Acleistorhinus Temporal range: Early Permian, | |
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Illustration of Acleistorhinus skull | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | † Parareptilia |
Order: | † Procolophonomorpha |
Family: | † Acleistorhinidae |
Genus: | † Acleistorhinus Daly, 1969 |
Type species | |
†Acleistorhinus pteroticus Daly, 1969 |
Acleistorhinus (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian (middle Kungurian stage) of Oklahoma. [1] 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 (also from the Early Permian of Oklahoma).
Acleistorhinus was first discovered and named by Eleanor Daly in 1969 in the Hennessey Formation of South Grandfield, Tillman county, Oklahoma. The name Acleistorhinus combines Greek rhin (ῥῑ́ν), meaning "nose," and akleistos, Greek for “unclosed.“
Although its total body length is unknown, an Acleistorhinus skull is about 3.5 centimetres long. [2] From the dorsal side, the Acleistorhinus skull appears to have a triangular outline. The surface of the skull is generally smooth with a few small, shallow circular pits. Anteriorly, the snout is gently rounded. [3] The elliptical external nares are each bordered by the maxilla. The tooth bearing portion of the premaxilla appears to be directed somewhat downward at its tip. Each premaxilla possess spaces for four teeth. The maxilla has a dorsal expansion immediately behind the nares forming the entire posterier border of the opening. This configuration resembles that of the procolophonids, and turtles, and results in the exclusion of the lacrimal from the posterior border of the nares. Slightly more than one third of the total length of the skull is contributed by the frontal. It is constricted anteriorly by the prefrontals, but otherwise expanded above the orbits. [4] Generally, in early amniotes the largest element was the occiput of the supraoccipital. In Acleistorhinus the supraoccipital is rather plate-like. The reduction in the overall size of the supraoccipital allows for the development of large post-termporal fenestra, a characteristic of Reptilia. [5]
The marginal dentition is composed of conical teeth that are slightly recurved. [6] No canine region is evident although the second maxillary tooth is slightly larger than the rest. The tooth-bearing portion of the maxilla extends posterior to the orbit. All the premaxillary teeth appear to be approximately the same size, and noticeably smaller than those on the maxilla. The maxillae have 11 and 13 on the right and left sides respectively, there is room for at least 17 teeth for each element. Smaller teeth are present along the sloping surface of the transverse flange, anterior to the large row of teeth. On the parasphenoid plate, two separate paired rows of teeth diverge posteriorly. The lateral-most rows sit on a ridge that runs the length of the main body of the parasphenoid. The tooth ridges are evident anteriorly but appear to terminate at the same level as the teeth on the transverse flange of the pterygoid.
The Early Permian is marked by terrestrial plant diversification, in which insects evolved rapidly as they followed the plants into new habitats. Acleistorhinus is widely believed to be an insectivore because its teeth are numerous, small and pointed. The back of the skull is wide resulting in the orbits being pushed forward. This would have offered a degree of binocular vision giving Acleistorhinus, a land-dwelling insectivore, depth perception necessary for hunting fast moving objects.
The genus Acleistorhinus belongs to the taxon parareptilia along with Millerettidae, Lanthanosuchidae to whom it is a sister taxa, Macroleter and Procolophonia. As of present, there is only one known species of Acleistorhinus, known as Acleistorhinus pteroticus.
A recent restudy, phylogenetic analysis, of Acleistorhinus indicates that this Early Permian amniote from North America, the oldest known member of parareptilia, is a sister taxon to Russian Lanchanosuchidae. [7] In addition, the results support Laurin and Reisz (1995) hypothesis that Parareptilia is a monophyletic group, while differing in the number of synapomorphies diagnosing the clade. [8] The recognition of Acleistorhinus and lanthanosuchids as sister-taxa presents new evidence for the hypothesis that parareptiles had a cosmopolitan distribution during the Paleozoic. This sister-group relationship is supported by twelve synapomorphies. Furthermore, when acknowledging Acleistorhinus, lanthanosuchids, and Macroleter nest within Parareptilia, it becomes evident through specific interrelationships within Amniota that Parareptilia is a monophyletic taxon.
Using the tenet of minimum divergence time most recently discussed by Norell (1992) and Westphalian (1993), the earliest parareptile must extend into at least the Westphalian (stage) of the Upper Carboniferous. This suggests that the all major amniote clades Diapsida, Synapsida and Parareptilia all diverged early in the evolutionary radiation of amniotes. At the very least parareptiles are more diverse and possess a richer fossil record than previously recognized.
Parareptilia |
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Acleistorhinus was discovered by Daly in 1969, in the Early Permian outcrops of the Hennessey Formation, the locality of South Grandfield of southwestern Oklahoma. [9] The Hennessey Formation is believed to be contemporaneous with the Richards Spur locality near Fort Sill, Oklahoma, as they both possess a mixed fauna, which is generally disarticulated and incomplete. In addition, over 200 skulls and 500 specimens have been collected from South Grandfield, only one specimen of Acleistorhinus is known. [10] [11] It is very likely that this taxon is an erratic and would not normally preserve in the depositional environment that characterizes much of the Lower Permian of North America.
The discovery of Acleistorhinus was far reaching because until the present only Synapida and Diapsida could trace their earliest known members to North America. Now parareptiles can also trace their earliest record from the same continent. [12] It is very likely that all three major amniote clades Diapsida, Synapsida, and Parareptilia all diverged early during the evolutionary radiation that characterizes much of the Early Permian.
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.
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.
Milleretta is an extinct genus of millerettid parareptile from the Late Permian of South Africa. Fossils have been found in the Balfour Formation. Milleretta was a moderately sized, lizard-like animal, about 60 centimetres (24 in) in length. It was probably insectivorous. Its only known species is Milleretta rubidgei, making Milleretta a monospecific genus.
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.
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.
Macroleter is an extinct genus of nycteroleterid parareptile which existed in Oklahoma and Russia during the upper Permian period. It was a quite generalized primitive reptile, in many ways resembling their amphibian ancestors. It was first named by paleontologists Tverdochlebova and Ivachnenko in 1984. According to classification by Michel Laurin and Robert R. Reisz, the genus is a parareptile, belonging to the same branch as Millerettidae, Procolophonidae and other generalized anapsid reptiles. The type species is Macroleter poezicus from Upper Permian of Russia.
Leptopleuron is an extinct genus of procolophonid that lived in the dry lands during the late Triassic in Elgin of northern Scotland and was the first to be included in the clade of Procolophonidae. First described by English paleontologist and biologist Sir Richard Owen, Leptopleuron is derived from two Greek bases, leptos for "slender" and pleuron for "rib," describing it as having slender ribs. The fossil is also known by a second name, Telerpeton, which is derived from the Greek bases tele for "far off" and herpeton for "reptile." In Scotland, Leptopleuron was found specifically in the Lossiemouth Sandstone Formation. The yellow sandstone it was located in was poorly lithified with wind coming from the southwest. The environment is also described to consist of barchan dunes due to the winds, ranging up to 20 m tall that spread during dry phases into flood plains. Procolophonoids such as Leptopleuron were considered an essential addition to the terrestrial ecosystem during the Triassic.
Labidosaurikos is a genus of extinct captorhinid anapsid reptile that lived around 279 to 272 million years ago during Kungurian age of the lower Permian. The American paleontologist John Willis Stovall first described Labidosaurikos in 1950, naming it "Labidosaurus like" for the striking similarity of the holotype skull of his specimen to the cranial anatomy of another captorhinid Labidosaurus hamatus. Labidosaurus or generally called "lipped lizard" is another genus of the family Captorhinidae whose name is derived from the Greek for "forceps lizard" based on τσιμπίδα and σαυρος ("lizard")
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.
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.
Australothyris is an extinct genus of basal procolophonomorph parareptile known from the Middle Permian of Tapinocephalus Assemblage Zone, South Africa. The type and only known species is Australothyris smithi. As the most basal member of Procolophonomorpha, Australothyris helped to contextualize the origin of this major parareptile subgroup. It has been used to support the hypotheses that procolophonomorphs originated in Gondwana and ancestrally possess temporal fenestrae, due to its large and fully enclosed temporal fenestra and South African heritage. It also possessed several unique features, including a high tooth number, long postfrontal, small interpterygoid vacuity, and a specialized interaction between the stapes and quadrate.
Ankyramorpha is an extinct clade of procolophonomorph parareptiles which lived between the early Cisuralian epoch and 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.
Feeserpeton is an extinct genus of parareptile from the Early Permian of Richard's Spur, Oklahoma. It is known from a single species, Feeserpeton oklahomensis, which was named in 2012 on the basis of a nearly complete skull. Feeserpeton is a member of the clade Lanthanosuchoidea and is one of the earliest parareptiles.
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.