverse Paleozoic tetrapod communities known. A common historical name for the site is Fort Sill, in reference to the nearby military base. Fossils were first reported at the quarry by workers in 1932, spurring a wave of collecting by local and international geologists. Early taxa of interest included the abundant reptile Captorhinus and microsaurs such as Cardiocephalus and Euryodus. Later notable discoveries include Doleserpeton, the most diverse assortment of parareptiles in the Early Permian, and the rare early diapsid Orovenator.