Microleter Temporal range: Early Permian, | |
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Skull diagram | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | † Parareptilia |
Order: | † Procolophonomorpha |
Node: | † Ankyramorpha |
Genus: | † Microleter Tsuji et al., 2010 |
Type species | |
†Microleter mckinzieorum Tsuji et al., 2010 |
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. [1]
The only known specimen of Microleter is a well-preserved skull and lower jaw designated as OMNH 71306, the holotype specimen. It was found at the Dolese Brothers limestone quarry near Richards Spur in Comanche County, Oklahoma. [1] Historically referred to as the Fort Sill locality, the quarry has preserved Early Permian (Artinskian, ~289-286 Ma) fissure fills in an Ordovician cave system. The fissure fills have yielded many other well preserved tetrapod fossils, including the most diverse assortment of Permian parareptiles in North America. [2]
Microleter mckinzieorum was named by paleontologists Linda A. Tsuji, Johannes Muller, and Robert R. Reisz in 2010. The genus name combined Greek "mikros" (small) with "-oleter" (murderer), a suffix common to parareptile genera. The specific name honors the McKinzie family, as the specimen was discovered by Mark McKinzie. [1]
Based on the skull's large orbits (eye holes) and weak sutures, the specimen was likely a juvenile. Most of the skull bones were externally textured by radiating pits and furrows, with both sparse large pits and numerous tiny pits as in basal lanthanosuchoids. The only smoothly textured bones of the skull roof were the maxilla, squamosal, and quadratojugal. The maxilla was long and narrow, possessing conical teeth which only differed from each other in a slight shortening trend towards the rear of the maxilla. The maxillary tooth count was 32 or 33, more than any other parareptile apart from Lanthanosuchus . [1] Like several other basal parareptiles, Microleter had teeth with plicidentine, a type of internally folded dentine which is most common in "labyrinthodont" amphibians. [3] Although the lacrimal was not complete, the internal texture of the overlapping maxilla indicates that it extended to the nares (nostril holes), a trait also observed in millerettids, pareiasaurs, and bolosaurids. [1]
The front edge of the orbit was thick due to an internal flange on the prefrontal, akin to that of procolophonids and Colobomycter . The upper edge of the orbit had a slight contribution by the frontal bone, though more restricted than that of lanthanosuchoids and procolophonoids. The jugal and particularly the postfrontal are both small and crescent-shaped. The postorbital, by contrast, is larger and extends to near the rear edge of the skull roof. Along with the jugal, squamosal, and quadratojugal, the postorbital edges a tall, narrow opening in the rear portion of the skull. This opening, termed a ventral temporal emargination, is likely homologous to the temporal opening present in various other parareptiles. [4] The slit-like opening of Microleter is proportionally similar to that of Nyctiphruretus , although differs in the participation of the postorbital in its border. However, other parareptiles with postorbital participation ( Australothyris and lanthanosuchids) have their opening fully surrounded by bone, while that of Microleter is open from below. The rear of the skull is not very long, with a broad parietal, small supratemporal, solitary postparietal, and tall, boxy squamosal and quadratojugal which are excavated along their rear edge. [1]
Most of the palate is obscured by overlapping bones. The palatine is very broad, while the transverse flange of the pterygoid is oriented forwards. What can be seen of the braincase indicates that Microleter had tubular paroccipital processes, rather than fan-shaped ones present in other parareptiles. The elongated dentary is ornamented with small pits, but its teeth are obscured and cannot be properly counted. The surangular possesses a folded ridge on its outer surface, and encompasses the front half of a hole at the rear of the jaw. The rear half of the hole is edged by the articular bone. Microleter is one of the few parareptiles to have preserved part of the sclerotic ring, which was formed by tall, concave plates. A few cervical vertebrae are the only fossilized postcranial elements, but they are poorly preserved. [1]
Microleter is one of the most basal parareptiles, and was originally described as the most basal parareptile from Laurasia. Before the description of Microleter, parareptiles were hypothesized to have originated in Gondwana. However, Microleter appears in Laurasia soon after the earliest known parareptiles, and spurred discussion supporting the origin of parareptiles in Laurasia. Other factors in support of a Laurasian origin are the early appearance of the Laurasian bolosaurians, [5] the fact that the other two clades of amniotes, synapsids and eureptiles, are both considered to have been Laurasian in origin, and evidence from varanopids that colonization of Gondwana by Laurasian amniotes was more common than the reverse process. [1]
The original description of Microleter used maximum parsimony and bayesian phylogenetic analyses to position it more derived than millerettids and less derived than lanthanosuchoids. In the parsimony analysis, it was in a polytomy with Australothyris and more derived taxa, which was resolved in the bayesian analysis to place Australothyris more basally. [1]
Cladogram from the bayesian analysis of Tsuji, Muller, & Reisz (2010): [1]
However, since the original description, various other analyses have each had slightly different conclusions on the position of Microleter, placing it as the sister taxon of Australothyris, [6] basal to Australothyris, [7] or as a basal ankyramorph more derived than lanthanosuchoids, [8] [5] [9] [10] [11]
The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.
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.
Cacops, is a genus of dissorophid temnospondyls from the Kungurian stage of the early Permian of the United States. Cacops is one of the few olsoniforms whose ontogeny is known. Cacops fossils were almost exclusively known from the Cacops Bone Bed of the Lower Permian Arroyo Formation of Texas for much of the 20th century. New material collected from the Dolese Brothers Quarry, near Richards Spur, Oklahoma in the past few decades has been recovered, painting a clearer picture of what the animal looked and acted like.
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.
Elginia is an extinct genus of pareiasaurid known from the Late Permian of Scotland and China. It was named for the area around Elgin in Scotland, which has yielded many fossils referred to as the Elgin Reptiles.
Procolophonidae is an extinct family of small, lizard-like parareptiles known from the Late Permian to Late Triassic that were distributed across Pangaea, having been reported from Europe, North America, China, South Africa, South America, Antarctica and Australia. The most primitive procolophonids were likely insectiovous or omnivorous, more derived members of the clade developed bicusped molars, and were likely herbivorous feeding on high fiber vegetation or durophagous omnivores. Many members of the group are noted for spines projecting from the quadratojugal bone of the skull, which likely served a defensive purpose as well as possibly also for display. At least some taxa were likely fossorial burrowers. While diverse during the Early and Middle Triassic, they had very low diversity during the Late Triassic, and were extinct by the beginning of the Jurassic.
Nyctiphruretus is an extinct genus of nyctiphruretid parareptile known from the Guadalupian series of European Russia.
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.
Colobomycter is an extinct genus of lanthanosuchoid parareptile known from the Early Permian of Oklahoma.
Hapsidopareion is an extinct genus of microsaur belonging to the family Hapsidopareiidae. Fossils have been found in the early Permian of Oklahoma.
Llistrofus is an extinct genus of early Permian microsaur within the family Hapsidopareiidae that is known from 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.
Mesenosaurus is an extinct genus of amniote. It belongs to the family Varanopidae. This genus includes two species: the type species Mesenosaurus romeri from the middle Permian Mezen River Basin of northern Russia, and Mesenosaurus efremovi from the early Permian (Artinskian) Richards Spur locality. M. romeri’s stratigraphic range is the middle to late Guadalupian while M. efremovi’s stratigraphic range is the Cisuralian.
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.
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.
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.
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.
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 nyctiphruretid 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.