Pampaphoneus

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Pampaphoneus
Temporal range: Guadalupian (Wordian), 268–265  Ma
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Pampaphoneus holotype skull.png
Pampaphoneus biccai holotype skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Suborder:† Dinocephalia
Family:† Anteosauridae
Subfamily:† Syodontinae
Genus:† Pampaphoneus
Cisneros et al., 2012
Type species
†Pampaphoneus biccai
Cisneros et al., 2012

Pampaphoneus is an extinct genus of carnivorous dinocephalian therapsid belonging to the family Anteosauridae. It lived 268 to 265 million years ago during the Wordian age of the Guadalupian (= middle Permian) period in what is now Brazil. Pampaphoneus is known by an almost complete skull with the lower jaw still articulated, discovered on the lands of the Boqueirão Farm, near the city of São Gabriel, in the state of Rio Grande do Sul. A second specimen from the same locality was reported in 2019 and 2020 but has not yet been described. It is composed of a skull associated with postcranial remains. It is the first South American species of dinocephalian to have been described. The group was previously known in South America only by a few isolated teeth and a jaw fragment reported in 2000 in the same region of Brazil. Phylogenetic analysis conducted by Cisneros and colleagues reveals that Pampaphoneus is closely related to anteosaurs from European Russia, indicating a closer faunal relationship between South America and Eastern Europe than previously thought, thus promoting a Pangea B continental reconstruction.

Contents

Etymology

The name of the genus comes from the Pampas, a region of vast plains typical of southern South America, from which the specimen originates, and from the Greek phoneus, meaning "killer", in reference to the predatory habits of the animal. The specific epithet honors José Bicca, the landowner of the farm where the fossil was found. [1]

Description

Reconstruction of the skull of Pampaphoneus biccai by Juan Carlos Cisneros. Pampaphoneus skull reconstruction.jpg
Reconstruction of the skull of Pampaphoneus biccai by Juan Carlos Cisneros.

The holotype (UFRGS PV386P) of Pampaphoneus is a skull measuring approximately 32 cm in length. As with all anteosaurs, the ventral margin of the premaxilla is inclined upwards and the postorbital bar is strongly curved anteroventrally so that the temporal fenestra undercuts orbit. The skull roof is slightly pachyostosed. The temporal fenestrae open widely on the skull roof where the insertion zone of the adductor musculature of the mandible extends to the pineal boss. The frontals contribute to the insertion zone of this musculature and also participate to the anterior edge of the pineal boss. These characteristics indicate that Pampaphoneus belongs to the clade Syodontinae. Within this group it is with the Russian species Syodon biarmicum that Pampaphoneus shares the most similarities. Like the latter, it possesses strongly recurved hook-like canines and very low postcanines with serrassions. The canines of the holotype are 7 cm in length. Pampaphoneus however differs from Syodon by its larger size, more robust snout, thickened postorbital forming a supraorbital boss, and a well-developed ridge extending from the pineal boss to the orbital rim. It also differs from other anteosaurs by its squamosal with a jugal process extending beyond the anterior edge of the temporal fenestra. [1] A peculiarity of the holotype that was thought to be a diagnostic feature is the presence of only four teeth per premaxilla (the fourth incisor, small in size, being however laterally masked by the maxilla) and eight postcanines. [1] However, a second skull (yet undescribed) was reported showing some differences with the holotype such as the presence of five premaxillary teeth and only seven postcanines. Since the other cranial elements are very similar, these differences are interpreted as intraspecific variation. [2] [3] Although being a Syodontinae, Pampaphoneus also shares several characters with the more derived Anteosaurinae. It thus has a well-developed medial crest on the skull roof, while a pronounced thickening of each postorbital forms a supraorbital boss similar to that of a subadult individual of the Russian Titanophoneus potens . In addition, the angular bone of the lower jaw bears a boss as in Anteosaurus and the two species of Titanophoneus. This angular boss is however much less developed in Pampaphoneus. [1]

Paleoecology

Life restoration of Pampaphoneus biccai by Juan Carlos Cisneros. Pampaphoneus life restoration.png
Life restoration of Pampaphoneus biccai by Juan Carlos Cisneros.

Remains of Pampaphoneus comes from the Morro Pelado Member of the Rio do Rasto Formation, outcropping on the lands of the Boqueirão farm in the municipality of São Gabriel (district of Catuçaba, State of Rio Grande do Sul). The fossils are preserved there in a fine pinkish sandstone and are covered with a dark crust of iron oxide. [4] Sediments of the Morro Pelado Member were deposited in fluvio-lacustrine and deltaic settings suggesting alluvial conditions with coalescing floodplains. Aeolian facies (fossil dunes) present in the upper part of the Morro Pelado Member attest to a progressive aridification of the environment. Palaeogeographically, southeastern Brazil was located at mid-latitudes between the 35th and 45th parallel south, where a warm temperate climate probably prevailed with a prolonged dry season. [5] The flora included forests of Glossopteris communis which occupied the floodplains and the overbank channels while the more humid biotopes were covered with dense mats of equisetids including equisetales ( Schizoneura and Phyllotheca ) and sphenophyllales ( Sphenophyllum paranaense ). [5] Besides Pampaphoneus, the Boqueirão farm site also contains the temnospondyl Konzhukovia sangabrielensis [6] and the small dicynodont Rastodon . [4] Other localities in the Morro Pelado Member have also yielded tetrapod remains. In Rio Grande do Sul, several sites around Aceguá in the district of Minuano have yielded the most complete fossils of the pareiasaur Provelosaurus . [7] The Fagundes farm locality (São Gabriel, Catuçaba district) yielded a fragmentary skull of Provelosaurus, [8] teeth of undetermined Anteosaurid, Titanosuchid and Tapinocephalid dinocephalians, [9] and an undetermined amphibian. [10] In the district of Tiarajú (also near São Gabriel) was found the basal anomodont Tiarajudens [11] [12] From the Serra do Cadeado area (near Ortigueira, State of Paraná), come the amphibian Australerpeton , [10] an undetermined species of the dicynodont Endothiodon , [13] (possibly E. bathystoma [14] ) as well as an indeterminate tapinocephalid dinocephalian showing similarities with the genera Moschops and Moschognathus . [10] However, there are uncertainties about the contemporaneity of all these sites. Vertebrate fossils from the Rio do Rasto Formation occur in scattered, isolated, and discontinuous outcrops due to dense vegetation cover, making it difficult to establish local correlations. The precise location of several ancient discoveries is uncertain and several taxa could come from different stratigraphic levels within the Rio do Rasto Formation. This is the case of the fossils of Paraná, where tapinocephalid and Endothiodon remains could come from two distinct levels. [10]

Age of the Rio do Rasto Formation

Estimating the age of the Rio do Rasto formation has long relied solely on the basis of biostratigraphic correlations. Since 2018, radiometric dating has made it possible to specify the age of part of the formation. [15] [16] Based on conchostracan and bivalve faunas, Holz and colleagues have suggested that the sedimentary succession of this formation extended from the Wordian (middle Guadalupian) to the Wuchiapingian (base of the Lopingian). [17] Among the tetrapods, the presence of dinocephalians in Fagundes and Boqueirão sites (State of Rio Grande do Sul) as well as in the region of Serra do Cadeado (State of Paraná) indicates a Guadalupian age for these localities, given that this synapsid group is restricted to this age in the rest of the world. In the Fagundes farm site, dinocephalians coexist with the pareiasaur Provelosaurus. The latter is present in the Aceguà area about 2 m above a 30 cm thick layer of bentonite radiometrically dated at 266 ± 5.4 million years. [16] [7] The broad error margin of this radiometric dating places the Brazilian dinocephalian sites in the Lower Roadian – Middle Capitanian time interval, confirming the Guadalupian age of this part of the Morro Pelado Member. [7] The other tetrapods of the Morro Pelado Member in Rio Grande do Sul suggest an age not exceeding Wordian: Tapinocephalids are only known from Wordian and Capitanian rocks, the basal anomodont Tiarajudens is closely related to the South African genus Anomocephalus of the Tapinocephalus Assemblage Zone whose age extends from the late Wordian to the late Capitanian, and the temnospondyl Konzhukovia , present at the Boqueirão farm site in São Gabriel, is also known in eastern European Russia in strata limited to the Wordian-Capitanian interval. [7] The probable Wordian age of the Morro Pelado Member is also reinforced by a radiometric age of the Serrinha Member (lower part of the Rio do Rasto formation) obtained from a volcanic ash layer which gave an age of 270.61 + 1.76/− 3.27 Ma corresponding to the Roadian (early Guadalupian). [15]

Classification

In describing Pampaphoneus, Cisneros et al. presented several cladograms confirming the recognition of the clades Anteosaurinae and Syodontinae erected a year earlier by Christian Kammerer. In all their analyzes, Pampaphoneus is identified as the most basal Syodontinae: [1]

Therapsida

Biarmosuchus tener

Dinocephalia

Estemmenosuchus uralensis

Ulemosaurus svijagensis

Tapinocaninus pamelae

Anteosauridae

Archaeosyodon praeventor

Anteosaurinae

Sinophoneus yumenensis

Titanophoneus adamanteus

Titanophoneus potens

Anteosaurus magnificus

Syodontinae

Pampaphoneus biccai

Notosyodon gusevi

Syodon biarmicum

Australosyodon nyaphuli

Palaeobiogeography

During the Permian, most of the landmasses were united into a single supercontinent, Pangea, which was roughly C-shaped. Its northern (Laurasia) and southern (Gondwana) parts were connected to the west but separated to the east by a very large oceanic bay - the Tethys. [18] A long string of microcontinents, grouped under the name of Cimmeria, divided the Tethys in two: the Paleo-Tethys in the north, and the Neo-Tethys in the south. [19] In the northern hemisphere, anteosaurs are known from eastern European Russia, Kazakhstan, and China, territories corresponding to mid-paleolatitudes between the 30th and 40th parallel north. [20] In the southern hemisphere, anteosaur remains are known in South Africa, Zambia and Zimbabwe, territories which were all located at high paleolatitudes located at the level of the 60th parallel south. [20] [21] The Brazilian localities were located a little further north, between the 35th and the 45th parallel south. [5] The close relationship of the Brazilian Pampaphoneus with some South African and Russian anteosaurs suggests dispersal through western Pangea rather than eastern Pangea via the Cathaysian Bridge. The latter included parts of northern and southern China, Korea, and Indochina and played an important role in the dispersal of therapsids in the late Permian and Triassic. However, this bridge was probably not yet formed in the Middle Permian and marine barriers broke up the different Cathaysian blocks, making it difficult for animals to disperse from Cathaysia to Southern Africa via Eastern Europe. Cisneros and colleagues suggest a seemingly easier migration route via western Pangea by favoring a Pangea B-type continental reconstruction where South America was juxtaposed with the Appalachia and was thus closer to Russia than in the Pangea A reconstruction. In the latter, the Appalachian and Mauritanides mountains would have been difficult natural barriers to cross. In contrast, in a Pangea B configuration, Brazil was not only closer to Eastern Europe, but the only mountain barrier along the way was the moderately high European Hercynides. Anteosaurs would have migrated from Russia (where the oldest specimens come from) to southern Africa, passing through eastern Europe and western Africa (bypassing the Mauritanides chain) and then through Brazil. [1] The discovery of probable anteosaur footprints in southern France, [N 1] which was then located at low paleolatitude, at the level of the 10th parallel north, also supports this hypothesis, the south-western Europe being in this migration corridor. [22] [23] [24] [25]

Notes

  1. ↑ These footprints, initially named Planipes caudatus and P. brachydactylus, are now grouped under the name Brontopus antecursor (cf. references Marchetti et al. 2019).

Related Research Articles

The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0.5 – 259.1 ± 0.4 Mya. The series saw the rise of the therapsids, a minor extinction event called Olson's Extinction and a significant mass extinction called the end-Capitanian extinction event.

<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.

<span class="mw-page-title-main">Anteosaur</span> Extinct clade of therapsids

Anteosaurs are a group of large, primitive carnivorous dinocephalian therapsids with large canines and incisors and short limbs, that are known from the Middle Permian of South Africa, Russia, China, and Brazil. Some grew very large, with skulls 50–80 centimetres (20–31 in) long, and were the largest predators of their time. They died out at the end of the Middle Permian, possibly as a result of the extinction of the herbivorous Tapinocephalia on which they may have fed.

<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. With its skull reaching 80–90 cm (31–35 in) in length and a body size estimated at more than 5 m (16 ft) in length, and 500 to 600 kg in weight, 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>Tapinocephalus</i> Assemblage Zone

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.

<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>Endothiodon</i> Extinct genus of dicynodonts

Endothiodon is an extinct genus of large dicynodont from the Late Permian. Like other dicynodonts, Endothiodon was an herbivore, but it lacked the two tusks that characterized most other dicynodonts. The anterior portion of the upper and lower jaw are curved upward, creating a distinct beak that is thought to have allowed them to be specialized grazers.

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

Anomocephalus is an extinct genus of primitive anomodonts and belongs to the clade Anomocephaloidea. The name is said to be derived from the Greek word anomos meaning lawless and cephalos meaning head. The proper word for head in Greek is however κεφαλή (kephalē). It is primitive in that it retains a complete set of teeth in both jaws, in contrast to its descendants, the dicynodonts, whose dentition is reduced to only a single pair of tusks, with their jaws covered by a horny beak similar to that of a modern tortoise. However, they are in no way closely related.

<i>Styracocephalus</i>

Styracocephalus platyrhynchus (Greek for “spiked-head”) 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>Konzhukovia</i> Genus of amphibians (fossil)

Konzhukovia is an amphibian genus that belongs to an extinct group of temnospondyls, the largest clade of basal tetrapods including about 198 genera, 292 species, and more than half of which were alive during the early Mesozoic period. The animal was a predator that lived about 260 million years ago, and could get up to about three meters in length. Specifically, Konzukovia lived during the Permian, between 252 and 270 million years ago according to the type of rock the fossil was found in. There are three species within this genus, K. vetusta, K. tarda, and K. sangabrielensis, the first two originating from Russia while the latest originating from Southern Brazil. The discovery of this specimen in Southern Brazil provided more evidence to support the idea that during this animals existence, there was a “biological corridor” because of the supercontinent Pangea, allowing these species to be found so far apart from each other. Konzhukovia belongs to the family Archegosauridae, a family consisted of large temnospondyls that most likely compare to modern day crocodiles. Since the discovery of the latest species, K. sangabrielensis, Pacheco proposes that there must be the creation of a new family, Konzhokoviidae, a monophyletic group in a sister-group relationship with Stereospondlyi in order to accommodate the three species. Konzhukovia skulls usually exhibit typical rhinesuchid features including an overall parabolic shape, small orbits located more posteriorly, and the pterygoids do not reach the vomer. These animals were long-snouted amphibians that had clear adaptations made for fish catching, as well as exemplifying aquatic features.

<i>Moschognathus</i> Extinct genus of therapsids that lived in the Guadalupian epoch

Moschognathus is an extinct genus of dinocephalian therapsid in the family Tapinocephalidae. The genus includes only the type species M. whaitsi, named by palaeontologist Robert Broom in 1914. It was a short-snouted tapinocephalid, closely related to and resembling the well-known genus Moschops, but its skull is less thickened overall has a relatively longer and shallower snout by comparison. Indeed, Moschognathus has typically been regarded as a junior synonym of Moschops since 1969 after Lieuwe Dirk Boonstra sunk Moschognathus into Moschops, albeit retained as its own doubtfully valid species. However, researchers in the 21st century have expressed doubt over this synonymy and suggested that Moschognathus is a distinct taxon after all, including first by Christian Kammerer in a 2009 Ph.D. thesis and formally in 2015 by Alessandra D. S. Boos and colleagues in 2015. Moschognathus has since began to re-enter scientific literature of dinocephalians as a valid name and treated distinct from Moschops.

<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.

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

Tiarajudens is an extinct genus of saber-toothed herbivorous anomodonts which lived during the Middle Permian period in what is now Rio Grande do Sul, Brazil. It is known from the holotype UFRGS PV393P, a nearly complete skull. The type species T. eccentricus was named in 2011.

<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">Syodontinae</span> Extinct subfamily of mammal ancestors

Syodontinae is a group of dinocephalian therapsids. It is one of two subfamilies in the family Anteosauridae, the other being Anteosaurinae. They are known from the Middle Permian Period of what is now Russia and South Africa. One of the best known syodontines is Syodon from Russia. The South African form Australosyodon, is one of the earliest known Gondwanan anteosaurs.

<span class="mw-page-title-main">Anteosaurinae</span> Extinct subfamily of therapsids

Anteosaurinae is an extinct subfamily of dinocephalian therapsids. It is one of two subfamilies in the family Anteosauridae, the other being Syodontinae.

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

Sinophoneus is an extinct genus of carnivorous dinocephalian therapsid belonging to the family Anteosauridae. It lived 272 to 270 million years ago at the beginning of the Middle Permian in what is now the Gansu Province in northern China. It is known by a skull of an adult individual, as well as by many skulls of juvenile specimens. The latter were first considered as belonging to a different animal, named Stenocybus, before being reinterpreted as immature Sinophoneus. Sinophoneus shows a combination of characters present in other anteosaurs. Its bulbous profile snout and external nostrils located in front of the canine are reminiscent of the basal anteosaur Archaeosyodon, while its massive transverse pterygoids processes with enlarged distal ends are more similar to the more derived anteosaurs Anteosaurus and Titanophoneus. First phylogenetic analyzes identified Sinophoneus as the most basal Anteosaurinae. A more recent analysis positioned it outside the Anteosaurinae and Syodontinae subclades, and recovers it as the most basal Anteosauridae.

<span class="mw-page-title-main">Rio do Rasto Formation</span> Geological formation in Brazil

The Rio do Rasto Formation is a Late Permian sedimentary geological formation in the South Region of Brazil. The official name is Rio do Rasto, although in some publications it appears as Rio do Rastro.

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

Rastodon is an extinct genus of dicynodont. Uniquely among dicynodonts, its tusks curve forward. The type and only species is R. procurvidens.

Rastosuchus is an extinct genus of stereospondyl temnospondyl within the family Rhinesuchidae. It contains one species, Rastosuchus hammeri, found in the Permian Rio do Rasto Formation of Brazil.

References

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  2. ↑ Anilson Costa Santos, M.; Bulsing, K.P.; Santos, A.T.; Ferraz, J.S.; Oliveira, N.; Protti, S.W.L.; Dos Santos Rodriguez, L.A.; Xavier, P.L.; De Queiroz, L.C.; Cisneros, J.C.; Pinheiro, F.L. (2019). "Um Esqueleto Parcial do Dinocefálio Anteossaurídea Pampaphoneus biccai (Permiano do Rio Grande do Sul)". Anais do XXVI Congresso Brasileiro de Paleontologia. Campinas: Galoá. 2: 270–271.
  3. ↑ Perceval Paz, C.; Anilson Costa Santos, M.; Lima Pinheiro, F. (2020). "O Primeiro Registro de Pampaphoneus biccai (Anteosauridae) Com Ossos Pós-Cranianos". Revista Multidisciplinar de Educação e Meio Ambiente. 1 (1): 133.
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  6. ↑ Pacheco, C.P.; Eltink, E.; Müller, R.T.; Dias-da-Silva, S.A. (2016). "A new Permian temnospondyl with Russian affinities from South America, the new family Konzhukoviidae, and the phylogenetic status of Archegosauroidea". Journal of Systematic Palaeontology. 15 (3): 241–256. doi:10.1080/14772019.2016.1164763. S2CID   87860271.
  7. 1 2 3 4 Cisneros, J.C.; Dentzien-Dias, P.; Francischini, H. (2021). "The Brazilian Pareiasaur revisited". Frontiers in Ecology and Evolution. 9: 758802. doi: 10.3389/fevo.2021.758802 .
  8. ↑ Cisneros, J.C.; Abdala, F.; Malabarba, M.C. (2005). "Pareiasaurids from the Rio do Rasto Formation, southern Brazil: Biostratigraphic implications for Permian faunas of the Paraná Basin". Revista Brasileira de Paleontologia. 8 (1): 13–24. doi: 10.4072/rbp.2005.1.02 .
  9. ↑ Langer, M.C. (2000). "The first record of dinocephalians in South America: Late Permian (Rio do Rasto Formation) of the Paraná Basin, Brazil". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 215: 69–95. doi:10.1127/njgpa/215/2000/69.
  10. 1 2 3 4 Boos, A.D.S.; Kammerer, C.F.; Schultz, C.L.; Paes Neto, V.D. (2015). "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. Bibcode:2015JSAES..63..375B. doi:10.1016/j.jsames.2015.09.003.
  11. ↑ Cisneros, J.C.; Abdala, F.; Rubidge, B.S.; Dentzien-Dias, P.C.; Bueno, A.O. (2011). "Dental Occlusion in a 260-Million-Year-Old Therapsid with Saber Canines from the Permian of Brazil". Science. 331 (6024): 1603–1605. Bibcode:2011Sci...331.1603C. doi:10.1126/science.1200305. PMID   21436452. S2CID   8178585.
  12. ↑ Cisneros, J.C.; Abdala, F.; Jashashvili, T.; Bueno, A.O.; Dentzien-Dias, P. (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): 1–27. Bibcode:2015RSOS....250090C. doi:10.1098/rsos.150090. PMC   4632579 . PMID   26587266.
  13. ↑ Boos, A.D.S.; Schultz, C.L.; Vega, C.S.; Aumond, J.J. (2013). "On the presence of the Late Permian dicynodont Endothiodon in Brazil". Palaeontology. 56 (4): 837–848. doi:10.1111/pala.12020. S2CID   129679835.
  14. ↑ Kammerer, C.F.; Ordoñez, M.A. (2021). "Dicynodonts (Therapsida: Anomodontia) of South America". Journal of South American Earth Sciences. 108: 103171. Bibcode:2021JSAES.10803171K. doi:10.1016/j.jsames.2021.103171. S2CID   233565963.
  15. 1 2 Francischini, H.; Dentzien-Dias, P.; Guerra-Sommer, M.; Menegat, R.; Santos, J.O.S.; Manfroi, J.; Schultz, C.L. (2018). "A Middle Permian (Roadian) lungfish aestivation burrow from the Rio do Rasto Formation (Paraná Basin, Brazil) and associated U-Pb dating". PALAIOS. 33 (2): 69–84. Bibcode:2018Palai..33...69F. doi:10.2110/palo.2017.050. S2CID   134435628.
  16. 1 2 Rocha-Campos, A.C.; Basei, M.A.S.; Nutman, A.P.; Santos, P.R.; Passarelli, C.R .; Canile, F.M.; Rosa, O.C.R.; Fernandes, M.T.; Santa Ana, H.; Veroslavsky, G. (2019). "U-Pb zircon dating of ash fall deposits from the Paleozoic Paraná Basin of Brazil and Uruguay: a reevaluation of the stratigraphic correlations". The Journal of Geology. 127 (2): 167–182. Bibcode:2019JG....127..167R. doi:10.1086/701254. hdl: 1885/202604 . S2CID   134671451.
  17. ↑ Holz, M.; França, A.B.; Souza, P.A.; Iannuzzi, R.; Rohn, R. (2010). "A stratigraphic chart of the Late Carboniferous/Permian succession of the eastern border of the Paraná Basin, Brazil, South America". Journal of South American Earth Sciences. 29 (2): 381–399. Bibcode:2010JSAES..29..381H. doi:10.1016/j.jsames.2009.04.004.
  18. ↑ McLoughlin, S. (2001). "The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism". Australian Journal of Botany. 49 (3): 271–300. doi:10.1071/BT00023.
  19. ↑ Şengör, A.M.C. (1987). "Tectonics of the Tethysides: orogenic collage development in a collisional setting". Annual Review of Earth and Planetary Sciences. 15: 214–244. Bibcode:1987AREPS..15..213C. doi:10.1146/annurev.ea.15.050187.001241.
  20. 1 2 Kemp, T.S. (2006). "The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis". Journal of Evolutionary Biology. 19 (4): 1231–1247. doi: 10.1111/j.1420-9101.2005.01076.x . PMID   16780524. S2CID   3184629.
  21. ↑ Bamford, M.K. (2016). "Fossil woods from the Upper Carboniferous to Lower Jurassic Karoo Basin and their environmental interpretation". In Linol, B.; de Wit, M.J. (eds.). Origin and evolution of the Cape Mountains and Karoo Basin. Cham: Regional Geology Reviews, Springer. pp. 159–167. ISBN   978-3-319-40858-3.
  22. ↑ Gand, G.; Demathieu, G.; Ballestra, F. (1995). "La palichnofaune de vertébrés tétrapodes du Permien supérieur de l'Esterel (Provence, France)" [The palichnofauna of tetrapod vertebrates from the Upper Permian of Esterel (Provence, France)]. Palaeontographica Abteilung A. 4–6 (4–6): 97–139. doi:10.1127/pala/235/1995/97. S2CID   247392880.
  23. ↑ Gand, G.; Garric, J.; Demathieu, G.; Ellenberger, P. (2000). "La palichnofaune de vertébrés tétrapodes du Permien supérieur du bassin de Lodève (Languedoc-France)". Palaeovertebrata. 29 (1): 1–82.
  24. ↑ Marchetti, L.; Klein, H.; Buchwitz, M.; Ronchi, A.; Smith, R.M.H.; De Klerk, W.J.; Sciscio, L.; Groenewald, G.H. (2019). "Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises". Gondwana Research. 72: 139–168. Bibcode:2019GondR..72..139M. doi: 10.1016/j.gr.2019.03.009 . S2CID   133781923.
  25. ↑ Olroyd, S.L.; Sidor, C.A. (2017). "A review of the Guadalupian (middle Permian) global tetrapod fossil record". Earth-Science Reviews. 171: 583–597. Bibcode:2017ESRv..171..583O. doi: 10.1016/j.earscirev.2017.07.001 .
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