Tapinocephalus Assemblage Zone

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Tapinocephalus Assemblage Zone
Stratigraphic range: Middle Permian to Late Permian [1]
Tapinocephalus DB.jpg
Tapinocephalus atherstonei
Type Biozone
Unit of Abrahamskraal Formation of the Beaufort Group
Underlies Pristerognathus Assemblage Zone
Overlies Eodicynodon Assemblage Zone
Thicknessup to 6,561.68 feet (2,000 m)
Location
Region Northern, Western & Eastern Cape
CountryFlag of South Africa.svg  South Africa
Extent Karoo Basin
Type section
Named for Tapinocephalus atherstonei
Named byWatson (1914), Keyser & Smith (1977-78), Kitching (1984)

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. [2] [3] [4]

Contents

The name of the biozone refers to Tapinocephalus atherstonei , a large herbivorous tapinocephalid dinocephalian therapsid. It is characterised by the presence of this dinocephalian species along with the appearance of other advanced tapinocephalid dinocephalians, and the large pareiasaur Bradysaurus baini . It is also the first biozone of the series where the dicynodont, Diictodon feliceps, species first appear.

History

The first fossils to be found in the Beaufort Group rocks that encompass the current eight biozones were discovered by Andrew Geddes Bain in 1856. [5] However, it was not until 1892 that it was observed that the geological strata of the Beaufort Group could be differentiated based on their fossil taxa. The initial undertaking was done by Harry Govier Seeley who subdivided the Beaufort Group into three biozones, [6] which he named (from oldest to youngest):

These proposed biozones Seeley named were subdivided further by Robert Broom between 1906 and 1909. [7] Broom proposed the following biozones (from oldest to youngest):

The rocks of the current Tapinocephalus Assemblage Zone were first included with those of the lower Eodicynodon Assemblage Zone under the name "Pareiasaurus beds" by Broom. [8] [9] Years later Lieuwe Dirk Boonstra redefined the boundaries of the Tapinocephalus Assemblage Zone. As a young man Boonstra collaborated with Broom on research of dinocephalians. [10] After embarking on further study of dinocephalian fossils and their biostratigraphy, Boonstra defined the lower, middle, and upper sections of this biozone. [11] [12] [13] [14] In the 1970s, Keyser and Smith proposed the renaming of the biozone to Dinocephalian Assemblage Zone. In 1984 James Kitching proposed to name the biozone after Tapinocephalus, which was accepted over Keyser and Smith’s proposal. However, the zoning of the biozone rocks remains as they were defined by Keyser and Smith. [15] [16]

Lithology

The Tapinocephalus Assemblage Zone correlates with the Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group. Outcrops of this biozone are known from the south-western and central margins of the Abrahamskraal Formation where it conformably overlies the Eodicynodon Assemblage Zone in its south-western localities. In its northern and eastern localities it inter-fingers with Ecca Group-aged deposits. This biozone is considered to be Middle Permian (Guadalupian) in age. [17]

The rocks of the Tapinocephalus Assemblage Zone consist mainly of maroon to greyish red or purple mudstone layers which exhibit blocky weathering at exposed outcrops. The mudstones contain calcareous nodules and sheet limestones, both are indicative of a warm and seasonally arid climate, revealing the presence of paleocalcretes and carbonate precipitation respectively in playa lakes. Paleosols are also commonly found in the mudstones, which indicates a lack of deposition for long periods of time. In some deposits the mudstone layers contain thin chert lenses which have been attributed to silicified tuff deposits. Alternating beds of light grey to dark greenish grey siltstone and greenish grey to light olive grey sandstones which weather to light orange grey. The siltstones frequently contain both symmetrical and asymmetrical ripple surfaces which indicate that paleocurrents traveled downstream in a northerly direction. Desiccation cracks which are infilled by fine sandstone are also found. The sandstones are fine-grained and mainly tabular, indicating that deposition of these sandstones was in a low-energy fluvial environment. The sandstones are capped in the upper sections of the biozone with mudstone clast conglomerates. [18] [19]

The depositional environment of the Tapinocephalus Assemblage Zone was formed by sedimentary material being deposited in the Karoo Basin (a retro-arc foreland basin) by vast, low-energy alluvial plains flowing northwards from a southerly source area in the rising Gondwanide mountains. The Gondwanides were the result of crustal uplift that had previously begun to take course due to subduction of the Palaeo-pacific plate beneath the Gondwanan Plate. Orogenic pulses from the growing Gondwanides mountain chain and associated subduction created accommodation space for sedimentation in the Karoo Basin where the deposits of the Tapinocephalus Assemblage zone, and all other succeeding assemblage zone deposits, were deposited over millions of years. [20] [21]

Paleontology

Bradysaurus baini Bradysausus baini copia.jpg
Bradysaurus baini

The Tapinocephalus Assemblage Zone is characterized by the presence of Tapinocephalus atherstonei and Bradysaurus baini . Vertebrate fossils found in this biozone are not especially common, however are most commonly discovered as articulated single specimens within or associated with the calcareous nodules in the mudstone layers. [22] As the name for this biozone suggests, it is renowned by paleontologists for its diverse Dinocephalian fossil species where almost all members of this family – the Anteosauridae, Titanosuchidae, and Tapinocephalidae – are represented. [23] Example species from these families are Anteosaurus magnificus , Jonkeria boonstrai , and Tapinocephalus atherstonei respectively. Other notable dinocephalian species which have been discovered from this biozone are the advanced tapinocephalids Struthiocephalus whaitsi and Moschops capensis, and the unusual Styracocephalus platyrhynchus . [24] [25] Unfortunately dinocephalian fossils are frequently found disarticulated with much of their postcrania missing. [26]

Despite the rarity of fossils, the fossil taxa that have been found from the Tapinocephalus Assemblage Zone are extremely diverse. Parareptile species such as the pareiasaur, Bradysaurus, and the perplexing putative pantestudine Eunotosaurus africanus are confined to this biozone. [27] One of the few pelycosaur species found in South African deposits, Elliotsmithia longiceps , [28] and the biarmosuchian Hipposaurus boonstrai are likewise found, including a variety of basal gorgonopsid, [29] [30] anomodont, [31] [32] and therocephalian species. [33] The dicynodont Diictodon feliceps first appeared in this biozone and remained ubiquitous until the Permian-Triassic boundary. [34] Other dicynodont species found include Robertia broomiana and Pristerodon brachyops . Finally, fossils of temnospondyl amphibians such as Rhinesuchoides tenuiceps , [35] remains of the fish Namaichthys digitata, invertebrate fossils of molluscs, invertebrate trackways and burrows, vertebrate footprints of therapsids, and a variety of plant fossils, namely of Glossopteris , Dadoxylon , and Schizoneura , have been recovered.

Correlation

Many dinocephalian species that are found in the Tapinochephalus Assemblage Zone have been found in formations in different countries which correlate in age. As dinocephalian fossils are only known from Middle Permian (Guadalupian) deposits, dinocephalians are good biostratigraphic markers for this period. Dinocephalian fossils, along with other therapsid species found in the Tapinocephalus Assemblage Zone, have been recovered from the Rio do Rasto Formation from the Paraná Basin in Brazil [36] and from the Madumabisa Mudstone in Zambia. [37] [38]

See also

Related Research Articles

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

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.

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

Diictodon is an extinct genus of pylaecephalid dicynodont that lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the Cistecephalus Assemblage Zone of the Madumabisa Mudstone of the Luangwa Basin in Zambia and the Tropidostoma Assemblage Zone of the Teekloof Formation, Tapinocephalus Assemblage Zone of the Abrahamskraal Formation, Dicynodon Assemblage Zone of the Balfour Formation, Cistecephalus Assemblage Zone of the Middleton or Balfour Formation of South Africa and the Guodikeng Formation of China. Roughly half of all Permian vertebrate specimens found in South Africa are those of Diictodon. This small herbivorous animal was one of the most successful synapsids in the Permian period.

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

Jonkeria is an extinct genus of dinocephalians. Jonkeria was a large and omnivorous animal, from the Tapinocephalus Assemblage Zone, Lower Beaufort Group, of the South African Karoo.

<span class="mw-page-title-main">Beaufort Group</span> Third of the main subdivisions of the Karoo Supergroup in South Africa

The Beaufort Group is the third of the main subdivisions of the Karoo Supergroup in South Africa. It is composed of a lower Adelaide Subgroup and an upper Tarkastad Subgroup. It follows conformably after the Ecca Group and unconformably underlies the Stormberg Group. Based on stratigraphic position, lithostratigraphic and biostratigraphic correlations, palynological analyses, and other means of geological dating, the Beaufort Group rocks are considered to range between Middle Permian (Wordian) to Early Triassic (Anisian) in age.

<i>Anteosaurus</i> Extinct genus of anteosaurid synapsid from the Permian

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.

<i>Cistecephalus</i> Assemblage Zone

The Cistecephalus Assemblage Zone is a tetrapod assemblage zone or biozone found in the Adelaide Subgroup of the Beaufort Group, a majorly fossiliferous and geologically important geological group of the Karoo Supergroup in South Africa. This biozone has outcrops located in the Teekloof Formation north-west of Beaufort West in the Western Cape, in the upper Middleton and lower Balfour Formations respectively from Colesberg of the Northern Cape to east of Graaff-Reinet in the Eastern Cape. The Cistecephalus Assemblage Zone is one of eight biozones found in the Beaufort Group, and is considered to be Late Permian in age.

<i>Daptocephalus</i> Assemblage Zone

The Daptocephalus Assemblage Zone is a tetrapod assemblage zone or biozone found in the Adelaide Subgroup of the Beaufort Group, a majorly fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. This biozone has outcrops located in the upper Teekloof Formation west of 24°E, the majority of the Balfour Formation east of 24°E, and the Normandien Formation in the north. It has numerous localities which are spread out from Colesberg in the Northern Cape, Graaff-Reniet to Mthatha in the Eastern Cape, and from Bloemfontein to Harrismith in the Free State. The Daptocephalus Assemblage Zone is one of eight biozones found in the Beaufort Group and is considered Late Permian (Lopingian) in age. Its contact with the overlying Lystrosaurus Assemblage Zone marks the Permian-Triassic boundary.

<i>Lystrosaurus</i> Assemblage Zone

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.

<i>Cynognathus</i> Assemblage Zone Biozone which correlates to the Burgersdorp Formation of the Beaufort Group

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.

<i>Pristerognathus</i> Assemblage Zone

The Pristerognathus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Abrahamskraal Formation and lowermost 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 not more than 300 metres (980 ft), occur just east of Sutherland through to Beaufort West in the south and Victoria West in the north. Exposures are also found west of Colesberg and south of Graaff-Reinet. The Pristerognathus Assemblage Zone is the third biozone of the Beaufort Group.

<i>Tropidostoma</i> Assemblage Zone

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.

<i>Eodicynodon</i> Assemblage Zone

The Eodicynodon Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the 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 620 metres (2,030 ft), occur south-east of Sutherland, north of Prince Albert, and south-east of Beaufort West. The Eodicynodon Assemblage Zone is the lowermost biozone of the Beaufort Group.

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

Tapinocaninus is an extinct genus of therapsids in the family Tapinocephalidae, of which it is the most basal member. Only one species is known, Tapinocaninus pamelae. The species is named in honor of Rubidge's mother, Pam. Fossils have been found dating from the Middle Permian.

<i>Styracocephalus</i>

Styracocephalus platyrhynchus is an extinct genus of dinocephalian therapsid that existed during the mid-Permian throughout South Africa, but mainly in the Karoo Basin. It is often referred to by its single known species Styracocephalus platyrhynchus. The Dinocephalia clade consisted of the largest land vertebrates and herbivores during the early to mid-Permian. This period is often also referred to as the Guadalupian epoch, approximately 270 to 260 million years ago.

<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>Criocephalosaurus</i> Extinct genus of therapsids

Criocephalosaurus is an extinct genus of tapinocephalian therapsids that lived in Southern Africa during the Guadalupian epoch of the Permian. They are the latest surviving dinocephalians, extending past the Abrahamskraal Formation into the lowermost Poortjie Member of the Teekloof Formation in South Africa. They are also regarded as the most derived of the dinocephalians, alongside Tapinocephalus, and the most abundant in the fossil record.

<span class="mw-page-title-main">Abrahamskraal Formation</span> Geological formation of the Beaufort Group in South Africa

The Abrahamskraal Formation is a geological formation and is found in numerous localities in the Northern Cape, Western Cape, and the Eastern Cape of South Africa. It is the lowermost formation of the Adelaide Subgroup of the Beaufort Group, a major geological group that forms part of the greater Karoo Supergroup. It represents the first fully terrestrial geological deposits of the Karoo Basin. Outcrops of the Abrahamskraal Formation are found from the small town Middelpos in its westernmost localities, then around Sutherland, the Moordenaarskaroo north of Laingsburg, Williston, Fraserburg, Leeu-Gamka, Loxton, and Victoria West in the Western Cape and Northern Cape. In the Eastern Cape outcrops are known from Rietbron, north of Klipplaat and Grahamstown, and also southwest of East London.

<span class="mw-page-title-main">Middleton Formation</span> Late middle Permian geological formation in the Eastern Cape

The Middleton Formation is a geological formation that extends through the Northern Cape, Western Cape, and Eastern Cape provinces of South Africa. It overlies the lower Abrahamskraal Formation, and is the eastern correlate, East of 24ºE, of the Teekloof Formation. Outcrops and exposures of the Middleton Formation range from Graaff-Reinet in the Eastern Cape onwards. The Middleton Formation's type locality lies near the small hamlet, Middleton, approximately 25 km south of Cookhouse. Other exposures lie in hillsides along the Great Fish River in the Eastern Cape. The Middleton Formation forms part of the Adelaide Subgroup of the Beaufort Group, which itself forms part of the Karoo Supergroup.

<span class="mw-page-title-main">Teekloof Formation</span> Late Permian geological formation that forms part of the Beaufort Group of South Africa

The Teekloof Formation is a geological formation that forms part of the Beaufort Group, one of the five geological groups that comprises the Karoo Supergroup in South Africa. The Teekloof Formation is the uppermost formation of Adelaide Subgroup deposits West of 24ºE and contains Middle to Late Permian-aged deposits and four biozones of the Beaufort Group. It overlies the Abrahamskraal Formation. The Teekloof Formation does not underlie other units other than the younger Karoo dolerites and sills that relate to the emplacement of the Early Jurassic Drakensberg Group to the east. Outcrops and exposures of the Teekloof Formation range from Sutherland through the mountain escarpments between Fraserburg and Beaufort West. The northernmost localities of the Teekloof Formation are found by Loxton, Victoria West and Richmond.

Phorcys is an extinct genus of gorgonopsian that lived during the Middle Permian period (Guadalupian) of what is now South Africa. It is known from two specimens, both portions from the back of the skull, that were described and named in 2022 as a new genus and species P. dubei by Christian Kammerer and Bruce Rubidge. The generic name is from Phorcys of Greek mythology, the father of the Gorgons from which the gorgonopsians are named after, and refers to its status as one of the oldest representatives of the group in the fossil record. Phorcys was recovered from the lowest strata of the Tapinocephalus Assemblage Zone (AZ) of the Beaufort Group, making it one of the oldest known gorgonopsians in the fossil record—second only to fragmentary remains of an indeterminate gorgonopsian from the older underlying Eodicynodon Assemblage Zone.

References

  1. ↑ Rubidge, B.S.; Kitching, J.W. (2003). "A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa". Palaeontology. 46 (1): 199–210. doi: 10.1111/1475-4983.00294 .
  2. ↑ Rubidge, B. S. (ed.) 1995b. Biostratigraphy of the Beaufort Group (Karoo Supergroup). South African Committee of Stratigraphy. Biostratigraphic Series 1. Pretoria, Council for Geoscience.
  3. ↑ van der Walt, Merrill; Day, Michael; Rubidge, Bruce; Cooper, Antony; Netterberg, Inge (2010-12-31). "A new GIS-based biozone map of the Beaufort Group (Karoo Supergroup), South Africa". Palaeontologia Africana. 45: 1–6.
  4. ↑ Jirah, Sifelani; Rubidge, Bruce S. (2014-12-01). "Refined stratigraphy of the Middle Permian Abrahamskraal Formation (Beaufort Group) in the southern Karoo Basin". Journal of African Earth Sciences. 100: 121–135. doi:10.1016/j.jafrearsci.2014.06.014. ISSN   1464-343X.
  5. ↑ Bain, Andrew Geddes (1845-02-01). "On the Discovery of the Fossil Remains of Bidental and other Reptiles in South Africa". Quarterly Journal of the Geological Society. 1 (1): 317–318. doi:10.1144/GSL.JGS.1845.001.01.72. ISSN   0370-291X. S2CID   128602890.
  6. ↑ Seeley, H. G. (1895). "Researches on the Structure, Organization, and Classification of the Fossil Reptilia. Part IX., Section 4. On the Gomphodontia". Philosophical Transactions of the Royal Society of London B. 186: 1–57. doi: 10.1098/rstb.1895.0001 . JSTOR   91793.
  7. ↑ Broom, R. (January 1906). "V.—On the Permian and Triassic Faunas of South Africa". Geological Magazine. 3 (1): 29–30. doi:10.1017/S001675680012271X. ISSN   1469-5081. S2CID   129265956.
  8. ↑ Broom, R., 1906. V.—On the Permian and Triassic Faunas of South Africa. Geological Magazine, 3(1), pp.29-30.
  9. ↑ Broom, R., 1912, December. On some new fossil reptiles from the Permian and Triassic beds of South Africa. In Proceedings of the Zoological Society of London (Vol. 82, No. 4, pp. 859-876). Oxford, UK: Blackwell Publishing Ltd.
  10. ↑ Boonstra, L.D. and Broom, R., 1936. Some features of the cranial morphology of the tapinocephalid deinocephalians. Bulletin of the AMNH; v. 72, article 2.
  11. ↑ Boonstra, Lieuwe D. (1938-03-01). "A Report on some Karroo Reptiles from the Luangwa Valley, Northern Rhodesia". Quarterly Journal of the Geological Society. 94 (1–4): 371–384. doi:10.1144/GSL.JGS.1938.094.01-04.14. ISSN   0370-291X. S2CID   129931146.
  12. ↑ Boonstra, L.D. (1963-05-01). "Early dichotomies in the Therapsids". South African Journal of Science. 59 (5). ISSN   0038-2353.
  13. ↑ Boonstra, L.D., 1968. The braincase, basicranial axis and median septum in the Dinocephalia.
  14. ↑ Boonstra, L.D., 1969. The fauna of the Tapinocephalus Zone (Beaufort beds of the Karoo).
  15. ↑ Keyser, A.W. and Smith, R.M.H., 1978. Vertebrate biozonation of the Beaufort Group with special reference to the western Karoo Basin. Geological Survey, Department of Mineral And Energy Affairs, Republic of South Africa.
  16. ↑ Keyser, A.W., 1979. A review of the biostratigraphy of the Beaufort Group in the Karoo Basin of South Africa. Geocongress, Geological Society of South Africa, 2, pp.13-31.
  17. ↑ Day, Michael Oliver; Rubidge, Bruce Sidney (2014-12-01). "A brief lithostratigraphic review of the Abrahamskraal and Koonap formations of the Beaufort Group, South Africa: Towards a basin-wide stratigraphic scheme for the Middle Permian Karoo". Journal of African Earth Sciences. 100: 227–242. doi:10.1016/j.jafrearsci.2014.07.001. ISSN   1464-343X.
  18. ↑ Rubidge, B. S. (ed.) 1995b. Biostratigraphy of the Beaufort Group (Karoo Supergroup). South African Committee of Stratigraphy. Biostratigraphic Series 1. Pretoria, Council for Geoscience.
  19. ↑ Oliver, Day, Michael (2014-03-04). Middle Permian continental biodiversity changes as reflected in the Beaufort Group of South Africa: a bio- and lithostratigraphic review of the Eodicynodon, Tapinocephalus and Pristerognathus assemblage zones (Thesis).{{cite thesis}}: CS1 maint: multiple names: authors list (link)
  20. ↑ Hancox, P.J; Rubidge, B.S (2001-01-01). "Breakthroughs in the biodiversity, biogeography, biostratigraphy, and basin analysis of the Beaufort group". Journal of African Earth Sciences. 33 (3–4): 563–577. doi:10.1016/S0899-5362(01)00081-1. ISSN   1464-343X.
  21. ↑ Rubidge, Bruce S.; Day, Michael O.; Barbolini, Natasha; Hancox, P. John; Choiniere, Jonah N.; Bamford, Marion K.; Viglietti, Pia A.; McPhee, Blair W.; Jirah, Sifelani (2016), "Advances in Nonmarine Karoo Biostratigraphy: Significance for Understanding Basin Development", Origin and Evolution of the Cape Mountains and Karoo Basin, Springer International Publishing, pp. 141–149, doi:10.1007/978-3-319-40859-0_14, ISBN   9783319408583
  22. ↑ Canoville, A. and Chinsamy, A., 2017. Bone Microstructure of Pareiasaurs (Parareptilia) from the Karoo Basin, South Africa: Implications for Growth Strategies and Lifestyle Habits. The Anatomical Record, 300(6), pp.1039-1066.
  23. ↑ Atayman, S., Rubidge, B.S. and Abdala, F., 2009. Taxonomic re-evaluation of tapinocephalid dinocephalians. Palaeontologia africana, 44, p.88Á90. Link: https://core.ac.uk/download/pdf/39675877.pdf#page=88
  24. ↑ Rubidge, Bruce; VAN DEN HEEVER, J.A (1997-01-01). "Morphology and systematic position of the dinocephalian Styracocephalus platyrhynchus (Amniota: Therapsida)". Lethaia. 30: 157–168. doi:10.1111/j.1502-3931.1997.tb00457.x.
  25. ↑ Rubidge, Bruce S.; Sidor, Christian A. (2002-07-08). "On the cranial morphology of the basal therapsids Burnetia and Proburnetia (Therapsida: Burnetiidae)". Journal of Vertebrate Paleontology. 22 (2): 257–267. doi:10.1671/0272-4634(2002)022[0257:otcmot]2.0.co;2. ISSN   0272-4634. S2CID   86207308.
  26. ↑ Day, Michael O.; Güven, Saniye; Abdala, Fernando; Jirah, Sifelani; Rubidge, Bruce; Almond, John (April 2015). "Youngest dinocephalian fossils extend the Tapinocephalus Zone, Karoo Basin, South Africa". South African Journal of Science. 111 (3–4): 1–5. doi: 10.17159/sajs.2015/20140309 . ISSN   0038-2353.
  27. ↑ Lyson, Tyler R.; Bever, Gabe S.; Scheyer, Torsten M.; Hsiang, Allison Y.; Gauthier, Jacques A. (2013-06-17). "Evolutionary Origin of the Turtle Shell". Current Biology. 23 (12): 1113–1119. doi: 10.1016/j.cub.2013.05.003 . ISSN   0960-9822. PMID   23727095.
  28. ↑ Reisz, Robert R.; Dilkes, David W.; Berman, David S (1998-09-15). "Anatomy and relationships ofElliotsmithia longicepsBroom, a small synapsid (Eupelycosauria: Varanopseidae) from the late Permian of South Africa". Journal of Vertebrate Paleontology. 18 (3): 602–611. doi:10.1080/02724634.1998.10011087. ISSN   0272-4634.
  29. ↑ Kammerer, Christian F. (2013-09-21), "A Redescription of Eriphostoma microdon Broom, 1911 (Therapsida, Gorgonopsia) from the Tapinocephalus Assemblage Zone of South Africa and a Review of Middle Permian Gorgonopsians", Early Evolutionary History of the Synapsida, Vertebrate Paleobiology and Paleoanthropology, Springer Netherlands, pp. 171–184, doi:10.1007/978-94-007-6841-3_11, ISBN   9789400768406
  30. ↑ Kammerer, C.F., Smith, R.M., Day, M.O. and Rubidge, B.S., 2015. New information on the morphology and stratigraphic range of the mid‐Permian gorgonopsian E riphostoma microdon Broom, 1911. Papers in Palaeontology, 1(2), pp.201-221.
  31. ↑ Cisneros, Juan Carlos; Abdala, Fernando; Jashashvili, Tea; de Oliveira Bueno, Ana; Dentzien-Dias, Paula (July 2015). "Tiarajudens eccentricus and Anomocephalus africanus, two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana". Royal Society Open Science. 2 (7): 150090. doi:10.1098/rsos.150090. ISSN   2054-5703. PMC   4632579 . PMID   26587266.
  32. ↑ Angielczyk, Kenneth D.; Rubidge, Bruce S.; Day, Michael O.; Lin, Florence (2016-02-06). "A reevaluation ofBrachyprosopus broomiandChelydontops altidentalis, dicynodonts (Therapsida, Anomodontia) from the middle PermianTapinocephalusAssemblage Zone of the Karoo Basin, South Africa". Journal of Vertebrate Paleontology. 36 (2): e1078342. doi:10.1080/02724634.2016.1078342. ISSN   0272-4634. S2CID   130520407.
  33. ↑ Boonstra, L.D., 1965. The girdles and limbs of the Gorgonopsia of the Tapinocephalus Zone. Annals of the South African Museum, 48(13), pp.237-249.
  34. ↑ ANGIELCZYK, KENNETH D.; RUBIDGE, BRUCE S. (September 2010). "A new pylaecephalid dicynodont (Therapsida, Anomodontia) from the Tapinocephalus Assemblage Zone, Karoo Basin, Middle Permian of South Africa". Journal of Vertebrate Paleontology. 30 (5): 1396–1409. doi:10.1080/02724634.2010.501447. ISSN   0272-4634. S2CID   129846697.
  35. ↑ Damiani, Ross J. (2004-01-01). "Temnospondyls from the Beaufort Group (Karoo Basin) of South Africa and Their Biostratigraphy". Gondwana Research. 7 (1): 165–173. doi:10.1016/S1342-937X(05)70315-4. ISSN   1342-937X.
  36. ↑ Boos, A.D.S.; Kammerer, C.F.; Schultz, C.L.; Paes Neto, V.D. (2015-11-01). "A tapinocephalid dinocephalian (Synapsida, Therapsida) from the Rio do Rasto Formation (Paraná Basin, Brazil): Taxonomic, ontogenetic and biostratigraphic considerations". Journal of South American Earth Sciences. 63: 375–384. doi:10.1016/j.jsames.2015.09.003. ISSN   0895-9811.
  37. ↑ Sidor, Christian A.; Angielczyk, Kenneth D.; Smith, Roger M. H.; Goulding, Adam K.; Nesbitt, Sterling J.; Peecook, Brandon R.; Steyer, J. Sébastien; Tolan, Stephen (2014-06-07). "Tapinocephalids (Therapsida, Dinocephalia) from the Permian Madumabisa Mudstone Formation (Lower Karoo, Mid-Zambezi Basin) of southern Zambia". Journal of Vertebrate Paleontology. 34 (4): 980–986. doi:10.1080/02724634.2013.826669. ISSN   0272-4634. S2CID   128431441.
  38. ↑ Sidor, Christian (2015-04-09). "The first biarmosuchian from the upper Madumabisa Mudstone Formation (Luangwa Basin) of Zambia". Palaeontologia Africana. 49: 1–7.
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