Acleistorhinus

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Acleistorhinus
Temporal range: Early Permian, 273.6–271.6  Ma
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Acleistorhinus four images.png
Illustration of Acleistorhinus skull
Scientific classification OOjs UI icon edit-ltr.svg
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).

Contents

Etymology

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.“

Description and Paleobiology

Skull

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 posterior 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]

Dentition

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.

Habitat and Diet

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.

Classification and Species

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

Discovery

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.

See also

Related Research Articles

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<span class="mw-page-title-main">Parareptilia</span> Extinct subclass of reptiles (306–201Ma ago)

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.

<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)

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<i>Macroleter</i> Extinct genus of 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.

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

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

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<i>Labidosaurikos</i> Extinct genus of tetrapods

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

Colobomycter is an extinct genus of acleistorhinid parareptile known from the Early Permian of Oklahoma.

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

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.

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

Acleistorhinidae is an extinct family of Late Carboniferous and Early Permian-aged parareptiles. It is defined as a node based clade including the last common ancestor of Acleistorhinus pteroticus and Colobomycter pholeter and all its descendants. Acleistorhinids are most diverse from the Richards Spur locality of the Early Permian of Oklahoma. Richards Spur acleistorhinids include Acleistorhinus, Colobomycter, Delorhynchus, Feeserpeton and Klastomycter. 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.

<i>Heleosaurus</i> Extinct genus of tetrapods

Heleosaurus scholtzi is an extinct species of basal synapsids, known as pelycosaurs, in the family of Varanopidae during the middle Permian. At first H. scholtzi was mistakenly classified as a diapsid. Members of this family were carnivorous and had dermal armor, and somewhat resembled monitor lizards. This family was the most geologically long lived, widespread, and diverse group of early amniotes. To date only two fossils have been found in the rocks of South Africa. One of these fossils is an aggregation of five individuals.

<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.

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

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.

Plemmyradytes is an extinct genus of dissorophoid temnospondyl from the early Permian. It is an amphibamiform from the Eskridge Formation exposures of Nebraska. The type species is Plemmyradytes shintoni. The genus name derives from the Greek plemmyris and dytes ('diver'), while the specific name honors John Shinton, a fossil preparator at the Denver Museum of Natural History where all known specimens of this taxon are reposited following collection in the late 20th century.

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

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.

<span class="mw-page-title-main">Lanthanosuchoidea</span> Extinct superfamily of reptiles

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.

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

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.

Huskerpeton is an extinct genus of recumbirostran from the Early Permian period. They belong to the order Microsauria, which was established in 1863 by Dawson, and was quickly expanded to include many different small taxa. They lived in what is now Nebraska and Kansas. The holotype of Huskerpeton was uncovered at the Eskridge formation in Nebraska, which is part of how it got its name.

References

  1. Daly, E. (1969). "A new Procolophonomorpha Procolophonoid reptile from the Early Permian of Oklahoma". Journal of Paleontology. 43 (3): 676–687.
  2. Reisz, R.R.; Macdougall, M.J.; Modesto, S. (2014). "A new species of the parareptile genus Delorhynchus, based on articulated skeletal remains from the Richards Spur, Lower Permian of Oklahoma". Journal of Vertebrate Paleontology. 34 (5): 1033–1043. doi:10.1080/02724634.2013.829844. S2CID   128459194.
  3. MacDougall, M.J.; Modesto, S.P.; Reisz, R.R. (2016). "A new reptile from the Richards Spur Locality, Oklahoma, USA, and patterns of Early Permian parareptile diversification". Journal of Vertebrate Paleontology. 36 (5): e1179641. doi:10.1080/02724634.2016.1179641. S2CID   89333948.
  4. Reisz, R.R.; Mueller, J.; Tsuji, L.; Scott, D. (2007). "The cranial osteology of Belebey vegrandis (Parareptilia: Bolosauridae), from the Middle Permian of Russia, and its bearing on reptilian evolution". Zoological Journal of the Linnean Society. 151 (1): 191–214. doi: 10.1111/j.1096-3642.2007.00312.x .
  5. Modesto, S.P. (1999). "Colobomycter pholeter from the Lower Permian of Oklahoma: a parareptile, not a protorothyridid". Journal of Vertebrate Paleontology. 19 (3): 466–472. doi:10.1080/02724634.1999.10011159.
  6. Anderson, J.M.; Cruickshank, A.R.I. (1978). "The biostratigraphy of the Permian and the Triassic Part 5. A review of the classification and distribution of Permo-Triassic tetrapods". Palaeontologia Africana. 21: 15–44.
  7. Case, E. C. (1911). "A Revision of the Cotylosauria of North America". Carnegie Institution of Washington. Publication. 145: 1–122.
  8. Laurin, M.; R. R. Reisz (1995). "A reevaluation of early amniote phylogeny". Biological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x. S2CID   29661691.
  9. Tsuji, L.A. (2006). "Cranial anatomy and phylogenetic affinities of the Permian parareptile Macroleter poezicus". Journal of Vertebrate Paleontology. 26 (4): 849–865. doi:10.1671/0272-4634(2006)26[849:CAAPAO]2.0.CO;2. S2CID   86266685.
  10. Gow, C.E. (1973). "The osteology and relationships of the Millerettidae (Reptilia: Cotylosauria)". Journal of Zoology. 167 (2): 219–264. doi:10.1111/j.1469-7998.1972.tb01731.x.
  11. Robert R. Reisz; Mark J. Macdougall; Sean P. Modesto (2014). "A new species of the parareptile genus Delorhynchus, based on articulated skeletal remains from Richards Spur, Lower Permian of Oklahoma". Journal of Vertebrate Paleontology. 34 (5): 1033–1043. doi:10.1080/02724634.2013.829844. S2CID   128459194.
  12. Fox, R. (1962). "Two new pelycosaurs from the lower Permian of Oklahoma". University of Kansas Publications, Museum of Natural History. 12 (6): 297–307.

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