Pentasaurus

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Pentasaurus
Temporal range: Late Triassic
Norian
Pentasaurus holotype.png
The holotype mandible of Pentasaurus goggai (NHMW 1876-VII-B-114)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Suborder: Anomodontia
Clade: Dicynodontia
Family: Stahleckeriidae
Subfamily: Placeriinae
Genus: Pentasaurus
Kammerer, 2018
Species:
P. goggai
Binomial name
Pentasaurus goggai
Kammerer, 2018

Pentasaurus is an extinct genus of dicynodont of the family Stahleckeriidae, closely related to the well known Placerias . It was found in the Lower Elliot Formation of South Africa, dated to the Norian of the Late Triassic period. The genus contains the type and only species, Pentasaurus goggai. Pentasaurus is named after the ichnogenus Pentasauropus , fossil footprints that were originally described from the lower Elliot Formation in 1970 decades before the body fossils of Pentasaurus itself were recognised. Pentasauropus footprints were likely made by dicynodonts, and in South Africa Pentasaurus itself was the likely trackmaker. The name reflects the fact that a large dicynodont was predicted to have existed in the lower Elliot Formation before any body fossils were recognised, and so Pentasaurus was named after its probable footprints. This is a reversal of the more typical occurrence where fossil footprints are named after their presumed trackmakers. The name of the species honours its collector Alfred Brown, nicknamed "Gogga", which means "bug" in Afrikaans.

Contents

Pentasaurus is also unique for being one of the only known dicynodonts to have coexisted with large-bodied sauropodomorph dinosaurs. Prior to its discovery, large sauropodomorphs and dicynodonts were thought to have never ecologically coexisted, with dicynodonts either going extinct prior to large sauropodomorphs evolving or being outcompeted and driven to extinction by the herbivorous dinosaurs. Pentasaurus challenges these assumptions, suggesting that dicynodonts and large sauropodomorphs could be contemporaneous, and may instead have been separated through habitat and dietary preferences. Furthermore, Pentasaurus expands the range of stahleckeriid dicynodonts in the Late Triassic into South Africa, refuting ideas that dicynodonts were geographically restricted by the Late Triassic.

Description

The fossil remains of Pentasaurus are fragmented and very incomplete, so its overall morphology is poorly understood. However, it was similarly sized to other large Late Triassic dicynodonts, such as the closely related Placerias, and likely resembled them in appearance.

Skull and mandible

Stereopairs of the skull roof of Pentasaurus viewed from above and below Pentasaurus skullroof.png
Stereopairs of the skull roof of Pentasaurus viewed from above and below

The only known bones from the skull of Pentasaurus is a portion from the roof of the skull, specifically the front end of the intertemporal region consisting of both frontal bones, the left postorbital and the preparietal (a bone unique to dicynodonts and some other therapsids). There are two depressed regions of the skull piece, one along the bottom left side and another cut into the back end, interpreted as representing a site for jaw muscle attachments at the edge of the temporal fenestra and for the pineal foramen, respectively. Otherwise, the skull is notably rough and strongly textured, particularly on a pair of mound-like rugosities at the back edge of the frontals. Frontal rugosities are absent in other kannemeyeriiforms and may be unique to Pentasaurus, although the bones around the eyes of Placerias are similarly rugose. [1] [2]

Only the very front of the mandible of Pentasaurus is known, mostly consisting of the fused mandibular symphysis (including the splenial) and part of the left dentary and angular, although the tip of the beak is missing. The mandible is robust, and like other dicynodonts was fused into a toothless beak. The beak was relatively short in Pentasaurus, with a steep front edge drawn into a mid-line ridge like those of other placeriines, although it is relatively shorter and lacks a pair of grooves typically found on either side of the ridge in other stahleckeriids. The beak is proportionately broader than that of Placerias, and its edges are smoothly rounded and curve towards the tip, unlike the sharp edged, flattened surfaces on the beaks of Placerias or Stahleckeria . A mid-line groove on the top surface of the beak, present in other kannemeyeriiforms, is unusually shallow in Pentasaurus, even more so than the notably shallow groove in Placerias. [1]

The lateral dentary shelf, a characteristic ridge on the jaws of dicynodonts for attaching jaw muscles, is unusual in Pentasaurus. It is remarkably robust and prominent, and the shelf sits uniquely far forward on the jaw compared to all other dicynodonts. Further unlike other dicynodonts, the shelf does not extend back to meet the mandibular fenestra either, instead curving up at its end from an otherwise horizontal position. Features such as the shape of the beak and lateral dentary shelf likely influenced the feeding style of dicynodonts, although the significance of the unusual morphology in Pentasaurus is currently unknown. [1]

Postcranial skeleton

The postcranial skeleton of Pentasaurus is mostly represented by the appendicular skeleton, including parts of the limbs, shoulders and pelvis, with only a single poorly preserved cervical vertebra from the neck representing the axial skeleton. [1]

Pelvic bones of Pentasaurus compared to those of the related Placerias, Zambiasaurus and Jachaleria Pentasaurus hips.png
Pelvic bones of Pentasaurus compared to those of the related Placerias , Zambiasaurus and Jachaleria

The shoulder is only known from the glenoid region of the fused scapulocoracoid and is typical for stahleckeriid dicynodonts, including a relatively large glenoid opening for the shoulder joint directed back and out to the side, indicating a sprawled forelimb posture. The forelimb itself is only known from the end of the humerus and the proximal portion of the ulna. The humerus is largely typical of kannemeyeriiform dicynodonts in shape, however, the distal end is unusually large and thickened compared to other stahleckeriids. The capitulum (part of the elbow) in the only known specimen is unossified and would have been cartilaginous, suggesting the specimen may be immature compared to similarly sized specimens of Placerias. The known portion of ulna is badly worn, but the roundness of its tip suggests that Pentasaurus had a separated olecranon process of the elbow, similar to Placerias and Ischigualastia but unlike some specimens of Stahleckeria. An unidentified long bone may represent a radius. [1]

The pelvis is known from a partial left pubis and ischium and is typical of kannemeyeriiforms, including notable 'twisting' of the pubic shaft. The only known hindlimb element is the proximal end of a right tibia, and likewise it is typical of kannemeyeriiform dicynodonts. Although similar in construction to sauropodomorph dinosaurs, the internal structure visible at the broken end shows extensive trabecular bone, characteristic of Triassic dicynodonts. [1]

History of discovery

The only known fossil remains of Pentasaurus were collected in the late 19th century by amateur palaeontologist Alfred "Gogga" Brown, an Englishman recluse who lived for much of his life in Aliwal North in the Eastern Cape Province, South Africa. His first fossil discoveries began in the 1860s, and his collections largely consisted of tetrapod fossils from the Middle Triassic Beaufort Group and Late Triassic–Early Jurassic Stormberg Group. After initially corresponding with and sending a crate of fossils to the geologist Sir Roderick Murchison in London, Brown's relationships with English and later Parisian scientists soured. However, he later accepted an invitation to have his fossils shipped to the Imperial Natural History Museum, now a part of the Natural History Museum, Vienna in Austria. The fossils of Pentasaurus were part of a lot accessioned in 1876 that was determined to have been collected from a single locality in the Norian aged lower Elliot Formation, based on the shared brown-grey colour of the bones and brown sandstone matrix surrounding them, as well as the association with fossils of the exclusively lower Elliot dinosaur Eucnemesaurus . [1]

Prior to the recognition of Pentasaurus body fossils, dicynodonts were inferred to have been present in the lower Elliot Formation from preserved footprints and track ways attributed to them. These ichnofossils were described in 1970 as Pentasauropus incredibilis, and were determined to have most likely been produced by dicynodonts from their large size and five short toes (pentadactyly). [3] Similar trace fossils were identified on other continents, including South and North America, where they were more confidently attributed to large dicynodonts. [4] However, the absence of dicynodont fossils in some localities known to produce Pentasauropus, as well as their younger age compared to known dicynodont genera, led palaeontologists Adrian Hunt and Spencer Lucas to question this identification. [1] [5]

Re-examination of Brown's collections in the 21st century revealed that a number of bones belonged to a large dicynodont, and in 2018 they were described by palaeontologist Christian Kammerer as a new genus and species, Pentasaurus goggai. The generic name was derived from the ichnogenus Pentasauropus due to the likely association between the tracks and their track maker. Such nomenclatural associations are not uncommon in palaeontology (e.g. Anchisauripus , Megalosauropus ). However, typically the ichnofossil is named after the presumed track maker, whereas this situation is reversed in Pentasaurus. The specific epithet is named in honour and recognition of Alfred Brown and his work, derived from his nickname 'Gogga' (pronounced /ˈxɒxə/ ), an Afrikaan word loosely translated to "bug" that the local villagers nicknamed him for this unusual habits. [1] [6]

Pentasaurus is only known from several isolated fragments of bone, of which a partial mandible (NHMW 1876-VII-B-114) was made the holotype specimen. However, because the bones were collected from one locality and that none of them are duplicated, as well as their similar sizes, it is possible that they all belong to only a single individual. Although disarticulated from each other, the known bones are characteristic of dicynodonts and cannot be referred to other large animals known from the Elliot Formation, such as 'rauisuchians' or sauropodomorph dinosaurs. The inferred association between Pentasaurus and Pentasauropus incredibilis tracks cannot be definitively proven, as there are no known bones from the hands and feet to compare with the shape of the tracks. However, Pentasaurus is the only known candidate dicynodont track-maker in the Elliot Formation, and as other Norian localities are only ever known to have supported one species of dicynodont, it is almost certain that Pentasaurus produced the Pentasauropus tracks originally identified from the Elliot Formation. [1]

Classification

Although fragmentary, the known remains of Pentasaurus are clearly dicynodont and are consistent with belonging to the only known family of Late Triassic dicynodonts, the Stahleckeriidae. Furthermore, a number of features of the lower jaw and humerus support its position in the stahleckeriid subfamily Placeriinae, and so as a close relative of the North American Placerias , Moroccan Moghreberia and the giant Polish Lisowicia . A phylogenetic analysis performed by Kammerer (2018) supported this relationship, however, it could not resolve the relationships within Placeriinae beyond a polytomy containing Pentasaurus and all other placeriines, as shown in the cladogram reproduced below. [1] Later phylogenetic analyses including Pentasaurus produced the same results. [7]

The beak of Pentasaurus compared to those of other stahleckeriids Stahleckeriid mandibles.png
The beak of Pentasaurus compared to those of other stahleckeriids
Stahleckeriidae
Placeriinae

Pentasaurus

Moghreberia

Placerias

Zambiasaurus

Stahleckeriinae

Sangusaurus

Stahleckeria

Eubrachiosaurus

Ischigualastia

Jachaleria

The precise relationships of Pentasaurus to other placeriines cannot be determined cladistically due to the incompleteness of the known remains, nonetheless, it can still be readily distinguished from other placeriines. Particularly, Pentasaurus is characterised by the rounder shape of the beak on its lower jaw, compared to the more angular, straight edged beaks of Placerias and Moghreberia. The robustness and position of the lateral dentary shelf far forward on the lower jaw is also unique to Pentasaurus among dicynodonts, as well as the thickness of its distal humerus. The paired mound-like rugosities over its frontals may also be a diagnostic trait of Pentasaurus. [1]

Palaeoecology

The lower Elliot Formation consists mostly of red-purple mudstones and thick, stacked layers of sandstone deposited by meandering river channels on a floodplain. The relatively narrow width (tens of metres) of the sandstone channels indicate that the river channels were stable and did not migrate across the floodplain, and sedimentary structures preserved within the sandstone indicate that the rivers were slow-moving and perennial. The stability of the channels suggests that riparian forests were present along the banks of these rivers, which is further supported by the relatively high abundance of fossilised wood fragments found in the lower Elliot Formation. These riparian environments were separated by extensive floodplains that were less well vegetated, but likewise they show little evidence for desiccation. Overall, the evidence is consistent with a humid to semi-arid climate in the lower Elliot Formation, and was a wetter environment than the dry, arid upper Elliot Formation of the proceeding Early Jurassic. [8] [9] [10]

The fauna of the lower Elliot Formation is poorly known compared to the upper Elliot Formation, but they show some similarities to other Norian-aged faunas globally. Only two others therapsid are known from the lower Elliot Formation, the large herbivore Scalenodontoides , a traversodontid cynodont and the small trithelodontid cynodont Elliotherium . Archosaurs are better represented, though mostly by several species of herbivorous sauropodomorph dinosaurs. Eucnemesaurus was collected together with Pentasaurus, and so very likely coexisted with it, but Pentasaurus may also have occurred with Blikanasaurus , Sefapanosaurus and Meroktenos , as well as perhaps Melanorosaurus and Plateosauravus depending on their stratigraphic positions. Predatory archosaurs are less well known, but include probable 'rauisuchian' remains, including portions of jaws and teeth that were collected with Pentasaurus. [11] Predatory theropod dinosaurs were also present, but are only known from isolated teeth and footprints. Temnospondyl amphibians are also known from the lower Elliot Formation, but consist of only of chigutisaurids. The footprint record from the lower Elliot Formation records a similar set of fauna to those known from body fossils, although a singular footprint may indicate that crocodylomorphs were also present. [3] [8] [12]

Biogeography

The lower Elliot sauropodomorph Melanorosaurus readi, a possible contemporary of Pentasaurus Melanorosaurus readi steveoc.jpg
The lower Elliot sauropodomorph Melanorosaurus readi, a possible contemporary of Pentasaurus

The coexistence of Pentasaurus with large sauropodomorph dinosaurs including as Eucnemesaurus in the Elliot Formation is significant, as dicynodonts and large sauropodomorphs are otherwise not known to have coexisted anywhere else in world. Prior to the discovery of Pentasaurus, it was even suggested that the two groups of herbivores never even coexisted at all, and that sauropodomorphs replaced dicynodonts as large herbivores in most Late Triassic ecosystems, either through ecological turnover or by direct competition. The coexistence of Pentasaurus and large sauropodomorphs in the lower Elliot Formation suggests that this may not be the case, and that the two groups may have instead been segregated by habitat preference, and may have fed upon different types of vegetation. [1]

Pentasaurus also represents the southernmost example of a placeriine in the world, refuting prior suggestions that placeriine dicynodonts were mostly restricted to the northern hemisphere and demonstrating instead that the group had a more global distribution. [1]

Related Research Articles

<span class="mw-page-title-main">Dicynodont</span> Extinct clade of therapsids

Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.

<i>Melanorosaurus</i> Sauropodomorph dinosaur genus from the Late Triassic period

Melanorosaurus is a genus of basal sauropodomorph dinosaur that lived during the Late Triassic period. A herbivore from South Africa, it had a large body and sturdy limbs, suggesting it moved about on all fours. Its limb bones were massive and heavy like the limb bones of true sauropods.

<i>Massospondylus</i> Sauropodomorph dinosaur genus from Early Jurassic South Africa and Botswana

Massospondylus is a genus of sauropodomorph dinosaur from the Early Jurassic. It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been named. Fossils have since been found at other locations in South Africa, Lesotho, and Zimbabwe. Material from Arizona's Kayenta Formation, India, and Argentina has been assigned to the genus at various times, but the Arizonan and Argentinian material are now assigned to other genera.

<i>Azendohsaurus</i> Genus of herbivorous Triassic reptile

Azendohsaurus is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, Azendohsaurus laaroussii, was described and named by Jean-Michel Dutuit in 1972 based on partial jaw fragments and some teeth from Morocco. A second species from Madagascar, A. madagaskarensis, was first described in 2010 by John J. Flynn and colleagues from a multitude of specimens representing almost the entire skeleton. The generic name "Azendoh lizard" is for the village of Azendoh, a local village near where it was first discovered in the Atlas Mountains. It was a bulky quadruped that unlike other early archosauromorphs had a relatively short tail and robust limbs that were held in an odd mix of sprawled hind limbs and raised forelimbs. It had a long neck and a proportionately small head with remarkably sauropod-like jaws and teeth.

Blikanasaurus is a genus of sauropodomorph dinosaur from the late Triassic of South Africa. The generic name Blikanasaurus is derived from Greek, meaning "lizard from Blikana". The species name cromptoni is taken from the surname of A.W. “Fuzz” Crompton, an American paleontologist who led numerous field expeditions in Elliot Formation outcrop localities in South Africa. Blikanasaurus is only known from partial hindlimb bones that were recovered from the lower Elliot Formation (LEF) in the Eastern Cape.

<i>Euskelosaurus</i> Extinct genus of dinosaur from late Triassic southern Africa

Euskelosaurus is a sauropodomorph dinosaur from the Late Triassic of South Africa and Lesotho. Fossils have only been recovered from the lower Elliot Formation in South Africa and Lesotho, and in one locality in Zimbabwe.

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

Sinokannemeyeria is a genus of kannemeyeriiform dicynodont that lived during the Anisian age of Middle Triassic period in what is now Shanxi, China.

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

Emydops is an extinct genus of dicynodont therapsids from the Middle Permian to Late Permian of what is now South Africa. The genus is generally small and herbivorous, sharing the dicynodont synapomorphy of bearing two tusks. In the following years, the genus grew to include fourteen species. Many of these species were erected on the basis of differences in the teeth and the positioning of the frontal and parietal bones. A 2008 study narrowed Emydops down to two species, E. arctatus and the newly described E. oweni.

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

Moghreberia is an extinct genus of dicynodont predicted to have lived only in the mid-Triassic, primarily during the early middle Carnian and found only in the Angara Basin of Morocco. Moghreberia belonged to the Stahleckeriidae family, a group of anomodont therapsids and is most commonly known by its species Moghreberia nmachouensis. Its name is derived from the Arabic phrase al-Maghrib al-Aqsa meaning “the far west”, a term used by Arabic scholars to refer to the approximate region of Morocco, the area in which this animal’s fossil was first discovered. The extinction of many dicynodonts has been attributed to pressures of the Carnian Pluvial Episode, which occurred around 234-232 Ma and generated major ecological and climate changes for years to come.

<span class="mw-page-title-main">Elliot Formation</span> Lithostratigraphic layer of the Stormberg Group in South Africa

The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, southern Free State, and in the eastern KwaZulu-Natal provinces of South Africa. Outcrops and exposures are also found in several localities in Lesotho such as Qacha's Neck, Hill Top, Quthing, and near the capital, Maseru. The Elliot Formation is further divided into the lower (LEF) and upper (UEF) Elliot formations to differentiate significant sedimentological differences between these layers. The LEF is dominantly Late Triassic (Norian-Hettangian) in age while the UEF is mainly Early Jurassic (Sinemurian-Pliensbachian) and is tentatively regarded to preserve a continental record of the Triassic-Jurassic boundary in southern Africa. This geological formation is named after the town of Elliot in the Eastern Cape, and its stratotype locality is located on the Barkly Pass, 9 km north of the town.

Sangusaurus is an extinct genus of large dicynodont synapsid with two recognized species: S. edentatus and S. parringtonii. Sangusaurus is named after the Sangu stream in eastern Zambia near to where it was first discovered + ‘saur’ which is the Greek root for lizard. Sangusaurus fossils have been recovered from the upper parts of the Ntawere Formation in Zambia and of the Lifua Member of the Manda Beds in Tanzania. The earliest study considered Sangusaurus a kannemeyeriid dicynodont, but more recent phylogenetic analyses place Sangusaurus within the stahleckeriid clade of Dicynodontia. Until recently, little work had been done to describe Sangusaurus, likely due to the fact that only four incomplete fossil specimens have been discovered.

Zambiasaurus is an extinct genus of dicynodonts that was discovered in the Middle Triassic (Anisian) Ntawere Formation of Zambia, southern Africa. It was a large dicynodont, reconstructed using several fossil fragments, in majority belonging to probably a juvenile Zambiasaurus submersus.

<span class="mw-page-title-main">Stormberg Group</span> Triassic/Jurassic geological group in the Karoo Supergroup in South Africa

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<i>Arcusaurus</i> Extinct genus of dinosaur from early Jurassic South Africa

Arcusaurus is an extinct genus of sauropodomorph dinosaur from the Early Jurassic of South Africa.

<i>Diodorus scytobrachion</i> Extinct species of reptile

Diodorus is a genus of silesaurid dinosauromorph that lived during the Late Triassic in what is now Morocco. Fossils were discovered in the Timezgadiouine Formation of the Argana Basin, and were used to name the new genus and species Diodorus scytobrachion. The genus name honors the mythological king Diodorus and the ancient historian Diodorus Siculus; the specific name is ancient Greek for 'leathery arm' and also honors the mythographer Dionysius Scytobrachion. The holotype specimen is a partial dentary bone (front of the lower jaw), and assigned specimens include isolated teeth, two humeri (upper arm bones), a metatarsal (foot bone), and femur (thigh bone).

<i>Lisowicia</i> Genus of giant dicynodont therapsid

Lisowicia is an extinct genus of giant dicynodont synapsid that lived in what is now Poland during the late Norian or earliest Rhaetian age of the Late Triassic Period, about 210–205 million years ago. Lisowicia is the largest known dicynodont, as well as the largest non-mammalian synapsid, and is estimated to have weighed between 5–6 tons, comparable in size to modern elephants. It was also one of the last dicynodonts, living shortly before their extinction at the end of the Triassic period. Fossils of a giant dicynodont were known from Poland since 2008, but Lisowicia was not named and officially described as a new species until late 2018.

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

Thliptosaurus is an extinct genus of small kingoriid dicynodont from the latest Permian period of the Karoo Basin in KwaZulu-Natal, South Africa. It contains the type and only known species T. imperforatus. Thliptosaurus is from the upper Daptocephalus Assemblage Zone, making it one of the youngest Permian dicynodonts known, living just prior to the Permian mass extinction. It also represents one of the few small bodied dicynodonts to exist at this time, when most other dicynodonts had large body sizes and many small dicynodonts had gone extinct. The unexpected discovery of Thliptosaurus in a region of the Karoo outside of the historically sampled localities suggests that it may have been part of an endemic local fauna not found in these historic sites. Such under-sampled localities may contain 'hidden diversities' of Permian faunas that are unknown from traditional samples. Thliptosaurus is also unusual for dicynodonts as it lacks a pineal foramen, suggesting that it played a much less important role in thermoregulation than it did for other dicynodonts.

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

Repelinosaurus is an extinct genus of dicynodont from the Purple Claystone Formation of Luang Prabang in Laos, Southeast Asia that lived at around the time of the Permian-Triassic boundary and possibly dates to the earliest Early Triassic. Its type and only known species is R. robustus. Repelinosaurus was originally described as the earliest known kannemeyeriiform dicynodont, supporting the idea of a more rapid radiation of the Triassic kannemeyeriiform dicynodonts during the Early Triassic following the Permian mass extinction. However, it may alternatively be more closely related to the Permian Dicynodon. The discovery of a potential early kannemeyeriiform in an understudied locality like Laos highlights the importance of such places in dicynodont research, which has been largely focused on historically important localities such as the Karoo Basin of South Africa.

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

Ufudocyclops is an extinct genus of stahleckeriid dicynodont from the Middle Triassic of South Africa. It was found in the Burgersdorp Formation, part of the uppermost Cynognathus Assemblage Zone of the Beaufort Group in the Karoo Basin. The type and only known species is U. mukanelai. It was a large, beaked herbivore like other Triassic dicynodonts, lacking tusks, and is mostly characterised by unique features of the skull. It is known from three specimens, two of which were previously referred to the Tanzanian dicynodont Angonisaurus. The separation of Ufudocyclops from Angonisaurus indicates that the Middle Triassic fauna of the Beaufort Group in South Africa was not part of a larger shared fauna with those of the Manda Beds in Tanzania, as was previously supposed, and suggests that they were separated as more localised faunas, possibly by geographic barriers or in time. Ufudocyclops then would have been a unique part of the uppermost Cynognathus Assemblage Zone in South Africa. It is also the oldest known member of the family Stahleckeriidae, and implies that the family was already diversifying in the Middle Triassic alongside other kannemeyeriiforms, not just in the Late Triassic after other families died out.

Taoheodon is an extinct genus of dicynodont therapsid from the Sunjiagou Formation in the Shanxi province of China, dated to the Wuchiapingian age of the Late Permian. Its type and only known species is T. baizhijuni. Taoheodon was a close relative of the well known Dicynodon, and may represent a biogeographical link between the South African Dicynodon and similar dicynodonts found in Laos.

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