Broomistega Temporal range: Early Triassic, | |
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Skeleton of a juvenile Broomistega putterilli shown in ventral (C) and dorsal (D) views | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Amphibia |
Order: | † Temnospondyli |
Suborder: | † Stereospondyli |
Family: | † Rhinesuchidae |
Genus: | † Broomistega Shishkin and Rubidge, 2000 |
Type species | |
Lydekkerina putterilli Broom, 1930 |
Broomistega is an extinct genus of temnospondyl in the family Rhinesuchidae. It is known from one species, Broomistega putterilli, which was renamed in 2000 from Lydekkerina putterilli Broom 1930. Fossils are known from the Early Triassic Lystrosaurus Assemblage Zone of the Beaufort Group in the Karoo Basin of present-day South Africa, a region that had been an enclave of Gondwana. Specimens of B. putterilli were once thought to represent young individuals of another larger rhinesuchid such as Uranocentrodon , but the species is now regarded as a paedomorphic taxon, possessing the features of juvenile rhinesuchids into adulthood. [1]
In 2013, a well-preserved skeleton of Broomistega was discovered alongside the skeleton of the cynodont Thrinaxodon (a mammal relative) in a cast of a burrow. The individual probably entered the burrow while the cynodont was in a state of aestivation (dormancy), and afterwards a flash flood filled the burrow with sediment to preserve both bodies together. [2]
Broomistega is the only rhinesuchid known from the Triassic Period. Its presence in the Early Triassic indicates that rhinesuchids survived the Permo-Triassic Mass Extinction about 252 million years ago. However, compared to the diversity of rhinesuchids that existed in the Permian Period, Broomistega is a very rare component of the Early Triassic Karoo fauna. It may have been the last surviving representative of the group, making it a relict taxon. [3]
The most complete skeleton of Broomistega, specimen BP/1/7200, was discovered in the sandstone cast of a burrow (BP/1/5558) after the cast was scanned at the European Synchrotron Radiation Facility in 2013. The cast was first found in the Karoo Basin of South Africa by paleontologist James Kitching in 1975, but was left unprepared for many years. Part of a skull of a cynodont was exposed on the surface of the cast, allowing Kitching to attribute it to the genus Thrinaxodon . Only after the synchrotron scanning was the Broomistega skeleton found. All bones are preserved except for a few phalanges of the right hind foot, and nearly all of the bones are articulated as they were in life. The skeletons do not show evidence of stiffening due to rigor mortis after death, but are pressed against the sides of the burrow as the animals would have been when alive. The skeleton of BP/1/7200 is preserved belly-up, resting on the right side of the Thrinaxodon skeleton. This position was probably the result of the individual being pushed on top of the Thrinaxodon by floodwater entering the burrow. [2] Broomistega is not thought to have been a burrowing animal and was instead semiaquatic. The Thrinaxodon individual preserved in BP/1/5558 was probably the original occupant of the burrow, and the Broomistega individual probably entered the burrow later. The joints of BP/1/7200 are not well developed, indicating that the individual was a juvenile with large amounts of cartilage in its skeleton. Several ribs on the right side of the skeleton are broken, indicating that the individual was severely injured before entering the burrow, probably by being crushed. The animal was still alive when it entered the burrow because the rib fractures show evidence of healing, but the injury probably impacted its ability to breathe and to move about. Two holes are present on the skull roof of BP/1/7200. Although they resemble bite marks, they do not match with the teeth of the Thrinaxodon specimen. [2]
The presence of two different species of large vertebrates in the same burrow is unusual. Modern examples of this association are usually the result of predator-prey interactions (for example, a predator storing the body of its prey in the burrow) or mutualistic relationships whereby the original occupant gains protection from predators by the presence of the second inhabitant. However, the benefit of cohabitation usually only works when there are multiple burrows, casting doubt on the possibility that the Thrinaxodon was benefiting from the presence of the Broomistega. Because the Broomistega skeleton lacks any sign of damage caused by the Thrinaxodon, the two were probably not predator and prey. The most likely explanation for the association as of 2013 is that the Thrinaxodon tolerated the Broomistega or was unable to remove it, possibly because it was aestivating. As is the case with many modern amphibians, the Broomistega probably entered the burrow to seek temporary shelter. During the Early Triassic the Karoo Basin was seasonally arid, so the Thrinaxodon may have been aestivating to conserve energy during a time when resource availability was low and the normally aquatic Broomistega may have entered to burrow to escape the hot and dry conditions of its environment. [2]
Cynognathus is an extinct genus of large-bodied cynodontian therapsids that lived in the Middle Triassic. It is known from a single species, Cynognathus crateronotus. Cynognathus was a predator closely related to mammals and had a southern hemispheric distribution. Fossils have so far been recovered from South Africa, Argentina, Antarctica, and Namibia.
Thrinaxodon is an extinct genus of cynodonts, including the species T. liorhinus which lived in what are now South Africa and Antarctica during the Early Triassic. Thrinaxodon lived just after the Permian–Triassic mass extinction event, its survival during the extinction may have been due to its burrowing habits.
Galesaurus is an extinct genus of carnivorous cynodont therapsid that lived between the Induan and the Olenekian stages of the Early Triassic in what is now South Africa. It was incorrectly classified as a dinosaur by Sir Richard Owen in 1859.
Rhinesuchus is a large temnospondyl. Remains of the genus are known from the Permian of the South African Karoo Basin's Tapinocephalus and Cistecephalus assemblage zones, both belonging to the Beaufort Group. The skull of Rhinesuchus had a flat triangular shape with blunt snout similar to some of the other large temnospondyls, and had a palate filled with small sharp teeth, suggesting that it hunted fish. Also, the small eyes were on top of the head suggesting that it approached its prey from below.
Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian stage, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. Measuring 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb), Anteosaurus was the largest known carnivorous non-mammalian synapsid and the largest terrestrial predator of the Permian period. Occupying the top of the food chain in the Middle Permian, its skull, jaws and teeth show adaptations to capture large prey like the giants titanosuchids and tapinocephalids dinocephalians and large pareiasaurs.
The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecological niche to modern crocodilians prior to the diversification of pseudosuchian archosaurs.
The Tapinocephalus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the middle Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 2,000 metres (6,600 ft), occur from Merweville and Leeu-Gamka in its southernmost exposures, from Sutherland through to Beaufort West where outcrops start to only be found in the south-east, north of Oudshoorn and Willowmore, reaching up to areas south of Graaff-Reinet. Its northernmost exposures occur around the towns Fraserburg and Victoria West. The Tapinocephalus Assemblage Zone is the second biozone of the Beaufort Group.
The Lystrosaurus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Adelaide and lower Tarkastad Subgroups of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. This biozone has outcrops in the south central Eastern Cape and in the southern and northeastern Free State. The Lystrosaurus Assemblage Zone is one of eight biozones found in the Beaufort Group, and is considered to be Early Triassic in age.
The Cynognathus Assemblage Zone is a tetrapod biozone utilized in the Karoo Basin of South Africa. It is equivalent to the Burgersdorp Formation, the youngest lithostratigraphic formation in the Beaufort Group, which is part of the fossiliferous and geologically important Karoo Supergroup. The Cynognathus Assemblage Zone is the youngest of the eight biozones found in the Beaufort Group, and is considered to be late Early Triassic (Olenekian) to early Middle Triassic (Anisian) in age. The name of the biozone refers to Cynognathus crateronotus, a large and carnivorous cynodont therapsid which occurs throughout the entire biozone.
The Tropidostoma Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the lower Teekloof Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 240 metres (790 ft), occur from east of Sutherland through to Beaufort West and Victoria West, to areas south of Graaff-Reinet. Its northernmost exposures occur west/north-west of Colesberg. The Tropidostoma Assemblage Zone is the fourth biozone of the Beaufort Group.
Rhinesuchidae is a family of tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians are probably descended from. Rhinesuchids can be differentiated from other temnospondyls by details of their skulls, most notably the interior structure of their otic notches at the back of the skull. They were among the earliest-diverging members of the Stereospondyli, a subgroup of temnospondyls with flat heads and aquatic habits. Although more advanced stereospondyls evolved to reach worldwide distribution in the Triassic period, rhinesuchids primarily lived in the high-latitude environments of Gondwana during the Guadalupian and Lopingian epochs of the Permian. The taxonomy of this family has been convoluted, with more than twenty species having been named in the past; a 2017 review recognized only eight of them to be valid. While several purported members of this group have been reported to have lived in the Triassic period, most are either dubious or do not belong to the group. However, at least one valid genus of rhinesuchid is known from the early Triassic, a small member known as Broomistega. The most recent formal definition of Rhinesuchidae, advocated by Mariscano et al. (2017) is that of a stem-based clade containing all taxa more closely related to Rhinesuchus whaitsi than to Lydekkerina huxleyi or Peltobatrachus pustulatus. A similar alternate definition is that Rhinesuchidae is a stem-based clade containing all taxa more closely related to Uranocentrodon senekalensis than to Lydekkerina huxleyi, Trematosaurus brauni, or Mastodonsaurus giganteus.
Lydekkerina is an extinct genus of stereospondyl temnospondyl. It is the type genus of the family Lydekkerinidae. Fossils have been collected from Early Triassic deposits in South Africa and Australia. The type species is L. huxleyi, first described in 1889. While most other stereospondyls were semiaquatic, Lydekkerina was exclusively terrestrial.
Progalesaurus is an extinct genus of galesaurid cynodont from the early Triassic. Progalesaurus is known from a single fossil of the species Progalesaurus lootsbergensis, found in the Lystrosaurus Assemblage Zone of the Balfour Formation. Close relatives of Progalesaurus, other galesaurids, include Galesaurus and Cynosaurus. Galesaurids appeared just before the Permian-Triassic extinction event, and disappeared from the fossil record in the Middle-Triassic.
Platycraniellus is an extinct genus of carnivorous cynodonts from the Early Triassic. It is known from the Lystrosaurus Assemblage Zone of the Normandien Formation in South Africa. P. elegans is the only species in this genus based on the holotype specimen from the Ditsong National Museum of Natural History in Pretoria, South Africa. Due to limited fossil records for study, Platycraniellus has only been briefly described a handful of times.
Trirachodon is an extinct genus of cynodonts. Fossils have been found in the Cynognathus Assemblage Zone of the Beaufort Group in South Africa and the Omingonde Formation of Namibia, dating back to the Early and Middle Triassic.
Lumkuia is an extinct genus of cynodonts, fossils of which have been found in the Cynognathus Assemblage Zone of the Beaufort Group in the South African Karoo Basin that date back to the early Middle Triassic. It contains a single species, Lumkuia fuzzi, which was named in 2001 on the basis of the holotype specimen BP/1/2669, which can now be found at the Bernard Price Institute in Johannesburg, South Africa. The genus has been placed in its own family, Lumkuiidae. Lumkuia is not as common as other cynodonts from the same locality such as Diademodon and Trirachodon.
Abdalodon is an extinct genus of late Permian cynodonts, known by its only species A. diastematicus.Abdalodon together with the genus Charassognathus, form the clade Charassognathidae. This clade represents the earliest known cynodonts, and is the first known radiation of Permian cynodonts.
The Katberg Formation is a geological formation that is found in the Beaufort Group, a major geological group that forms part of the greater Karoo Supergroup in South Africa. The Katberg Formation is the lowermost geological formation of the Tarkastad Subgroup which contains the Lower to Middle Triassic-aged rocks of the Beaufort Group. Outcrops and exposures of the Katberg Formation are found east of 24 degrees on wards and north of Graaff-Reniet, Nieu Bethesda, Cradock, Fort Beaufort, Queensdown, and East London in the south, and ranges as far north as Harrismith in deposits that form a ring around the Drakensberg mountain ranges.
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.
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.