Kunpania | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Clade: | † Bidentalia |
Infraorder: | † Dicynodontoidea |
Genus: | † Kunpania Sun, 1978 |
Species: | †K. scopulusa |
Binomial name | |
†Kunpania scopulusa Sun, 1978 | |
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.
The fossil remains of Kunpania were collected from the uppermost layers of the Quanzijie Formation in the Bogda Mountains of Jimsar in Xinjiang, China. This discovery was notable, as the majority of dicynodont fossils from the Bogda Mountains come from the younger Guodikeng Formation above it, and indeed Kunpania was the only tetrapod known from the formation until the femur of a chroniosuchian was discovered in 2020. [1] The precise age of these deposits is uncertain, although they are typically thought to be from the Capitanian age at the end of the Guadalupian (or Middle Permian) based on palynology, cyclostratigraphy and constrained radiometric dating in surrounding rock units. However, it is possible that the uppermost Quanzijie Formation dates to the Wuchiapingian in the Lopingian (Late Permian), albeit no younger than 254.22 million years old. [2]
Kunpania was first described in 1978 by palaeontologist Ailing Sun from the type and only known specimen IVPP V4695. The specimen originally only consisted of a partial snout missing the tip, a broken lower jaw and part of the forelimb including the right scapula, coracoid plate and humerus. Additionally, a partial right clavicle and a piece of the left tibia with the same specimen number were later recovered from the Institute of Vertebrate Paleontology and Paleoanthropology collections that matched the preservation style of the holotype, and so likely belong to the same specimen. [2] In 1998 and 2001, palaeontologist Spencer G. Lucas synonymised Kunpania into Dicynodon as a distinct species, D. scopulusa, though he later considered the species to be dubious (a nomen dubium ) in 2005. [3] [4] [5] The species was again considered valid by Jörg Fröbisch in 2009, who again regarded it as a species of Dicynodon. [6] In their 2011 revision of the taxonomy of Dicynodon, palaeontologists Christian Kammerer, Kenneth Angielczyk, and Fröbisch tentatively concluded that 'D.' scopulusa may be a valid taxon, but that its status was ultimately uncertain owing to its incompleteness. [7]
Kunpania was fully redescribed in 2021 by Angielczyk, Jun Liu and Wan Yang, including further preparation of the underside of the skull that revealed additional features. Angielczyk and colleagues concluded that Kunpania was indeed a wholly distinct genus and species of dicynodont based on a unique combination of physical traits and comparisons with other dicynodonts. [2]
Kunpania was a relatively large dicynodont, with a preserved basal skull length of 307 millimetres (12.1 in) and an estimated complete basal length of 442 millimetres (17.4 in) based on Daptocephalus . Like other bidentalian dicynodonts, Kunpania had no teeth and a beak except for two tusks in the upper jaws. The tusks themselves are mostly missing in the only known specimen, but the caniniform processes that housed them are angled strongly forwards, and so the tusks sat ahead of the eyes on the snout. Although the very tip is missing, the preserved shape of the snout suggests that the beak was rounded-off and somewhat hooked. Likewise, the top surface of the skull is damaged, so it is unknown if Kunpania possessed nasal bosses like some other dicynodonts. [2]
The secondary palate on the roof of the mouth, composed of the premaxillary bones, is unusually smooth in Kunpania, with only a single midline ridge at the back. The crista oseophagea, a bony process of the pterygoid on the roof of the mouth, is unusually robust and blocky, unlike the blade-like structure more typical of bidentalians and more like those of more primitive dicynodonts. [2]
The mandible resembles those of other dicynodonts, with a squared-off beak that is upturned at the tip. The lateral dentary shelf, a site for jaw muscle attachment, is well developed and overhangs the mandibular fenestra. The shelf is unusually thick compared to other dicynodontoids, and is angled outwards in a triangular shape from above, more similar to its shape in the unrelated Dicynodontoides (a non-dicynodontoid dicynodont). Just behind the mandibular fenestra, there is an unusual backwards-curving ridge on the angular bone. This ridge sits where the reflected lamina (a bony structure potentially used in hearing) [8] usually attaches in other dicynodonts, however they are usually attached by a vertical ridge, unlike the horizontally curving structure in Kulpania. The reflected lamina may then have been unusually shaped. [2]
The scapular is robust, with a glenoid (shoulder socket) that faced back and out to the sides in life. The humerus of the upper arm possesses a pointed triangular process jutting from the back side of the bone, likely the attachment point for the latissimus dorsi muscle. In all other dicynodonts this process is usually low and flattened, and so the exaggerated size of this process may indicate a specialised function of the forelimbs in Kunpania. The head of the humerus may also suggest a specialisation, as it is notably narrower than is typical for dicynodonts. [2]
Kunpania was originally classified as a member of the family Dicynodontidae by Sun in 1978, and later as a species of Dicynodon itself by later authors. Kunpania would not be included in a phylogenetic analysis until its redescription in 2021, where it was found to be an early diverging member of the clade Dicynodontoidea and most closely related to the South African genus Sintocephalus .
The cladogram produced by Angielczyk et al. (2021) is reproduced and simplified below: [2]
Kunpania is diagnosed by the absence of ridges on the palate at the front of the mouth, a robust and block-like crista oseophagea, a prominent and thick lateral dentary shelf on the lower jaw, a small backwards-curving projection of the angular bone, and the large, triangular shaped attachment for the latissimus dorsi on the humerus. [2]
As an early dicynodontoid, Kunpania could reveal information about the evolution and radiation of bidentalian dicynodonts. If the Capitanian age is correct, it would indicate that bidentalian dicynodonts had radiated earlier than the fossil record indicates, before the Capitanian mass extinction event, and that the early history of bidentalians is otherwise unknown in the fossil record. It would also create extensive ghost lineages amongst bidentalians, including for the three traditional lineages of cryptodonts, 'elphids', and at least three lineages of dicynodontoids. [2]
Alternatively, an early to middle Wuchiapingian age would be more consistent with the known record of bidentalian dicynodonts, being a contemporary of early cryptodonts and still the oldest known dicynodontoid. The latter is also in keeping with its relatively basal phylogenetic position and would fill the ghost lineage between derived dicynodontoids and cryptodonts. This scenario would suggest bidentalians originated before the end-Capitanian extinction but only radiated afterwards, rapidly spreading across Pangaea to Laurasia and Gondwana. A third hypothesis, that Kunpania is mid-late Wuchiapingian in age, would result in no change to the known pattern of bidentalian evolution. It would also require that the age of the upper Quanzijie Formation to be significantly underestimated and drastically shorten the period of time between it and the overlying Guodikeng Formation. [2]
Paleosols (fossilised soils) from the uppermost Quanzijie Formation are mostly of water-logged gleysols, indicating a wet humid to subhumid climate. The only other known tetrapod contemporary with Kunpania is an indeterminate chroniosuchian, a semi-aquatic predatory reptiliomorph. [1] [2]
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.
Robertia is an extinct genus of small herbivorous dicynodonts from the Middle to Late Permian of South Africa, between 260 and 265 million years ago. It is a monospecific genus, consisting of the type-species R. broomiana, which was classified by Lieuwe Dirk Boonstra in 1948 and named in honor of Robert Broom for his study of South African mammal-like reptiles.
Cistecephalus is an extinct genus of dicynodont therapsid from the Late Permian of southern Africa. It was a small, specialised, burrowing dicynodont, possibly with habits similar to a modern mole. The head was flattened and wedge-shaped, the body long, and the forelimbs very strong, with similarities in structure to the forelimb of modern burrowing mammals.
Dicynodon is a genus of dicynodont therapsid that flourished during the Upper Permian period. Like all dicynodonts, it was an herbivorous animal. This synapsid was toothless, except for prominent tusks, hence the name. It probably cropped vegetation with a horny beak, much like a tortoise, while the tusks may have been used for digging up roots and tubers.
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.
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.
Daptocephalus is an extinct genus of dicynodont synapsid, which was found in Late Permian strata, in a biozone known precisely for the presence of fossils of this dicynodont, the Daptocephalus Assemblage Zone, in the Karoo Basin in South Africa. An additional species, D. huenei, is known from the Usili Formation in Tanzania and was formerly assigned to the genus Dicynodon before a study in 2019 recognised that the type specimen belonged to Daptocephalus.
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.
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.
Lystrosauridae is a family of dicynodont therapsids from the Permian and Triassic time periods. It includes two genera, Lystrosaurus and Kwazulusaurus. Kwazulusaurus includes a single species, K. shakai, from the Late Permian of South Africa and Lystrosaurus includes many species from the Late Permian and Early Triassic of South Africa, India, and Antarctica.
Pylaecephalidae is a family of dicynodont therapsids that includes Diictodon, Robertia, and Prosictodon from the Permian of South Africa. Pylaecephalids were small burrowing dicynodonts with long tusks. The family was first named in 1934 and was redefined in 2009. Diictodontidae and Robertiidae are considered junior synonyms of Pylaecephalidae; although Pylaecephalus itself is considered a junior synonym of Diictodon, the name Pylaecephalidae predates these names and therefore takes priority.
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.
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.
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.
Parasumina is an extinct genus of anomodont known from the late Capitanian age at the end of the middle Permian period of European Russia. The type and only species is Parasuminia ivakhnenkoi. It was closely related to Suminia, another Russian anomodont, and was named for its resemblance. Little is known about Parasuminia as the only fossils are of fragmentary pieces of the skull and jaw, but the known remains suggest that its head and jaws were deeper and more robust than those of Suminia, and with shorter, stouter teeth. However, despite these differences they appear to have been similar animals with a similarly complex method of processing vegetation.
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.
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.
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.
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.