Australothyris

Last updated

Australothyris
Temporal range: Middle Permian, 265–260  Ma
O
S
D
C
P
T
J
K
Pg
N
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Parareptilia
Order: Procolophonomorpha
Genus: Australothyris
Modesto et al., 2009 [1]
Type species
Australothyris smithi
Modesto et al., 2009

Australothyris is an extinct genus of basal procolophonomorph parareptile known from the Middle Permian (middle Capitanian stage) 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. [1]

Contents

Discovery

Australothyris is known from a single specimen discovered at the Beukesplaas farm by Robert Smith in 1995. The fossil site at the Beukesplaas farm contains a diverse parareptile and synapsid fauna positioned in the Middle Permian Tapinocephalus Assemblage zone of the upper Abrahamskraal Formation. This specimen, SAM-PK-K8302, included most of a skull and portions of the rest of the skeleton, which had mostly been eroded away prior to its discovery. It was initially referred to Owenetta based on the numerous teeth and long postfrontal, until a reexamination revealed a temporal fenestra, which was absent in owenettids. In the wake of this restudy, the specimen was recognized as a new taxon, which was named Australohyris smithi by Sean P. Modesto, Diane M. Scott, and Robert R. Reisz in 2009. The generic name translates to "southern opening" in recognition that it supports the hypothesis that parareptiles originated in Gondwana and went through a phase of evolution where they possessed a temporal fenestra, an opening in the skull behind the eyes. The specific name honors Robert Smith. [1]

Description

Portions of the snout and upper skull have been weathered away, but many notable features are preserved. The maxilla contains 31 teeth, an unusually high number which is only surpassed by Microleter [2] and Lanthanosuchus among parareptiles. The teeth are small, slender, and conical, retaining roughly the same size and shape except for a subtle decrease in size towards the rear of the maxilla. The prefrontal is simple, hosting a small buttress in front of the orbits (eye holes) and being dissimilar in shape to that of procolophonids. Although the frontal does contact the upper edge of the orbit as in other amniotes, it lacks the distinct lappet observed in lanthanosuchoids. A distinct lateral temporal fenestra is present behind the orbit, completely surrounded by the jugal, quadratojugal, postorbital and squamosal. Other parareptiles with lateral temporal fenestrae (apart from lanthanosuchids) typically exclude the postorbital from its edge through contact between the jugal and squamosal, or have an open lower edge due to a loss of contact between the jugal and quadratojugal. The postfrontal is uniquely elongated, as its rear branch contacts the boxy supratemporal bone and separates the postorbital from the parietal. The pineal foramen is large, similar in size to that of procolophonids and bolosaurids. The quadrates are massive, being quite broad but also not very tall as in Acleistorhinus . Minor ornamentation is present on several bones, including broad grooves (on the nasal), shallow pits (on the jugal), clusters of knobs and furrows (on the postorbital), and low mounds (on the squamosal). [1]

The vomers possess an array of ridges, the largest being at the edge of the choanae. There are also teeth in small rows or solitary positions on the vomers. The palatines are characteristically large, possessing several low ridges covered with small teeth. The pterygoids were notably broad, owing to extensive contact with each other along the midline of the palate. As a consequence, the interpterygoid vacuity (gap between the pterygoids) is short, restricted to a triangular opening in front of the parabasisphenoid. Tooth rows occur along the inner edge of the pterygoids, on the main underside of the bones, and at the transverse flanges at their rear. The branches of the pterygoids leading to the quadrates are offset from the transverse flanges by a distinct notch. Overall the palate most closely resembles that of Lanthanosuchus. Uniquely, Australothyris even possesses patches of teeth on the basipterygoid processes of the parabasisphenoid. [1]

The rest of the braincase was fairly typical. The basioccipital was broad, with poorly-developed basitubera and kidney-shaped occipital condyle, and the exoccipitals do not meet at the midline of the foramen magnum. The supraoccipital was also broad and fused with the opisthotics in its lower portion, while the upper portion of the bone was overlapped by the small postparietal and tabular bones of the skull roof. The opisthotics were thickest at the base and generally similar to those of Milleretta . They each connected to a thick yet complex stapes which possessed a conspicuous footplate, stapedial foramen, and a dorsal process. A knob on the outer edge of the stapes likely connected to a characteristic spur on the quadrate. What can be seen of the lower jaw indicates that it was primarily formed by the dentary in its front half, and the low, elongated surangular and angular in its rear half. The coronoid had a low peak and the tall articular had a small retroarticular process. Only one tooth was exposed, and it was similar to those of the maxilla, albeit smaller. [1]

The articulated postcranial skeleton is weathered to the point that only portions of the cervical vertebrae and interclavicle are in good enough condition to describe. The cervicals had slight excavations on their outer surface, robust neural arches, and low neural spines, with that of the axis overhanging its predecessor. The interclavicle is anchor-shaped (like ankyramorph, or "anchor-form" parareptiles), but in contrast to ankyramorphs, the center of the interclavicle is thicker than the front edge. Overall the postcranium is congruent with that known for Milleretta. [1]

Classification

The original describers of Australothyris used a phylogenetic analysis designed by Muller & Tsuji (2007) to investigate its relations to other parareptiles. The analysis found that it had an optimal position as a relatively basal parareptile, specifically the sister taxon to Ankyramorpha (the group containing lanthanosuchoids and more derived parareptiles). This was nevertheless more derived than mesosaurs and millerettids, and the paper's authors assigned the name Procolophonomorpha to parareptiles more derived than millerettids. Australothyris was recovered as the first branch of Procolophonomorpha, suggesting that the group as a whole originated simultaneously with the evolution of a large, fully enclosed temporal fenestra in parareptiles. [1] However, the subsequent discovery of Microleter , which had a roughly equivalent phylogenetic position and a much more restricted temporal emargination, casts doubts on this hypothesis for the origin of temporal fenestration. Certain millerettids have also been observed to possess temporal fenestrae. [2] The position of Australothyris also supports another hypothesis which argues that procolophonomorphs evolved in Gondwana (southern Pangea) before spreading to and diversifying in more northern regions, [1] although Microleter, known from Oklahoma, once more casts doubt on this hypothesis. [2]

Cladogram after Modesto, Scott, & Reisz (2009). [1]

Parareptilia

Related Research Articles

The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.

<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.

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

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.

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

Askeptosaurus is an extinct genus of askeptosauroid, a marine reptile from the extinct order Thalattosauria. Askeptosaurus is known from several well-preserved fossils found in Middle Triassic marine strata in what is now Italy and Switzerland.

<i>Mesenosaurus</i> Extinct genus of synapsids

Mesenosaurus is an extinct genus of synapsid belonging 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.

<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>Polonosuchus</i> Extinct genus of reptiles

Polonosuchus is a genus of rauisuchid known from the late Triassic of Poland. It was a huge predator about 5–6 metres in length and, like all rauisuchians, was equipped with a large head of long sharp teeth. The legs were placed almost underneath the body, unlike most reptiles, which would have made it quite fast and a powerful runner. The appearance was very similar to that of the more known Postosuchus, of North America, and shared with the latter the ecological niche of the apex predator.

<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.

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

Jesairosaurus is an extinct genus of early archosauromorph reptile known from the Illizi Province of Algeria. It is known from a single species, Jesairosaurus lehmani. Although a potential relative of the long-necked tanystropheids, this lightly-built reptile could instead be characterized by its relatively short neck as well as various skull features.

Leninia is an extinct genus of basal ophthalmosaurine ichthyosaur known from the late Early Cretaceous of western Russia. Leninia was first named by Valentin Fischer, Maxim S. Arkhangelsky, Gleb N. Uspensky, Ilya M. Stenshin and Pascal Godefroit in 2013 and the type species is Leninia stellans. It was named for Vladimir Lenin, one of the leaders of the Communist Revolution in Russia, but not directlу: the museum where fossils is housed is located within the Lenin Memorial and Lenin school complex in Ulyanovsk; accordingly, the generic name reflects the geohistorical location of the find.

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

Palatodonta is an extinct genus of neodiapsid reptile known from the early Middle Triassic of the Netherlands. It was initially described in 2013 as a basal placodontiform closely related to a group of marine reptiles called placodonts, characterized by their crushing teeth and shell-like body armor. Under this interpretation, Palatodonta is transitional between placodonts and less specialized reptiles. Like placodonts, it has a row of large teeth on its palate, but while these teeth are thick and blunt in placodonts, Palatodonta has palatal teeth that are thin and pointed. A 2023 study instead classified it as a sauropterygomorph and the sister taxon to Eusaurosphargis. In other words, it is close to, but not within, Sauropterygia.

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.

<i>Bulbasaurus</i> Extinct genus of dicynodonts

Bulbasaurus is an extinct genus of dicynodont that is known from the Lopingian epoch of the Late Permian period of what is now South Africa, containing the type and only species B. phylloxyron. It was formerly considered as belonging to Tropidostoma; however, due to numerous differences from Tropidostoma in terms of skull morphology and size, it has been reclassified the earliest known member of the family Geikiidae, and the only member of the group known from the Tropidostoma Assemblage Zone. Within the Geikiidae, it has been placed close to Aulacephalodon, although a more basal position is not implausible.

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

Avicranium is a genus of extinct drepanosaur reptile known from the Chinle Formation of the late Triassic. The type species of Avicranium is Avicranium renestoi. "Avicranium" is Latin for "bird cranium", in reference to its unusual bird-like skull, while "renestoi" references Silvio Renesto, a paleontologist known for studies of Italian drepanosaurs.

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

Vadasaurus is an extinct genus of rhynchocephalian closely related to the aquatic pleurosaurids. Although this genus was not as specialized as the eel-like pleurosaurs for aquatic life, various skeletal features support the idea that it had a semiaquatic lifestyle. The type species, Vadasaurus herzogi, was described and named in 2017. It was discovered in the Solnhofen Limestone in Germany, which is dated to the Late Jurassic. The generic name "Vadasaurus" is derived from "vadare", which is Latin for "to go" or "to walk forth", and "saurus", which means "lizard". "Vadare" is the root of the English word "wade", which is the reason it was chosen for this genus, in reference to its perceived semiaquatic habits. The specific name, "herzogi", refers to Werner Herzog, a Bavarian filmmaker.

<i>Kadimakara australiensis</i> Extinct species of reptile

Kadimakara is an extinct genus of early archosauromorph reptile from the Arcadia Formation of Queensland, Australia. It was seemingly a very close relative of Prolacerta, a carnivorous reptile which possessed a moderately long neck. The generic name Kadimakara references prehistoric creatures from Aboriginal myths which may have been inspired by ice-age megafauna. The specific name K. australiensis relates to the fact that it was found in Australia. Prolacerta and Kadimakara were closely related to the Archosauriformes, a successful group which includes archosaurs such as crocodilians, pterosaurs, and dinosaurs.

Fraxinisaura is an extinct genus of basal lepidosauromorph reptile known from the Middle Triassic of Germany. The only known species is Fraxinisaura rozynekae. It possessed an elongated snout, unique features of the teeth, and an ilium which was intermediate in orientation between sphenodontians and squamates. Based on characteristics of the maxilla, it is considered a close relative of Marmoretta from the Middle Jurassic of the United Kingdom, resolving a ghost lineage between that genus and other Triassic basal lepidosauromorphs.

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

Polymorphodon is an extinct genus of archosauriform reptile from the Middle Triassic of Germany. The only known species is Polymorphodon adorfi, discovered in Lower Keuper deposits at a quarry in Eschenau, Germany. Polymorphodon is notable for its heterodont dentition, with long and conical premaxillary teeth followed by thin maxillary teeth with large serrations. Maxillary teeth near the back of the mouth are short and leaf-shaped, similar to some living and extinct reptiles with a herbivorous or omnivorous diet. This may suggest that Polymorphodon had some reliance on plants in its diet, a rarity among basal archosauriforms, most of which are carnivores.

<i>Vetusodon</i> Extinct genus of cynodonts

Vetusodon is an extinct genus of cynodonts belonging to the clade Epicynodontia. It contains one species, Vetusodon elikhulu, which is known from four specimens found in the Late Permian Daptocephalus Assemblage Zone of South Africa. With a skull length of about 18 centimetres (7.1 in), Vetusodon is the largest known cynodont from the Permian. Through convergent evolution, it possessed several unusual features reminiscent of the contemporary therocephalian Moschorhinus, including broad, robust jaws, large incisors and canines, and small, single-cusped postcanine teeth.

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

Taytalura is an extinct genus of lepidosauromorph reptile from the Late Triassic of Argentina. It contains a single species, Taytalura alcoberi, which is based on a well-preserved skull from the fossiliferous Ischigualasto Formation. As a lepidosauromorph, Taytalura is a distant relative of modern lepidosaurs such as sphenodontians and squamates. Taytalura did not belong to any group of modern lepidosaurs, since it bears unique features, such as unfused bones in the skull roof and teeth which all sit loosely in a deep groove without sockets. Regardless, Micro-CT scanning reveals features of the skull previously only seen in rhynchocephalians. This suggests that the ancestral condition of the skull in lepidosaurs was more similar to sphenodonts than to squamates.

References

  1. 1 2 3 4 5 6 7 8 9 10 Sean P. Modesto; Diane M. Scott & Robert R. Reisz (2009). "A new parareptile with temporal fenestration from the Middle Permian of South Africa". Canadian Journal of Earth Sciences. 46 (1): 9–20. Bibcode:2009CaJES..46....9M. doi:10.1139/E09-001.
  2. 1 2 3 Linda A. Tsuji; Johannes Muller; Robert R. Reisz (2010). "Microleter mckinzieorum gen. et sp. nov. from the Lower Permian of Oklahoma: the basalmost parareptile from Laurasia". Journal of Systematic Palaeontology. 8 (2): 245–255. doi:10.1080/14772010903461099. S2CID   129529082.