Taoheodon

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Taoheodon
Temporal range: Late Permian
~ Wuchiapingian
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Suborder: Anomodontia
Clade: Dicynodontia
Infraorder: Dicynodontoidea
Genus: Taoheodon
Liu, 2020
Species:
T. baizhijuni
Binomial name
Taoheodon baizhijuni
Liu, 2020

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.

Contents

Description

Taoheodon was a medium-sized dicynodont (basal skull length over 10 centimetres (3.9 in) long) currently only known from an incomplete skull and lower jaw. Based on the anatomy of other closely related dicynodonts such as Dicynodon, Taoheodon was probably a squat, sprawling quadruped with a short tail and a proportionally large head. Like other dicynodonts, Taoheodon was almost entirely toothless, sporting only a pair of tusks and a tortoise-like beak. [1] [2]

Skull

Like other dicynodonts, Taoheodon had a short skull with large temporal fenestra at the back, large orbits and a short snout, which in Taoheodon was proportionately short even for a dicynodont. Its skull is slightly longer than wide, with elongated temporal fenestra. The external nostril is rounded and not especially large for a dicynodont, but the area of bone behind and beneath it is hollowed out and concave compared to the rest of the snout. The nasal bones along the roof of the snout are relatively flat, but are nonetheless rugosely textured and bore a single weakly developed boss of tough skin or keratin on top of the snout. Likewise, the lacrimal and prefrontal bone form a distinct boss that bulges out to the side in front of each eye. By comparison, the postorbital bar behind the eyes is smooth and unornamented. The caniniform process housing the tusk is directed downwards from the snout, and sits entirely in front of the eyes. The pineal foramen (or "third eye") on the roof of the skull is large and positioned relatively far back. [1]

Lower jaw

The mandible of Taoheodon is mostly known from part of the dentary, with portions of the angular, surangular and splenial bones. The dentary is large and robust, with a rough, pitted surface texture at its front and along the top surface, corresponding to the horny beak typical of dicynodonts. The tip of the lower jaw is missing, so the exact shape of the beak is unknown; however, a low and wide curved ridge defines a clear edge between the side and front faces of the beak. Like other dicynodonts, the angular supports a prominent reflected lamina, which may have supported the eardrum in non-mammalian therapsids. In Taoheodon, the reflected lamina is large and rounded, facing down and back from the mandibular fenestra. [1] [3]

History of discovery

The holotype and only known specimen of Taoheodon, IVPP V 25335, was discovered in the valley of a tributary of the Tao He river, running through the lower part of the Sunjiagou Formation. The Sunjiagou Formation has been dated to the Late Permian (Lopingian epoch), although the exact age of the lower beds has been debated; either representing the late Wuchiapingian age or early Changhsingian. The lower Sunjiagou Formation is composed of grey to greenish-grey mudstones and fine grained sandstones, although the fossil of Taoheodon itself was found contained within an eroded rock nodule. This erosion resulted in the loss of the zygomatic arches and the tip of the snout from the specimen which had been exposed prior to collection, and the specimen has also been slightly compressed from top to bottom during fossilisation. The specimen was described in 2020 by Jun Liu as a new genus and species, Taoheodon baizhijuni. Taoheodon was named for the nearby Tao He river where it was discovered, combined with the Ancient Greek -odous for 'tooth', a common suffix in dicynodont generic names. The specific name is in honour of the fossil collector Bai Zhijun who discovered the specimen. [1] [4]

Classification

Taoheodon was a member of the dicynodont infraorder Dicynodontoidea, and is distinguished from all other dicynodonts by three unique autapomorphies: the top of postorbital bars behind the eyes have a shallow depression (fossa) where they meet the rest of the skull, the basisphenoid (a part on the underside of the braincase) slopes anterodorsally at a shallow angle in the basisphenoid-basioccipital tubera, and unlike other dicynodonts it lacks a keel on the pterygoid bones of the palate. Taoheodon was included in an updated phylogenetic analysis of dicynodonts using the combined datasets of Olivier et al. (2019) [5] and Kammerer (2019). [6] Within Dicynodontoidea, Taoheodon was found to group within a clade containing Dicynodon and very similar taxa that Liu identified as the "core-Dicynodon" clade.

The cladogram produced by Liu (2020), simplified and focused on the relationships of dicynodontoids, is shown below: [1]

Dicynodontoidea

Keyseria

Daqingshanodon

Sintocephalus

Basilodon

Syops

Euptychognathus

Lystrosaurus

Peramodon

Daptocephalus

Dinanomodon

Turfanodon

Dicynodon

Delectosaurus

Vivaxosaurus

Taoheodon

Counillonia

Repelinosaurus

Kannemeyeriiformes

"Core-Dicynodon" clade

The results of the analysis are almost identical to the cladograms produced from the previous studies, however, the position of the two Laotian dicynodonts Counillonia and Repelinosaurus differs from their original descriptions. The two Laotian genera were found to clade together with Taoheodon in the "core-Dicynodon" clade, contrasting with the analysis of Olivier and colleagues which originally found Repelinosaurus to be the basalmost kannemeyeriiform. Liu found Taoheodon and the Laotian dicynodonts to share a number of features, including notably short snouts, pineal foramens placed further back on the roof of the skull, anteriorly inclined occiputs, a fairly straight suture between the nasals and frontals, lacking the postfrontal bones, fairly flat postorbitals in the temporal area, and a large fossa on the ventral surface of the intertemporal bar. [1] [5]

Palaeobiogeography

The close relationship of Taoheodon to the Laotian dicynodonts suggests that there was a direct link between the North China Block, South China Block and Indochina Block that created a corridor of land for dicynodonts in Northern China to disperse into Laos on the Indochina Block and speciate. This paleobiogeographic inference has implications for the timing of the collisions between these landmasses, which although uncertainly dated, have typically been inferred to have occurred later during the Triassic period. The presence of a clade of closely related dicynodonts between these landmasses suggests that they were connected in some way by the end of the Permian. Furthermore, their position within a "core-Dicynodon" clade indicates that Taoheodon was part of a lineage of dicynodonts that could freely migrate from North China through Russia to South Africa. [1]

Palaeoecology

In the lower part of the Sunjiagou Formation, Taoheodon coexisted with the diminutive pareiasaurian parareptile Sanchuansaurus (a possible a relative of the better known Elginia ), a probable large cryptodont dicynodont, and the late surviving semi-aquatic predatory embolomere Seroherpeton . [4] [7] [8] The lower part of the Sunjiagou Formation is largely composed of fluvial and lacustrine sediments with infrequent coal seams that indicate a wet, swampy environment with abundant lake-shore habitats that preserve vertical burrows and rich, intensive bioturbation by aquatic organisms. [9] [10] Plant remains include Ningxiaites specialis , a partially deciduous conifer with an evergreen canopy that would shed only some of its leaves annually. [11]

Related Research Articles

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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>Euchambersia</i> Extinct genus of therapsid from Late Permian South Africa

Euchambersia is an extinct genus of therocephalian therapsids that lived during the Late Permian in what is now South Africa and China. The genus contains two species. The type species E. mirabilis was named by paleontologist Robert Broom in 1931 from a skull missing the lower jaw. A second skull, belonging to a probably immature individual, was later described. In 2022, a second species, E. liuyudongi, was named by Jun Liu and Fernando Abdala from a well-preserved skull. It is a member of the family Akidnognathidae, which historically has also been referred by as the synonymous Euchambersiidae.

<i>Biseridens</i> Extinct genus of therapsids

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

Dimacrodon is an extinct genus of non-mammalian synapsid from the latest Early Permian San Angelo Formation of Texas. It is distinguished by toothless, possibly beaked jaw tips, large lower canines and a thin bony crest on top of its head. Previously thought to be an anomodont therapsid related to dicynodonts, it was later found to lack any diagnostic features of anomodonts or even therapsids and instead appears to be a 'pelycosaur'-grade synapsid of uncertain classification.

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

Dicynodontoides is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name Dicynodontoides references its “dicynodont-like” appearance due to the caniniform tusks featured by most members of this infraorder. Kingoria, a junior synonym, has been used more widely in the literature than the more obscure Dicynodontoides, which is similar-sounding to another distantly related genus of dicynodont, Dicynodon. Two species are recognized: D. recurvidens from South Africa, and D. nowacki from Tanzania.

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

Odontocyclops is an extinct genus of Dicynodonts that lived in the Late Permian. Dicynodonts are believed to be the first major assemblage of terrestrial herbivores. Fossils of Odontocyclops have been found in the Karoo Basin of South Africa and the Luangwa Valley of Zambia. The phylogenetic classification of Odontocyclops has been long under debate, but most current research places them as their own genus of Dicynodonts and being very closely related to Rhachiocephalus and Oudenodon.

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

Pelanomodon is an extinct genus of dicynodont therapsids that lived in the Late Permian period. Fossil evidence of this genus is principally found in the Karoo Basin of South Africa, in the Dicynodon Assemblage Zone. Lack of fossil record after the Late Permian epoch suggests that Pelanomodon fell victim to the Permian-Triassic extinction event.

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

Tropidostoma is a medium-sized herbivorous oudenodontid dicynodont therapsid that lived during the Late Permian (Lopingian) period in South Africa. The first Tropidostoma fossil was described by Harry Govier Seeley in 1889. Later two subspecies were identified. Tropidostoma fossils are an index fossil in a biozone of the Karoo Basin known as the Tropidostoma Assemblage Zone. This biozone is characterized by the presence of this species in association with another dicynodont species, Endothiodon uniseries.

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

Kombuisia is a genus of dicynodont from Early to Middle Triassic of South Africa and Antarctica. Two species were described for the genus: Kombuisia frerensis (type) and Kombuisia antarctica.

Syops is an extinct genus of dicynodont therapsid. The type species S. vanhoepeni was first named in 1938 as Dicynodon vanhoepeni. Fossils of the genus have been found in the Cistecephalus Assemblage Zone in the Usili Formation of the Ruhuhu Basin, Tanzania and the Upper Madumabisa Mudstone Formation of the Luangwa Basin, Zambia. Its phylogenetic placement is somewhat uncertain, with multiple different studies finding it as either a basal geikiid, rhachiocephalid a dicynodontoid more derived than the most basal genera but less derived than Lystrosauridae, or a lystrosaurid.

Daqingshanodon is an extinct genus of dicynodont therapsid from the Late Permian of Inner Mongolia, China. The type species D. limbus was described in 1989 from a single skull found in the Naobaogou Formation. Daqingshanodon belongs to a group of dicynodonts called cryptodonts. It is the smallest known cryptodont, and the only one known from China. Like other cryptodonts, it has a pair of rounded nasal bosses above its nostrils and a ridge of bone on the upper jaw called the postcaniniform process. Daqingshanodon has a pair of elongated, recurved tusks extending from its beak-like snout. It is distinguished from other dicynodonts by the presence of a distinct ridge running along the side of the skull from below the eye socket to the area around the tusks. The skull of Daqingshanodon is less than 10 centimetres (3.9 in) long, yet this specimen is thought to have been an adult on the basis of its well-developed nasal bosses.

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

Jimusaria is an extinct genus of dicynodont therapsid from the Late Permian (Changhsingian) of China. The type species J. sinkianensis from the Guodikeng Formation in Xinjiang, was originally named as a species of Dicynodon, the first from Asia, but was given its own genus in 1963 before being sunk back into Dicynodon in 1988. The genus was resurrected in 2011 by palaeontologist Christian Kammerer in a taxonomic revision of the genus Dicynodon. Jimusaria was a mid-sized dicynodont, and was similar in appearance to the South African Dicynodon, but differed from it in features such as its narrower snout. A second species, Jimusaria monanensis was described from the Naobaogou Formation of northern China in 2023.

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

Turfanodon is an extinct genus of dicynodont therapsid from the Late Permian Sunan, Guodikeng, and Naobaogou Formations of China. The holotype of T. bogdaensis was discovered between 1963-1964 and was originally named in 1973 by A. Sun with the type species Turfanodon bogdaensis, Turfanodon was reclassified as a junior synonym of the related Dicynodon in 1988 by G. M. King. T. bogdaensis remained a species of Dicynodon for over two decades before the genus was reinstated in 2011 in a revision of the taxonomy of Dicynodon by palaeontologist Christian Kammerer. A second species from Inner Mongolia, T. jiufengensis, was named in 2021 by palaeontologist Jun Liu from a nearly complete skeleton and other referred bones. Turfanodon was a relatively large dicynodont, and similar in appearance to the related Daptocephalus from South Africa.

Kunpania is an extinct genus of dicynodont therapsid from the Quanzijie Formation of Xinjiang, China. The type and only species is K. scopulusa, and it is known only by a single incomplete specimen including parts of the skull and forelimb. Since its initial description in 1978 by palaeontologist Ailing Sun, it has sometimes been considered to be another species of Dicynodon by other researchers, or potentially undiagnostic. However, a redescription in 2021 reaffirmed its distinctiveness, including a uniquely well developed muscle attachment on the humerus. Kunpania is perhaps the oldest known member of the derived dicynodont group Dicynodontoidea, potentially dating to the Middle Permian period during the Capitanian, and so may fill a knowledge gap in the history of dicynodont evolution.

<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 Aulacocephalodon, although a more basal position is not implausible.

<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>Counillonia</i> Extinct genus of dicynodonts

Counillonia is an extinct genus of dicynodont therapsid from the area 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 C. superoculis. Counillonia was related to the Triassic dicynodonts such as Lystrosaurus and the Kannemeyeriiformes that survived the Permian mass extinction, but it was more closely related to the Permian genus Dicynodon than to either of these lineages. Counillonia may then possibly represent another line of dicynodonts that survived the Permian mass extinction into the Triassic period, depending on its age. The discovery of Counillonia in Laos and its unexpected evolutionary relationships hint at the less well understood geographies of dicynodont diversity across the Permo-Triassic boundary outside of well explored regions like the Karoo Basin in South Africa.

<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.

Kembawacela is an extinct genus of cistecephalid dicynodont from the Late Permian of East Africa. The genus contains two known species, the type species Kembawacela kitchingi from the Madumabisa Mudstone Formation of Zambia described in 2019, and a second species, K. yajuwayeyi, from the Chiweta Beds of Malawi described in 2022. Like other cistecephalids, Kembawacela was specialised for a fossorial, burrowing lifestyle similar to modern day moles. It is unique amongst cistecephalids for the presence of a pair of tusks in the upper jaw, characteristic of many other dicynodonts but lost in other cistecephalids. It is likely that Kembawacela was a locally endemic species of cistecephalid in the Luangwa Basin of Zambia.

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