Kannemeyeriiforms Temporal range: Triassic, | |
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Restoration of Stahleckeria potens | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Infraorder: | † Dicynodontoidea |
Clade: | † Kannemeyeriiformes von Huene, 1948 |
Subgroups | |
Kannemeyeriiformes is a group of large-bodied Triassic dicynodonts. As a clade, Kannemeyeriiformes has been defined to include the species Kannemeyeria simocephalus and all dicynodonts more closely related to it than to the species Lystrosaurus murrayi .
Despite being the most species-rich group of dicynodonts in the Triassic Period, kannemeyeriiforms exhibit much less diversity in terms of their anatomy and ecological roles than the dicynodonts from the Permian Period.
Lystrosauridae is thought to be the most closely related group (sister taxon) to Kannemeyeriiformes, and since the earliest lystrosaurids are known from the Late Permian, the divergence of these two groups must have occurred at least as far back as this time, implying that a long ghost lineage must exist. [1] Although no kannemeyeriiforms have been found in the Late Permian yet, the recent discovery of Sungeodon helps fill a gap in the early fossil record of the group by showing that kannemeyeriiforms diversified right after the Permian-Triassic extinction event. [2]
Four families have been included in the group: Kannemeyeriidae, Shansiodontidae, Stahleckeriidae, and Dinodontosauridae. [3] Occasionally some of these families are not recognized, with most kannemeyeriiforms being placed in Kannemeyeriidae. [4] Recent phylogenetic analyses suggest that these families represent similar body plans or morphotypes rather than true evolutionary groupings. [1]
Below is a cladogram from Szczygielski & Sulej (2023): [5]
Kannemeyeriiformes | |
Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida. The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.
Therapsida is a clade composing of a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, resulting in a more "standing" quadripedal posture, as opposed to the lower sprawling posture of many reptiles and amphibians.
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.
Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores. Anomodonts were very diverse during the Middle Permian, including primitive forms like Anomocephalus and Patranomodon and groups like Venyukovioidea and Dromasauria. Dicynodonts became the most successful and abundant of all herbivores in the Late Permian, filling ecological niches ranging from large browsers down to small burrowers. Few dicynodont families survived the Permian–Triassic extinction event, but one lineage (Kannemeyeriiformes) evolved into large, stocky forms that became dominant terrestrial herbivores right until the Late Triassic, when changing conditions caused them to decline, finally going extinct during the Triassic–Jurassic extinction event.
Stahleckeriidae is a family of dicynodont therapsids whose fossils are known from the Triassic of North America, South America, Asia and Africa.
Dinodontosaurus is a genus of dicynodont therapsid. It was medium to large dicynodont of the Triassic and had a beak corneum. It lived in the Middle Triassic but disappeared in the Upper Triassic.
Dolichuranus is an extinct genus of dicynodont therapsids from the Middle Triassic Omingonde Formation of Namibia and the Ntawere Formation of Zambia.
Wadiasaurus is an extinct genus of dicynodont from the family Kannemeyeria, that lived in herds from the early to Middle Triassic. Substantial fossorial evidence of W. indicus was recovered from Yerrapalli Formation of the Pranhita-Godavari valley, India, and it is so far the only Kannemeyeriid known for certain from India. The Kannemeyeriiformes underwent a significant diversification during the middle Triassic, with roughly 40 known species distributed worldwide. All Kannemeyeriiformes were medium to large bodied, graviportal herbivores with relatively erect posture and gait. Wadiasaurus indicus is currently the only known species of Wadiasaurus.
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.
Vinceria is an extinct genus of kannemeyeriiform dicynodont in the family Shansiodontidae. Fossils of the genus have been found in the Anisian Cerro de las Cabras Formation and Carnian Río Seco de la Quebrada Formation of Argentina. One species, V. argentinensis, named in 1966, was moved to its own genus, Acratophorus, in 2021. Another species, V. vieja, was merged with Acratophorus argentinensis in 2021, leaving V. andina as the only species in the genus.
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.
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.
Sungeodon is an extinct genus of dicynodont therapsid from the Early Triassic of China. It is known from a single type species, Sungeodon kimkraemerae, which was named in 2014. Sungeodon is the earliest member of a group of dicynodonts called Kannemeyeriiformes, which would radiate later in the Triassic to become the dominant large herbivores of terrestrial ecosystems. Before its discovery no kannemeyeriiform dicynodonts were known from the Early Triassic. The presence of Sungeodon in the earliest Triassic Jiucaiyuan Formation indicates that dicynodonts diversified soon after the Permian-Triassic extinction event, mirroring the explosive radiations of other tetrapod groups such as archosaurs soon after the extinction.
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.
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.
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.
Woznikella is an extinct genus of kannemeyeriiform dicynodont from the Late Triassic of Poland and possibly Germany of Europe. The type and only known species is W. triradiata.