Anteosaurus: Difference between revisions

Anteosaurus; Temporal range: Capitanian, ; ~265–260 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	A. magnificus skull (SAM-PK-11296) on display at the Iziko Museum in Cape Town, South Africa
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Synapsida
Clade:	Therapsida
Suborder:	†Dinocephalia
Family:	†Anteosauridae
Subfamily:	†Anteosaurinae
Clade:	†Anteosaurini
Genus:	†Anteosaurus; Watson, 1921
Species:	†A. magnificus
Binomial name
	†Anteosaurus magnificus; Watson, 1921
Synonyms
	Genus synonymy Eccasaurus (?); Broom, 1909 ; Titanognathus ; Broili & Schröder, 1935 ; Dinosuchus ; Broom, 1936 ; Broomosuchus ; Camp, 1942 ; Micranteosaurus ; Boonstra, 1954 ; Paranteosaurus ; Boonstra, 1954 ; Pseudanteosaurus ; Boonstra, 1954 ; Species synonymy Eccasaurus priscus (?); Broom, 1909 ; Anteosaurus minor ; Broom, 1929 ; Titanognathus lotzi ; Broili & Schröder, 1935 ; Dinosuchus vorsteri ; Broom, 1936 ; Broomosuchus vorsteri ; Camp, 1942 ; Anteosaurus abeli ; Boonstra, 1952 ; Anteosaurus vorsteri ; Boonstra, 1953 ; Titanosuchus lotzi ; Boonstra, 1953 ; Anteosuchus acutirostris ; Boonstra, 1954 ; Anteosaurus crassifrons ; Boonstra, 1954 ; Anteosaurus cruentus ; Boonstra, 1954 ; Anteosaurus laticeps ; Boonstra, 1954 ; Anteosaurus levops ; Boonstra, 1954 ; Anteosaurus lotzi ; Boonstra, 1954 ; Anteosaurus major ; Boonstra, 1954 ; Anteosaurus minusculus ; Boonstra, 1954 ; Paranteosaurus primus ; Boonstra, 1954 ; Pseudanteosaurus minor ; Boonstra, 1954 ; Micranteosaurus parvus ; Boonstra, 1954 ;

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Anteosaurus (meaning "Antaeus lizard") is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian (= Middle Permian) during the Capitanian age, 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.

As in many other dinocephalians the cranial bones of Anteosaurus are pachyostosed, but to a lesser extent than in tapinocephalid dinocephalians. In Anteosaurus, pachyostosis mainly occurs in the form of horn-shaped supraorbital protuberances. According to some paleontologists this structure would be implicated in intraspecific agonistic behaviour, including head-pushing probably during the mating season. On the contrary, other scientists believe that this pachyostosis served to reduce cranial stress on the bones of the skull when biting massive prey.

Young Anteosaurus started their life with fairly narrow and lean skulls, and as it grew up bones of the skull became progressively thickened (process known as pachyostosis), creating the characteristic robust skull roof of Anteosaurus. The study of its inner ear revealed that Anteosaurus was a largely terrestrial, agile predator with highly advanced senses of vision, balance and coordination. It was also very fast and would have been able to outrun competitors and prey alike thanks to its advanced adaptations. Its body was well-suited to projecting itself forward, both in hunting and evidently in head-butting.

Anteosaurus and all other dinocephalians became extinct about 260 million years ago in a mass extinction at the end of the Capitanian in which the large Bradysaurian pareiasaurs also disappeared.^[1] The reasons of this extinction are obscure, although some research have shown a temporal association between the extinction of dinocephalian and an important volcanism event in China (known as the Emeishan Traps).^[2]^[1]

Etymology

Some confusion surrounds the etymology of the name Anteosaurus. It is often translated as meaning "before lizard", "previous lizard" or "primitive lizard", from the Latin prefix ante which means "before". The zoologist and paleontologist David Meredith Seares Watson gave no explanation when he named Anteosaurus in 1921. According to Ben Creisler,^[3] the prefix does not come from the Latin ante, but would refer to a Giant of the Greek mythology, Antaios, which once Latinized give Antaeus or more rarely Anteus. The type specimen of Anteosaurus is an incomplete skull that Watson had initially classified in the genus Titanosuchus, named after the Titans of the Greek mythology. Once this specimen recognized as belonging to a different genus, the name dedicated to Antaeus established another connection with a giant of Greek mythology.^[3]

Description

Size

Anteosaurus is one of the largest known carnivorous non-mammalian synapsid and anteosaurid, measuring around 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb).^[4]^[5] Juvenile specimen BP/1/7074 has an estimated body mass of about 71 kg (157 lb), showing extreme disparity in size with adult Anteosaurus.^[6]

Skull

The skull of Anteosaurus is large and massive, measuring between 80 and 90 centimetres (31 and 35 in) in the largest specimens (TM265 and SAM-PK-11293),^[7]^[8] ^[9] with an heavily pachyostosed skull roof showing a frontal boss more of less developed. The main features of the skull are the massively pachyostosed postfrontals that form strong horn-like bosses projected laterally. A boss, characteristically oval in shape, is also present on the angular bone of the lower jaw. The morphology of this angular boss is different between each anteosaurids species. In Anteosaurus the boss is oval in shape, roughly the same thickness throughout its length, with blunt anterior and posterior edges. Some individuals may have also a jugal boss more of less pronounced. Like other anteosaurids, the postorbital bar is strongly curved anteroventrally in such way that the temporal fenestra undercuts the orbit. An additional typical character of anteosaurs is the premaxilla oriented upwards at an angle of about 30 to 35° with respect to the ventral edge of the maxilla. However, unlike most anteosaurs in which the ventral margin of the premaxilla is directed upwards in a straight line, in Anteosaurus the anterior end of the premaxilla is curved ventrally, producing a concave alveolar border of the region preceding the canines. The skull shows also a concave dorsal snout profile. On the top of the skull, the pineal boss is exclusively formed by the parietals as is the case in other anteosaurines (and in more basal anteosaurs such as Archaeosyodon and Sinophoneus) while this boss is made up of both frontals and parietals in the other anteosaur subgroup, the syodontines.^[10] Contrary to what is observed in the latter, the frontals and the pineal boss of the anteosaurines do not participate in the attachment site of the mandibular adductor musculature.^[10] On the palate, the transverse processes of pterygoids are massively enlarged at their distal end, giving them a palmate shape in ventral view, as is the case in Titanophoneus and Sinophoneus. As in other anteosaurs, two prominent palatal bosses carried several small teeth.^[7]^[8]^[10] In Anteosaurus (and in other anteosaurines), these two palatal bosses are well separated from each other while in syodontines the two bosses are very close or interconnected.^[11]^[9]

Dentition

The dentition of Anteosaurus is composed of long to very long incisors, a large canine, and some small postcanines. In addition, some small teeth are present on both palatine bosses. There are five upper and four lower incisors, but even in the same skull the number in the two halves is mostly different. The incisors intermesh together. Like other anteosaurids, the first incisor of each premaxilla form together a pair that passes in between the lower pair formed by the first incisor of each dentary. The canines are well individualized. The upper canine is large and very massive, but is proportionally shorter than in some gorgonopsians of the Late Permian. The upper and lower canines did not intermesh. When the jaws were closed, the lower canines passed on the lingual side of the fifth upper incisor. Behind the canines, there are 4 to 8 small and relatively robust postcanines. Although smaller than the incisors and canines, these postcanines are proportionately more massive, with a thick base and a more conical general shape. Some postcanines of the upper jaws have a peculiar implantation. The most posterior are canted postero-laterally : the last three to four postcanine teeth are out-of-plane with the rest of the tooth row, being directed strongly backwards and somewhat outwards. Other smaller teeth were located on two prominences of the palate, the palatal bosses, which are semilunar or reniform in shape. These palatal teeth were recurved and most often implanted in a single curved row (a specimen however shows a double row). These teeth were used to hold meat during the swallowing process.^[8]^[12]^[13]^[10]

Postcranial skeleton

Reconstruction of Anteosaurus magnificus based on the skeletal proportions of the Russian anteosaur Titanophoneus potens.

Postcranial material of Anteosaurus is very rare and no complete skeletons are known. Only some associated or isolated bones (girdles and limbs bones, and some vertebrae), and more rarely some articulated remains have been found.^[14]^[15]^[16] An articulated left hand belonging to a juvenile individual shows that the manual phalangeal formula is 2-3-3-3-3 as in mammals.^[16] This hand (as well as an incomplete foot) was first considered by Lieuwe Dirk Boonstra as belonging to the right side of the animal. Boonstra himself corrected this mistake later by correctly identifying these remains as the left hand and foot. He also thought that digit III had four phalanges.^[14]^[15]^[17] Tim Rowe and J.A. van den Heever later showed that this was not the case, this digit having three phalanges.^[16] The manus have a digit I (the innermost) much smaller than the others. The digits III to V are the longest, the digit V (the outermost) being the most robust. The foot is only partially known, but also has a smaller digit I.^[17]^[16] Based on more complete skeletons of the Russian anteosaur Titanophoneus, the limbs would be rather long with a somewhat semi-erect posture. The tail is longer than in herbivorous Tapinocephalids dinocephalians.^[18]^[19]

Paleobiology

Skull variations and agonistic behaviour

The numerous skulls of Anteosaurus show a wide range of variation in cranial proportions and extent of pachyostosis. Most specifically the development of the postfrontal "horns" and the frontal boss is particularly variable between specimens. Some have both the "horns" and the boss massively pachyostosed, others have well-developed "horns" but a weak or nonexistent boss, and some others have a very weakly developed "horns" and boss. Even the heavily pachyostosed specimens show between them some variations. Some have "horns" relatively small compared to the boss, while others have postfrontal "horns" very massive. Some of these variations can be attributed to ontogenetic changes. In adults specimens the variations of the development of the frontal boss (to very weak to very strong) can be a sexually dimorphic feature, because in dinocephalians the frontal bosses have been implicated in head-butting and pushing behaviour.^[20]^[10]

Various authors have suggested the existence of agonistic behavior in Anteosaurus based on head-butting and/or demonstration involving canines. According to Herbert H. Barghusen, Anteosaurus does not use its teeth during intraspecific combat because both animals were able of doing severe damage to each other with their massive canines and incisors. The alternative head pushing strategy reduced the risk of fatal injuries in both combatants. The contact area of the skull roof during head combat included the most posterior part of the nasal bones, part of the prefrontal, and the entire frontal and postfrontal on either side. The thickened and laterally extended postfrontals horn-like bosses reduced the chance of the head of one opponent slipping past the head of the other.^[20]

More recently, Julien Benoit and colleagues have shown that the head of Anteosaurus had a natural posture that was less tilted downwards than that of the tapinocephalids and that, unlike the latter, it does not line up ideally with the vertebral column to optimize a head-to-head combat. This peculiarity associated with the presence of a pachyostosis less developed than that of the tapinocephalids and the retention of a large canine led these authors to suggest an agonistic behavior in which Anteosaurus more likely used its large canines for displays and/or during confrontation involving bites.^[6]

According to Christian Kammerer, the pachyostosis of Anteosaurus would have mainly allowed the skull to resist the cranial stress generated by the powerful external adductor muscles during the bite on a large prey, as has been suggested in other macropredators with a thickened supraorbital region such as rubidgeine gorgonopsians, mosasaurs, some thalattosuchians, sebecosuchians, rauisuchians and various large carnivorous dinosaurs.^[10]^[9]

All these authors, however, do not exclude a multiple use of this pachyostosis and the existence in Anteosaurus of a head-butting behaviour requiring however less energy than that of the Tapinocephalidae.^[10]^[6]

Ontogeny

Ashley Kruger and team in 2016 described a juvenile specimen of Anteosaurus (BP/1/7074), providing details into the ontogeny of this anteosaurid. Analyzed allometry between this specimen and others suggests that the cranial ontogeny of Anteosaurus was characterized by a rapid growth in the temporal region, a significant difference in the development of the postorbital bar and suborbital bar between juveniles and adults, as well as a notorious pachyostosis (bone thickening) during development, which ultimately modified the skull roof of adults. Consequently, pachyostosis was responsible for thickening important skull bones such as the frontal and postfrontal which were of great importance in the overall paleobiology and behavior of Anteosaurus. Kruger and team noted that these differences, when compared, are extreme between juvenile and mature Anteosaurus individuals.^[21]

In 2021 Mohd Shafi Bhat histologically studied several skeletal remains of specimens referred to Anteosaurus, finding three growth stages. The first growth stage is characterized by the predominance of highly vascularized, uninterrupted fibrolamellar bone tissue in the inner bone cortex, which suggests rapid formation of new bone during early ontogeny. A second stage of growth in Anteosaurus is represented by periodic/seasonal interruptions in the bone formation, indicated by the deposition of lines of arrested growth. Third and last reported growth stage by the team features the development of lamellar bone tissue with rest lines in the peripheral part of the bone cortex, which indicates that Anteosaurus slowed down growth at advanced age.^[22]

Habitat preference and diet

Boonstra in 1954 indicated that the overall dentition of Anteosaurus—characterized by prominent canines, elongated incisors, and relatively weak postcanines—reflects a specialized carnivore, and that this anteosaurid did not rely on chewing and shearing when feeding, but rather it was well-adapted for tearing flesh chunks from prey. In addition, Boonstra noted that some of the flesh material was likely held and/or torn by the recurved palatal dentition.^[8] Later in 1955, Boonstra indicated that anteosaurids had a crawling locomotion similar to crocodiles, based mostly on their hip joint and femur morphology, useful in a semiaquatic setting.^[23]

In 2008 Mivah F. Ivakhnenko analyzed a vast majority of Permian therapsid skulls, and suggested that anteosaurs, such as Anteosaurus, were strict semiaquatic piscivorous (fish-eater) synapsids, mostly similar to modern-day otters.^[24] Christian F. Kammerer in 2011 questioned this proposal, given that numerous anatomical traits of anteosaurs make this life-style unlikely. The typical dentition of piscivore animals include elongate, numerous, strongly recurved, and very sharp teeth in order to hold and kill fast-moving fish prey. In addition, the jaws of piscivores are commonly elongated and narrow for quick snatchs and minimal water resistance when shaking prey. Unlike these traits, the skull morphology of most anteosaurs—specifically anteosaurids—is extremely robust with deep jaws, and the teeth are bulbous and blunt, with only the canine being the recurved-most tooth. Kammerer instead indicated that anteosaurids like Anteosaurus likely preyed on large terrestrial dinocephalians, such as the gigantic titanosuchids and tapinocephalids. He also noted that anteosaurid teeth are mostly similar to that of large tyrannosaurids (postcanines robust bases, faceted surfaces, and obliquely angled serrations), whose dentition is interpreted as bone-crunching. Accordingly, bone-crunching may also have been employed by anteosaurids and an important component in their diet.^[10]

In 2020 Kévin Rey with colleagues analyzed stable oxygen isotope compositions of phosphate from teeth and bones from pareiasaurs and Anteosaurus, in order to estimate their affinity for water dependence. Obtained results showed similar δ18Op values between pareiasaurs, Anteosaurus, and therocephalians, with a wide range of extant terrestrial species, which indicated a terrestrial preference for these synapsids. However, it was noted that the δ18Op values were slightly lower in Anteosaurus, casting doubt for this interpretation. Nevertheless, Rey with colleagues concluded that a larger sample size may result in a more robust conclusion for Anteosaurus.^[25]

Bhat and team in 2021 noted that most skeletal elements of Anteoaurus are characterized by relatively thickened bone walls, extensive secondary bone reconstruction and the complete infilling of the medullary cavity. Combined, these traits indicate that Anteosaurus was mostly adapted for a terrestrial life-style. However a radius and femur have open medullary cavities with struts of bony trabeculae. The team suggested that it is conceable that Anteosaurus may have also occasionally inhabited shallow and short-lived pools, in a similar manner to modern-day hippopotamuses.^[22]

An in-depth study of the brain of juvenile Anteosaurus specimen BP/1/7074 published in 2021 disproves the idea that this dinocephalian was a sluggish, crocodilian-like predator. Studies by Benoit et al. using x-ray imaging and 3-D reconstructions showcase that Anteosaurus was a fast, agile animal in spite of its great size. Its inner ears were larger than those of its closest relatives and competitors, showcasing that it was well-suited to the role of an apex predator that could outrun both its rivals and prey alike. It was also determined that the area of the brain of Anteosaurus that was responsible for coordinating the movements of the eyes with the head was exceptionally large; an important feature in ensuring it could track its prey accurately. As a result, Anteosaurus was well-adapted to swift hunting and fast attacking strikes on land.^[6]

Geographic and stratigraphic range

South Africa

The fossils of Anteosaurus magnificus come mainly from the Abrahamskraal Formation as well as from the basal part of the Teekloof Formation of the Beaufort Group in the Karoo Basin, South Africa. The species appears in the middle part of the Abrahamskraal Formation (Kornplaats member) and continues in the rest of the formation (Swaerskraal, Moordenaars, and Kareskraal members). Its last representatives come from the base of the Teekloof formation (in the lower strata of the Poortjie member).^[26]^[1] More than 30 localities are known, most of them being localized in the Western Cape province (Beaufort West, Prince Albert and Laingsburg). Some localities are also known near the towns of Sutherland and Fraserburg in the southern end of the Northern Cape province (Karoo Hoogland).^[8]^[27]^[10] and at least one specimen (BP/1/7061) was found near Grahamstown in the Eastern Cape Province (Makana).^[28]^{[nb 1]} A skull discovered in the same province in 2001 was also tentatively ascribed to a juvenile specimen of Anteosaurus.^[29]^[30] However, the complete preparation of this skull, made later, revealed that it belonged to a tapinocephalid dinocephalian.^[31]^{[nb 2]}

The Middle Permian Abrahamskraal Formation is biostratigraphically subdivided in two faunal zone : the Eodicynodon Assemblage Zone which is the oldest one with an essentially Wordian age, and the Tapinocephalus Assemblage Zone, which is mainly Capitanian in age. Anteosaurus belongs to the Tapinocephalus Assemblage Zone which is characterized by the abundance and the diversification of the dinocephalians therapsids. Since 2020, this zone is divided into two subzones : a lower Eosimops - Glanosuchus subzone and an upper Diictodon - Styracocephalus subzone, both of which contain Anteosaurus fossils.^[26] Like all other South African dinocephalians, Anteosaurus was presumed extinct at the top of the Abrahamskraal Formation. However, remains of Anteosaurus and two other dinocephalian genera (Titanosuchus and Criocephalosaurus) have been found in the basal portion of the Poortjie Member of the overlying Teekloof Formation. These discoveries greatly expanded both the stratigraphic range of these three dinocephalian genera and the upper limit of the Tapinocephalus Assemblage Zone that reaches the base of the Teekloof Formation.^[32]^[1] In the latter, the remains of these three dinocephalians were found in an interval of 30 metres (98 ft) above a level dated to 260.259 ± 0.081 million years ago, representing the Upper Capitanian.^[1] Other radiometric dating have constrained the base of the Tapinocephalus Assemblage Zone (Leeuvlei Member in the middle part of the Abrahamskraal Formation) to be older than 264.382 ± 0.073 Ma and placed the boundary between the two subzones at 262.03 ± 0.15 Ma.^[33] The upper part of the Abrahamskraal Formation (top of the Karelskraal Member) gave an age of 260.226 ± 0.069 Ma which is consistent with the age of 260.259 ± 0.081 of the base of the Teekloof Formation.^[33] These datings show that the age of the Tapinocephalus Assemblage Zone extends from Late Wordian to Late Capitanian (based on Guadalupian radiometric ages obtained in 2020 from the type locality of the Guadalupe Mountains in west Texas).^[33]^[34]

Russia?

The genus Anteosaurus is possibly present in Russia based on a fragmentary cranial remain found in the 19th century in the Republic of Tatarstan (Alexeyevsky District). This specimen, firstly interpreted as a snout boss of a dicynodont (named Oudenodon rugosus), was later correctly identified by Ivan Efremov as an angular boss of an anteosaurid. The shape of this boss clearly differs from those of others Russian anteosaurids, so this specimen was attributed to a new species of the genus Titanophoneus (and named Titanophoneus rugosus). More recently, Christian Kammerer showed that the shape of this boss differs markedly from the lenticular bosses of the Russian anteosaurs T. potens and T. adamanteus. In contrast the angular boss of T. rugosus is very similar to the Anteosaurus morphotype, so this specimen can be the first representative of the genus Anteosaurus in Russia. The dermal sculpturing of the boss, with prominent furrows, is different from that observed in few well preserved A. magnificus specimens. According to Kammerer, as the range of variation in dermal sculpturing between Anteosaurus individuals is no well known, it is more reasonable to consider provisionally Titanophoneus rugosus as a nomen dubium (maybe an Anteosaurus sp.). Only the discovery of more complete Russians specimens with the rugosus morphotype will clarify the relationship of this taxon with Anteosaurus.^[10]

Paleoenvironment

Paleogeography and paleoclimate

At the time of Anteosaurus, most of the landmasses were united in one supercontinent, Pangaea. It 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 Sea.^[35] 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.^[36] The territory that would become the South African Karoo was located much further south than today, at the level of the 60th parallel south.^[37] Although located close to the Antarctic Circle, the climate prevailing at this latitude during most of the Permian was temperate with distinct seasons.^[38]^[39]^[40] There are uncertainties about the temperatures that prevailed in South Africa during the Middle Permian. Previously, this region of the world had undergone significant glaciation during the Upper Carboniferous.^[41] Subsequently, the Lower Permian had first seen the retreat of glaciers and the emergence of subpolar tundra and taiga-like vegetation (dominated by Botrychiopsis and Gangamopteris),^[42] then the introduction of warmer and wetter climatic conditions that allowed the development of the Mesosaurus fauna and the Glossopteris flora.^[41] The scientists who studied the climate of that time found very different results on the thermal ranges that existed in the ancient Karoo. At the end of the 1950s, Edna Plumstead compared the Karoo to today's Siberia or Canada, with a highly seasonal climate including very cold winters and temperate summers supporting the Glossopteris flora, which would have been restricted to sheltered basins.^[43] Later, other studies, mainly based on climate models, also suggested a cold temperate climate with high thermal amplitude between summer (+15 to +20 °C) and winter (-20 to -25 °C).^[44]^[45] More recent studies also indicate a temperate climate, but with much less severe winters than those previously suggested. Keddy Yemane thus suggested that the vast river system and the many giant lakes present at the time throughout southern Africa must have significantly moderated the continentality of the Karoo climate during most of the Permian.^[46] Paleobotanical studies focusing on the characteristic morphology of plant leaves and the growth rings of fossil woods also indicate a seasonal climate^[39]^[37] with summer temperatures of up to 30 °C and free-frost winters.^[39] According to Richard Rayner, the high southern latitudes experienced very hot and humid summers, with an average of 18 hours of light per day for more than four months during which precipitation was comparable to the annual amount falling in the present-day tropics. These conditions were extremely conducive to rapid growth in plants such as Glossopteris.^[39] The habit in Glossopteris of losing its leaves at the beginning of the bad season would be linked to a shorter duration of daylight rather than the existence of very cold winter temperatures.^[39] From the geochemical study of sediments from several Karoo sites, Kay Scheffler also obtains a temperate climate (with mean annual temperatures of about 15 to 20 °C), with free-frost winter, but with an increase in aridity during the Middle Permian.^[41]

Paleoecology

The sediments of the Abrahamskraal Formation consists of a succession of sandstones, and versicolor siltstones and mudstones, deposited by large rivers that flowed from south to north from the Gondwanide mountain range. These large rivers of variable sinuosity drained a vast alluvial plain that sloped gently down to the northeast toward the Ecca sea (a former landlocked sea), while in receding phase.^[47]^[48]^[49]^[50] The landscape was composed of marshy land, interrupted by rivers, lakes, woods and forests. Many fossil traces (footprints, ripple marks, mudcracks) indicate that swampy areas, which were the most extensive habitat, were frequently exposed to the open air and should not often be deeply flooded.^[38] The vegetation was dominated by the deciduous pteridosperm Glossopteris, which formed woodlands and large forests concentrated along the streams and on the uplands.^[38]^[39] Large horsetails (2–3 m (6.6–9.8 ft) high), such as Schizoneura and Paraschizoneura, formed bamboo-like stands that grew in and around swamps. Herbaceous horsetails (Phyllotheca) and ferns carpeted the undergrowth and small lycopods occupied the wetter areas.^[38]^[39]

Aquatic fauna included the lamellibranch Palaeomutela, the palaeonisciformes fishes Atherstonia, Bethesdaichthys, Blourugia, Namaichthys and Westlepis, and large freshwater predators, the temnospondyl amphibians Rhinesuchoides and Rhinesuchus.^[51]^[52] The terrestrial fauna was particularly diverse and dominated by the therapsids.^{[nb 3]} Anteosaurus occupied the top of the food chain there. It shared its environment with many other carnivorous tetrapods. Other large predatory animals included the lion-sized Lycosuchid therocephalians Lycosuchus and Simorhinella,^[53] and the Scylacosaurid therocephalian Glanosuchus.^[54] Medium-sized carnivorous were represented by the basal biarmosuchian Hipposaurus,^[55] the more derived biarmosuchian Bullacephalus,^[56] the scylacosaurids Ictidosaurus,^[54] Scylacosaurus,^[57] and Pristerognathus,^[26] and the small and basal gorgonopsian Eriphostoma.^[58]^[59] The small predator guild (mainly insectivorous forms) included the therocephalians Alopecodon,^[26] and Pardosuchus,^[26] the small monitor-like varanopids Elliotsmithia,^[60]^[57] Heleosaurus,^[61]^[62]^[63] and Microvaranops,^[64] the millerettid Broomia,^[26] the procolophonomorph Australothyris,^[65] and the lizard-like Eunotosaurus^[66] of uncertain affinities (variously considered as a parareptile,^[67] a pantestudine^[68]^[69] or a caseid synapsid^[70]).

Herbivorous were also numerous and diversified. Large-sized vegetarians were mainly represented by numerous dinocephalians including the tapinocephalids Agnosaurus,^[26]^[71] Criocephalosaurus,^[72]^[73]^[32]^[71] Mormosaurus,^[26]^[71] Moschognathus,^[26]^[71] Moschops,^[74]^[75]^[76]^[71] Riebeeckosaurus,^[75]^[77] Struthiocephalus^[78]^[79]^[76]^[80] Struthionops,^[26] and Tapinocephalus,^[72]^[81]^[76]^[71] the Styracocephalid Styracocephalus,^[82]^[83]^[84] and the huge titanosuchids Jonkeria and Titanosuchus.^[85]^[7]^[57]^[26] Other large herbivores that were not synapsids included the large bradysaurian pareiasaurs represented by Bradysaurus, Embrithosaurus and Nochelesaurus, whose dentition very different from that of herbivorous dinocephalians indicates that the two groups occupied clearly distinct ecological niches.^[86]^[87]^[88]^[89]^[90] The small to medium-sized forms included basal anomodonts (the non-dicynodonts Anomocephalus,^[91]^[92] Galechirus, Galeops and Galepus^[93]^[57]^[26]) and numerous dicynodonts (Brachyprosopus,^[94] Colobodectes,^[95]^[96] Pristerodon,^[57] and the Pylaecephalids Diictodon,^[97]^[57] Eosimops,^[98] Prosictodon,^[99] and Robertia,^[57]).

Classification and phylogeny

Named by Watson in 1921, Anteosaurus was longtime classified as a 'Titanosuchian Deinocephalian', and it is only in 1954 that Boonstra separated the Titanosuchians in two families: Jonkeridae (a junior synonym of Titanosuchidae) and Anteosauridae.^[100]^[8] At about the same time, Efremov erected the family Brithopodidae in which he includes the fragmentary Brithopus and the better known forms Syodon and Titanophoneus.^[101] Much later, Hopson and Barghusen argued that Brithopodidae should be discontinued and that the Russian taxa Syodon, Titanophoneus and Doliosauriscus should be placed with Anteosaurus in Anteosauridae. These authors placed also Anteosauridae in the new group Anteosauria for distinguished them of the other major dinocephalian group the Tapinocephalia in which they included the titanosuchids and the tapinocephalids. They also created the taxa Anteosaurinae, containing Anteosaurus and the Russian forms Titanophoneus and Doliosauriscus, and the Anteosaurini containing only the giant forms Anteosaurus and Doliosauriscus.^[102] Gilian King retained the incorrectly spelled 'Brithopidae' (including the subfamilies 'Brithopinae' and Anteosaurinae) and placed both Brithopidae and Titanosuchidae (including Titanosuchinae and Tapinocephalinae) in the superfamily Anteosauroidea.^[103] Later Ivakhnenko considered Brithopodidae as invalid and united Anteosauridae and Deuterosauridae (only known by the Russian Deuterosaurus) in the superfamily Deuterosauroidea.^[104]^[105] More recently Kammerer in its systematic revision of the anteosaurs (in which Doliosauriscus become a junior synonym of Titanophoneus) demonstrated that the wastebasket genus Brithopus is a nomen dubium composed both of remains of indeterminate estemmenosuchid-like tapinocephalian and indeterminate anteosaurian, so invalidating the Brithopodidae. He proposed also the first phylogenetic analysis including all anteosaurid taxa. This and other modern phylogenetic analysis of anteosaurs recovers a monophyletic Anteosauridae containing two major clades, Syodontinae and Anteosaurinae. In the Kammerer analysis, the Chinese Sinophoneus is the most basal anteosaurine and the sister-group of an unresolved trichotomy including Titanophoneus potens, T. adamanteus and Anteosaurus.^[10]

Below the cladogramm of Kammerer published in 2011 :

	Anteosaurus magnificus

	Titanophoneus adamanteus

	Titanophoneus potens

Sinophoneus yumenensis

Syodontinae

	Syodon biarmicum

	Australosyodon nyaphuli

Notosyodon gusevi

Archaeosyodon praeventor

Microsyodon orlovi

Tapinocephalia

	Tapinocaninus pamelae

	Estemmenosuchus mirabilis

Biarmosuchus tener

In describing the new Brazilian anteosaur Pampaphoneus, Cisneros et al. presented another cladogram confirming the recognition of the clades Anteosaurinae and Syodontinae. In the cladogram of the Fig. 2. of the main paper, which does not include the genus Microsyodon, Titanophoneus adamanteus is recovered as the sister taxon of a clade composed of Titanophoneus potens and Anteosaurus. However, in the four cladograms of the Fig. S1, presented in the Supporting Information of the same article, and including Microsyodon, Anteosaurus is recovered as the sister taxon of both species of Titanophoneus. These four cladograms differ only by the position of Microsyodon.^[11]

The cladogram of Cisneros et al. published in the main paper and excluding the genus Microsyodon. T. adamanteus is here the sister taxon of a clade composed of T. potens and Anteosaurus :^[11]

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

One of the four cladograms of Cisneros et al. published in the Supporting Information of the same article, and including Microsyodon. In all these cladogram, Anteosaurus is recovered as the sister taxon of both species of Titanophoneus :^[11]

	Ulemosaurus

	Tapinocaninus

Anteosaurus

	T. potens

	T. adamanteus

Syodontinae

Pampaphoneus

Notosyodon

Syodon

	Microsyodon

	Australosyodon

In resdescribing the Chinese anteosaur Sinophoneus, Jun Lui presented a new cladogram in which Sinophoneus is recovered as the most basal Anteosauridae and so excluded of the Anteosaurinae. Anteosaurus being also positioned as the sister-taxon of Titanophoneus potens and T. adamanteus.

The cladogramm of Jun Liu in 2013:^[106]

Therapsida

Biarmosuchus tener

Dinocephalia

Estemmenosuchus uralensis

	Ulemosaurus svijagensis

	Tapinocaninus pamelae

Anteosauridae

Sinophoneus yumenensis

Archaeosyodon praeventor

Anteosaurinae

Anteosaurus magnificus

	Titanophoneus adamanteus

	Titanophoneus potens

Syodontinae

Pampaphoneus biccai

Notosyodon gusevi

Syodon biarmicum

	Australosyodon nyaphuli

	Microsyodon orlovi

Genus synonymy

As defined by Lieuwe Dirk Boonstra, Anteosaurus is "a genus of anteosaurids in which the postfrontal forms a boss of variable size overhanging the dorso-posterior border of the orbit." On this basis he synonymised six of the seven genera named from the Tapinocephalus zone: Eccasaurus, Anteosaurus, Titanognathus, Dinosuchus, Micranteosaurus, and Pseudanteosaurus. Of these, he says, Dinosuchus and Titanognathus can safely be considered synonyms of Anteosaurus. Eccasaurus, with a holotype of which the cranial material consists of only few typical anteosaurid incisors, appears to be only determinable as to family. The skull fragment forming the holotype of Pseudanteosaurus can best be considered as an immature specimen of Anteosaurus. Micranteosaurus, the holotype of which contains a small snout, was previously considered a new genus on account of its small size but is better be interpreted as a young specimen of Anteosaurus. And likewise, the large number of species attributed to the genus Anteosaurus can also be considered synonyms. Boonstra still considers as valid the genus Paranteosaurus, which is defined as a genus of anteosaurids in which the postfrontal is not developed to form a boss. This is probably an example of individual variation and hence another synonym of Anteosaurus.^[55]

Species synonymy

Anteosaurus was once known by a large number of species, but the current thinking on this is that they merely represent different growth stages of the same type species, A. magnificus.^[55]^[10]

We have 32 skulls of Anteosaurus, of which 16 are reasonably well preserved and on them ten species have been named. To differentiate between the species the following main characters have. been used: the number, size and shape of the teeth, skull size, shape and the nature of the pachyostosis. On re-examination it has become clear that the crowns of the teeth are seldom well preserved; basing the count for the dental formula on the preserved roots is unreliable. as this is affected by age and tooth generation; size of skull is a function of age and also possibly sex; skull shape is greatly affected by post-mortem deformation, and the variability in the pachyostosis, which may be specific in some respects, can just as well be the result of...physiological processes. Specific diagnosis consisting of the enumeration of differences of degree in features such as the above can hardly be considered as sufficient indication of the existence of discrete species....A. magnificus thus has the following synonyms: abeli, acutirostrus, crassifrons, cruentus, laticeps, levops, lotzi, major, minor, minusculus, parvus, priscus and vorsteri.
— Boonstra.^[55]

Possible synonyms

Archaeosuchus

Archaeosuchus cairncrossi is a dubious species of anteosaur from the Tapinocephalus Assemblage Zone. It was named by Broom in 1905 on the basis of a partial maxilla. It was interpreted as a titanosuchid by Boonstra, but Kammerer determined it was an anteosaur indistinguishable from Anteosaurus and Titanophoneus. As Anteosaurus magnificus appears to be the only valid large anteosaur in the Tapinocephalus Assemblage Zone, Archaeosaurus cairncrossi is very likely to be based on a specimen of it, but due to poor preservation, the specimen lacks any features that would allow the synonymy to be proven.^[10]

Eccasaurus

Eccasaurus priscus is a dubious species of anteosaur from the Tapinocephalus Assemblage Zone. It was named by Robert Broom in 1909 on the basis of a fragmentary skeleton, of which Broom only described the humerus. As with Archaeosuchus cairncrossi, Eccasaurus priscus is very likely to be synonymous with Anteosaurus magnificus. As Eccasaurus was named before Anteosaurus, a petition to the ICZN would be needed to preserve the name Anteosaurus magnificus if the synonymy were to be proven.^[10]

Notes

^ This specimen was originally discovered in the Koonap Formation which was considered a lateral equivalent of the Abrahamskraal Formation east of the 24th meridian east. The two formations were amalgamated in 2016 because lithologically indistinguishable (cf. references Cole & al. 2016).
^ This skull was first attributed to Moschops capensis before being reassigned to Moschognathus whaitsi in 2020 by Saniye Neumann in a thesis work (cf. references).
^ Several taxa from the lower part of the Tapinocephalus zone, such as the biarmosuchian Pachydectes, the dicynodont Lanthanostegus, and the gorgonopsian Phorcys, have been found in strata located several hundred meters below the oldest known specimens of Anteosaurus. This particular fauna could represent a new zone or sub-zone not yet recognized as such and located between the Eodicynodon and Tapinocephalus Assemblage Zones (Day & Rubidge 2020, Rubidge & al. 2021).

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External links

Media related to Anteosaurus at Wikimedia Commons
Data related to Anteosaurus at Wikispecies
"Therapsida: Anteosauria". Palaeos. Archived from the original on November 24, 2010.
"Synapsida". Paleofile. Archived from the original on January 8, 2023.

[29] This specimen was originally discovered in the Koonap Formation which was considered a lateral equivalent of the Abrahamskraal Formation east of the 24th meridian east. The two formations were amalgamated in 2016 because lithologically indistinguishable (cf. references Cole & al. 2016).

[33] This skull was first attributed to Moschops capensis before being reassigned to Moschognathus whaitsi in 2020 by Saniye Neumann in a thesis work (cf. references).

[55] Several taxa from the lower part of the Tapinocephalus zone, such as the biarmosuchian Pachydectes, the dicynodont Lanthanostegus, and the gorgonopsian Phorcys, have been found in strata located several hundred meters below the oldest known specimens of Anteosaurus. This particular fauna could represent a new zone or sub-zone not yet recognized as such and located between the Eodicynodon and Tapinocephalus Assemblage Zones (Day & Rubidge 2020, Rubidge & al. 2021).

[Day&Rubidge2021-1] Day, M.O.; Rubidge, B.S. (2021). "The Late Capitanian Mass Extinction of Terrestrial Vertebrates in the Karoo Basin of South Africa". Frontiers in Earth Science. 9: 15. Bibcode:2021FrEaS...9...15D. doi:10.3389/feart.2021.631198.

[day2015a-2] Day, M.O.; Ramezani, J.; Bowring, S.A.; Sadler, P.M.; Erwin, D.H.; Abdala, F.; Rubidge, B.S. (2015). "When and how did the terrestrial mid-Permian mass extinction occur? Evidence from the tetrapod record of the Karoo Basin, South Africa". Proceedings of the Royal Society B. 282 (1811): 20150834. doi:10.1098/rspb.2015.0834. PMC 4528552. PMID 26156768.

[dml-3] "Misunderstood Giants: Geosaurus, Anteosaurus, Otozoum". Archives of the Dinosaur Mailing List. Ben Kreisler. Retrieved 30 October 2014.

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[5] Prothero, Donald R. (18 April 2022). "20. Synapsids: The Origin of Mammals". Vertebrate Evolution: From Origins to Dinosaurs and Beyond. Boca Raton: CRC Press. doi:10.1201/9781003128205-4. ISBN 978-0-36-747316-7. S2CID 246318785.

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@@ Line 13: / Line 13: @@
 {{collapsible list|bullets = true
  |title=<small>Genus synonymy</small>
-|''[[Eccasaurus]]'' (?)<br><small>Broom, 1909</small>
+|''Eccasaurus'' (?)<br><small>Broom, 1909</small>
 |''Titanognathus'' <br><small>Broili & Schröder, 1935</small>
 |''Dinosuchus'' <br><small>Broom, 1936</small>
@@ Line 23: / Line 23: @@
 {{collapsible list|bullets = true
  |title=<small>Species synonymy</small>
-|''[[Eccasaurus|Eccasaurus priscus]]'' (?)<br><small>Broom, 1909</small>
+|''Eccasaurus priscus'' (?)<br><small>Broom, 1909</small>
 |''Anteosaurus minor'' <br><small>Broom, 1929</small>
 |''Titanognathus lotzi'' <br><small>Broili & Schröder, 1935</small>
@@ Line 45: / Line 45: @@
 }}<!--|access-date=24 October 2015-->
-'''''Anteosaurus''''' (meaning "[[Antaeus]] reptile") is an [[extinct]] [[genus]] of large carnivorous [[dinocephalia]]n [[synapsid]]. It lived at the end of the [[Guadalupian]] (= Middle Permian) 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 {{convert|5|-|6|m|ft|abbr=on}} long and weighing about {{convert|600|kg|lb|abbr=on}}, ''Anteosaurus'' was the largest known [[carnivorous]] [[synapsid|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 [[Titanosuchidae|titanosuchids]] and [[Tapinocephalidae|tapinocephalids]] dinocephalians and large [[Pareiasauromorpha|pareiasaurs]].
+'''''Anteosaurus''''' (meaning "[[Antaeus]] lizard") is an [[extinct]] [[genus]] of large carnivorous [[dinocephalia]]n [[synapsid]]. It lived at the end of the [[Guadalupian]] (= Middle Permian) during the [[Capitanian]] age, 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 {{convert|5|-|6|m|ft|abbr=on}} long and weighing about {{convert|600|kg|lb|abbr=on}}, ''Anteosaurus'' was the largest known [[carnivorous]] [[synapsid|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 [[Titanosuchidae|titanosuchids]] and [[Tapinocephalidae|tapinocephalids]] dinocephalians and large [[Pareiasauromorpha|pareiasaurs]].
 As in many other dinocephalians the cranial bones of ''Anteosaurus'' are [[pachyostosis|pachyostosed]], but to a lesser extent than in tapinocephalid dinocephalians. In ''Anteosaurus'', pachyostosis mainly occurs in the form of horn-shaped supraorbital protuberances. According to some paleontologists this structure would be implicated in intraspecific [[agonistic behaviour]], including head-pushing probably during the mating season. On the contrary, other scientists believe that this pachyostosis served to reduce cranial stress on the bones of the skull when biting massive prey.
@@ Line 62: / Line 62: @@
 === Skull ===
+The [[skull]] of ''Anteosaurus'' is large and massive, measuring between {{convert|80|and|90|cm|0}} in the largest specimens (TM265 and SAM-PK-11293),<ref name="boonstra1953a"/><ref name="boonstra1954a">{{cite journal|last=Boonstra|first=L.D.|title=The cranial structure of the titanosuchian: Anteosaurus|journal=Annals of the South African Museum|year=1954|volume=42|issue=|pages=108–148|url=http://www.biodiversitylibrary.org/page/40889579#page/152/mode/1up}}</ref> <ref name="Angielczyk&Kammerer2018">{{cite book|last=Angielczyk |first=K.D. |author2=Kammerer, C.F. |year=2018 |chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree |pages=178 |title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics |editor1-last=Zachos |editor1-first=F.E. |editor2=Asher, R.J. |publisher=de Gruyter |location=Berlin |isbn=978-3-11-027590-2 }}</ref> with an heavily [[Pachyostosis|pachyostosed]] skull roof showing a frontal boss more of less developed. The main features of the skull are the massively pachyostosed postfrontals that form strong horn-like bosses projected laterally. A boss, characteristically oval in shape, is also present on the [[angular bone|angular]] bone of the lower jaw. The morphology of this angular boss is different between each anteosaurids species. In ''Anteosaurus'' the boss is oval in shape, roughly the same thickness throughout its length, with blunt anterior and posterior edges. Some individuals may have also a [[jugal bone|jugal]] boss more of less pronounced. Like other anteosaurids, the [[postorbital bone|postorbital]] bar is strongly curved anteroventrally in such way that the [[Skull#Temporal fenestrae|temporal fenestra]] undercuts the orbit. An additional typical character of anteosaurs is the premaxilla oriented upwards at an angle of about 30 to 35° with respect to the ventral edge of the maxilla. However, unlike most anteosaurs in which the ventral margin of the premaxilla is directed upwards in a straight line, in ''Anteosaurus'' the anterior end of the premaxilla is curved ventrally, producing a concave alveolar border of the region preceding the canines. The skull shows also a concave dorsal snout profile. On the top of the skull, the [[Parietal eye|pineal]] boss is exclusively formed by the [[Parietal bone|parietals]] as is the case in other anteosaurines (and in more [[Basal (phylogenetics)|basal]] anteosaurs such as ''[[Archaeosyodon]]'' and ''[[Sinophoneus]]'') while this boss is made up of both frontals and parietals in the other anteosaur subgroup, the syodontines.<ref name="kammerer2011">{{cite journal|last1=Kammerer |first1=C. F. |title=Systematics of the Anteosauria (Therapsida: Dinocephalia) |journal=Journal of Systematic Palaeontology |year=2011 |volume=9 |issue=2 |pages=261–304 |doi=10.1080/14772019.2010.492645 |bibcode=2011JSPal...9..261K |s2cid=84799772 }}</ref> Contrary to what is observed in the latter, the [[Frontal bone|frontals]] and the pineal boss of the anteosaurines do not participate in the attachment site of the mandibular [[Anatomical terms of motion#Abduction and adduction|adductor]] musculature.<ref name="kammerer2011"/> On the palate, the transverse processes of pterygoids are massively enlarged at their distal end, giving them a palmate shape in ventral view, as is the case in ''[[Titanophoneus]]'' and ''[[Sinophoneus]]''. As in other anteosaurs, two prominent palatal bosses carried several small teeth.<ref name="boonstra1953a">{{cite journal|last=Boonstra|first=L.D.|title=A suggested clarification of the taxonomic status of the South African titanosuchians|journal=Annals of the South African Museum|year=1953|volume=42|issue=|pages=19–28|url=http://www.biodiversitylibrary.org/page/40889428#page/49/mode/1up}}</ref><ref name="boonstra1954a"/><ref name="kammerer2011"/> In ''Anteosaurus'' (and in other anteosaurines), these two palatal bosses are well separated from each other while in syodontines the two bosses are very close or interconnected.<ref name="Cetal12">{{cite journal |last=Cisneros |first=J.C. |author2=Abdala, F. |author3= Atayman-Güven, S. |author4= Rubidge, B.S. |author5= Şengör, A.M.C. |author6= Schultz, C.L. |year=2012 |title=Carnivorous dinocephalian from the Middle Permian of Brazil and tetrapod dispersal in Pangaea |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=109 |issue=5 |pages=1584–1588 |doi=10.1073/pnas.1115975109 |pmid=22307615 |pmc=3277192 |bibcode=2012PNAS..109.1584C |url=http://www.pnas.org/content/109/5/1584.full.pdf?with-ds=yes |doi-access=free }}</ref><ref name="Angielczyk&Kammerer2018"/>
-[[File:Annals of the South African Museum = Annale van die Suid-Afrikaanse Museum (1953) (17798589344).jpg|thumb|left|This skull of  ''A. magnificus'' (SAM-PK-11296), measuring 70 cm in length, was formerly the [[holotype]] of ''A. abeli''.<ref name="boonstra1953a"/><ref name="boonstra1954a">{{cite journal|last=Boonstra|first=L.D.|title=The cranial structure of the titanosuchian: Anteosaurus|journal=Annals of the South African Museum|year=1954|volume=42|issue=|pages=108–148|url=http://www.biodiversitylibrary.org/page/40889579#page/152/mode/1up}}</ref> It presents a very pronounced pachyostosis. The frontal boss is particularly massive, so much so that postfrontals boss, although well developed, seem very small in comparison. The angular boss, oriented obliquely, is clearly visible at the back of the lower jaw. This specimen was discovered near [[Sutherland, Northern Cape|Sutherland]] in the [[Northern Cape]] province.<ref name="kammerer2011"/>]]
-The [[skull]] of ''Anteosaurus'' is large and massive, measuring between {{convert|80|and|90|cm|0}} in the largest specimens (TM265 and SAM-PK-11293),<ref name="boonstra1953a"/><ref name="boonstra1954a"/><ref name="Angielczyk&Kammerer2018">{{cite book|last=Angielczyk |first=K.D. |author2=Kammerer, C.F. |year=2018 |chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree |pages=178 |title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics |editor1-last=Zachos |editor1-first=F.E. |editor2=Asher, R.J. |publisher=de Gruyter |location=Berlin |isbn=978-3-11-027590-2 }}</ref> with an heavily [[Pachyostosis|pachyostosed]] skull roof showing a frontal boss more of less developed. The main features of the skull are the massively pachyostosed postfrontals that form strong horn-like bosses projected laterally. A boss, characteristically oval in shape, is also present on the [[angular bone|angular]] bone of the lower jaw. The morphology of this angular boss is different between each anteosaurids species. In ''Anteosaurus'' the boss is oval in shape, roughly the same thickness throughout its length, with blunt anterior and posterior edges. Some individuals may have also a [[jugal bone|jugal]] boss more of less pronounced. Like other anteosaurids, the [[postorbital bone|postorbital]] bar is strongly curved anteroventrally in such way that the [[Skull#Temporal fenestrae|temporal fenestra]] undercuts the orbit. An additional typical character of anteosaurs is the premaxilla oriented upwards at an angle of about 30 to 35° with respect to the ventral edge of the maxilla. However, unlike most anteosaurs in which the ventral margin of the premaxilla is directed upwards in a straight line, in ''Anteosaurus'' the anterior end of the premaxilla is curved ventrally, producing a concave alveolar border of the region preceding the canines. The skull shows also a concave dorsal snout profile. On the top of the skull, the [[Parietal eye|pineal]] boss is exclusively formed by the [[Parietal bone|parietals]] as is the case in other anteosaurines (and in more [[Basal (phylogenetics)|basal]] anteosaurs such as ''[[Archaeosyodon]]'' and ''[[Sinophoneus]]'') while this boss is made up of both frontals and parietals in the other anteosaur subgroup, the syodontines.<ref name="kammerer2011">{{cite journal|last1=Kammerer |first1=C. F. |title=Systematics of the Anteosauria (Therapsida: Dinocephalia) |journal=Journal of Systematic Palaeontology |year=2011 |volume=9 |issue=2 |pages=261–304 |doi=10.1080/14772019.2010.492645 |bibcode=2011JSPal...9..261K |s2cid=84799772 }}</ref> Contrary to what is observed in the latter, the [[Frontal bone|frontals]] and the pineal boss of the anteosaurines do not participate in the attachment site of the mandibular [[Anatomical terms of motion#Abduction and adduction|adductor]] musculature.<ref name="kammerer2011"/> On the palate, the transverse processes of pterygoids are massively enlarged at their distal end, giving them a palmate shape in ventral view, as is the case in ''[[Titanophoneus]]'' and ''[[Sinophoneus]]''. As in other anteosaurs, two prominent palatal bosses carried several small teeth.<ref name="boonstra1953a">{{cite journal|last=Boonstra|first=L.D.|title=A suggested clarification of the taxonomic status of the South African titanosuchians|journal=Annals of the South African Museum|year=1953|volume=42|issue=|pages=19–28|url=http://www.biodiversitylibrary.org/page/40889428#page/49/mode/1up}}</ref><ref name="boonstra1954a"/><ref name="kammerer2011"/> In ''Anteosaurus'' (and in other anteosaurines), these two palatal bosses are well separated from each other while in syodontines the two bosses are very close or interconnected.<ref name="Cetal12">{{cite journal |last=Cisneros |first=J.C. |author2=Abdala, F. |author3= Atayman-Güven, S. |author4= Rubidge, B.S. |author5= Şengör, A.M.C. |author6= Schultz, C.L. |year=2012 |title=Carnivorous dinocephalian from the Middle Permian of Brazil and tetrapod dispersal in Pangaea |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=109 |issue=5 |pages=1584–1588 |doi=10.1073/pnas.1115975109 |pmid=22307615 |pmc=3277192 |bibcode=2012PNAS..109.1584C |url=http://www.pnas.org/content/109/5/1584.full.pdf?with-ds=yes |doi-access=free }}</ref><ref name="Angielczyk&Kammerer2018"/>
 === Dentition ===
-The dentition of ''Anteosaurus'' is composed of long to very long [[incisor]]s, a large [[canine tooth|canine]], and some small postcanines. In addition, some small teeth are present on both [[palatine bone|palatine]] bosses. There are five upper and four lower incisors, but even in the same skull the number in the two halves is mostly different. The incisors intermesh together. Like other anteosaurids, the first incisor of each premaxilla form together a pair that passes in between the lower pair formed by the first incisor of each dentary. The canines are well individualized. The upper canine is large and very massive, but is proportionally shorter than in some [[gorgonopsia]]ns of the Late Permian. The upper and lower canines did not intermesh. When the jaws were closed, the lower canines passed on the lingual side of the fifth upper incisor. Behind the canines, there are 4 to 8 small and relatively robust postcanines. Although smaller than the incisors and canines, these postcanines are proportionately more massive, with a thick base and a more conical general shape. Some postcanines of the upper jaws have a peculiar implantation. The most posterior are canted postero-laterally : the last three to four postcanine teeth are out-of-plane with the rest of the tooth row, being directed strongly backwards and somewhat outwards. Other smaller teeth were located on two prominences of the palate, the palatal bosses, which are semilunar or reniform in shape. These palatal teeth were recurved and most often implanted in a single curved row (a specimen however shows a double row). These teeth were used to hold meat during the [[swallowing|swallowing process]].<ref name="boonstra1954a"/><ref name="Rubidge1991">{{cite journal|last=Rubidge|first=B.S.|title=A new primitive dinocephalian mammal-like reptile from the Permian of Southern Africa|year=1991|volume=34|issue=3|pages=547–559|url=https://www.palass.org/publications/palaeontology-journal/archive/34/3/article_pp547-559}}</ref><ref name="vanValkenburgh2002">{{cite journal|last1=van Valkenburgh |first1=B. |author2=Jenkins, I. |title=Evolutionary Patterns in the History of Permo-Triassic and Cenozoic Synapsid Predators |journal=Paleontological Society Papers |date=2002 |volume=8 |issue= |pages=267–288 |doi=10.1017/S1089332600001121 |citeseerx=10.1.1.729.1135 }}</ref><ref name="kammerer2011"/>
+The dentition of ''Anteosaurus'' is composed of long to very long [[incisor]]s, a large [[canine tooth|canine]], and some small postcanines. In addition, some small teeth are present on both [[palatine bone|palatine]] bosses. There are five upper and four lower incisors, but even in the same skull the number in the two halves is mostly different. The incisors intermesh together. Like other anteosaurids, the first incisor of each premaxilla form together a pair that passes in between the lower pair formed by the first incisor of each dentary. The canines are well individualized. The upper canine is large and very massive, but is proportionally shorter than in some [[gorgonopsia]]ns of the Late Permian. The upper and lower canines did not intermesh. When the jaws were closed, the lower canines passed on the lingual side of the fifth upper incisor. Behind the canines, there are 4 to 8 small and relatively robust postcanines. Although smaller than the incisors and canines, these postcanines are proportionately more massive, with a thick base and a more conical general shape. Some postcanines of the upper jaws have a peculiar implantation. The most posterior are canted postero-laterally : the last three to four postcanine teeth are out-of-plane with the rest of the tooth row, being directed strongly backwards and somewhat outwards. Other smaller teeth were located on two prominences of the palate, the palatal bosses, which are semilunar or reniform in shape. These palatal teeth were recurved and most often implanted in a single curved row (a specimen however shows a double row). These teeth were used to hold meat during the [[swallowing|swallowing process]].<ref name="boonstra1954a"/><ref name="Rubidge1991">{{cite journal |last=Rubidge |first=B.S. |year=1991 |title=A new primitive dinocephalian mammal-like reptile from the Permian of Southern Africa |url=https://www.palass.org/publications/palaeontology-journal/archive/34/3/article_pp547-559 |journal=Palaeontology |volume=34 |issue=3 |pages=547–559}}</ref><ref name="vanValkenburgh2002">{{cite journal|last1=van Valkenburgh |first1=B. |author2=Jenkins, I. |title=Evolutionary Patterns in the History of Permo-Triassic and Cenozoic Synapsid Predators |journal=Paleontological Society Papers |date=2002 |volume=8 |issue= |pages=267–288 |doi=10.1017/S1089332600001121 |citeseerx=10.1.1.729.1135 }}</ref><ref name="kammerer2011"/>
 === Postcranial skeleton ===
@@ Line 74: / Line 73: @@
 ==Paleobiology==
 === Skull variations and agonistic behaviour ===
-[[File:Anteosaurus magnificus holotype.jpg|thumb|left|Holotype (BMNH R3595) skull roof of ''A. magnificus''. In this specimen the postfrontal bosses are particularly massive. This skull was discovered near [[Beaufort West]] in the [[Western Cape]] Province.<ref name="kammerer2011"/>]][[File:Annals of the South African Museum - Annale van die Suid-Afrikaanse Museum (1953) (18233550770).jpg|thumb|right|Skull of ''A. magnificus'' (SAM-PK-11694). This weakly pachyostosed specimen is 56.5 cm long. Formerly the holotype of ''A. cruentus''. It was discovered near [[Laingsburg Local Municipality|Laingsburg]] in the [[Western Cape]] province.]]
+[[File:Anteosaurus magnificus holotype.jpg|thumb|left|Holotype (BMNH R3595) skull roof of ''A. magnificus''. In this specimen the postfrontal bosses are particularly massive. This skull was discovered near [[Beaufort West]] in the [[Western Cape]] Province.<ref name="kammerer2011"/>]]
 The numerous skulls of ''Anteosaurus'' show a wide range of variation in cranial proportions and extent of pachyostosis. Most specifically the development of the postfrontal "horns" and the [[frontal bone|frontal]] boss is particularly variable between specimens. Some have both the "horns" and the boss massively pachyostosed, others have well-developed "horns" but a weak or nonexistent boss, and some others have a very weakly developed "horns" and boss. Even the heavily pachyostosed specimens show between them some variations. Some have "horns" relatively small compared to the boss, while others have postfrontal "horns" very massive. Some of these variations can be attributed to [[Ontogeny|ontogenetic]] changes. In adults specimens the variations of the development of the frontal boss (to very weak to very strong) can be a [[Sexual dimorphism|sexually dimorphic]] feature, because in dinocephalians the frontal bosses have been implicated in head-butting and pushing behaviour.<ref name="barghusen1975"/><ref name="kammerer2011"/>
 Various authors have suggested the existence of agonistic behavior in ''Anteosaurus'' based on head-butting and/or demonstration involving canines. According to Herbert H. Barghusen, ''Anteosaurus'' does not use its teeth during intraspecific combat because both animals were able of doing severe damage to each other with their massive canines and incisors. The alternative head pushing strategy reduced the risk of fatal injuries in both combatants. The contact area of the skull roof during head combat included the most posterior part of the [[nasal bone]]s, part of the [[prefrontal bone|prefrontal]], and the entire frontal and postfrontal on either side. The thickened and laterally extended postfrontals horn-like bosses reduced the chance of the head of one opponent slipping past the head of the other.<ref name="barghusen1975">{{cite journal|last=Barghusen |first=H.R. |title=A review of fighting adaptations in dinocephalians (Reptilia, Therapsida) |journal=Paleobiology |year=1975 |volume=1 |issue= 3|pages=295–311 |doi=10.1017/S0094837300002542 |jstor=2400370 |bibcode=1975Pbio....1..295B |s2cid=87163815 |url=http://www.jstor.org/stable/2400370 }}</ref>
+[[File:Anteosaurus SAM-PK-K360 skull.jpg|thumb|right|Skull of  ''A. magnificus'' (SAM-PK-K360) with strong postfrontal bosses but no frontal boss. This specimen, exhibited in the [[Iziko South African Museum]] of [[Cape Town]], comes from Nuwelande, near [[Fraserburg]] in [[Northern Cape]] province.<ref name="kammerer2011"/>]]
 More recently, Julien Benoit and colleagues have shown that the head of ''Anteosaurus'' had a natural posture that was less tilted downwards than that of the tapinocephalids and that, unlike the latter, it does not line up ideally with the vertebral column to optimize a head-to-head combat. This peculiarity associated with the presence of a pachyostosis less developed than that of the tapinocephalids and the retention of a large canine led these authors to suggest an agonistic behavior in which ''Anteosaurus'' more likely used its large canines for displays and/or during confrontation involving bites.<ref name=Benoit2021>{{cite journal|last1=Benoit |first1=J.|last2=Kruger |first2=A.|last3=Jirah |first3=S.|last4=Fernandez |first4=V. |last5=Rubidge |first5=B. S.|date=2021|title=Palaeoneurology and palaeobiology of the dinocephalian therapsid Anteosaurus magnificus|journal=Acta Palaeontologica Polonica |volume=66 |doi=10.4202/app.00800.2020|doi-access=free|url=https://www.app.pan.pl/archive/published/app66/app008002020.pdf}}</ref>
-[[File:Annals of the South African Museum - Annale van die Suid-Afrikaanse Museum (1953) (18394843986).jpg|thumb|left|Skull of  ''A. magnificus'' (SAM-PK-4340), formerly classified in the species ''A. abeli''. In this individual, (skull length = 68 cm) the frontal boss and the posfrontal bosses are highly developed.<ref name="boonstra1953a"/><ref name="boonstra1954a"/> This specimen comes from Leeurivier near [[Beaufort West]] in the [[Western Cape]] province.<ref name="kammerer2011"/>]][[File:Anteosaurus SAM-PK-K360 skull.jpg|thumb|right|Skull of  ''A. magnificus'' (SAM-PK-K360) with strong postfrontal bosses but no frontal boss. This specimen, exhibited in the [[Iziko South African Museum]] of [[Cape Town]], comes from Nuwelande, near [[Fraserburg]] in [[Northern Cape]] province.<ref name="kammerer2011"/>]]
 According to Christian Kammerer, the pachyostosis of ''Anteosaurus'' would have mainly allowed the skull to resist the cranial stress generated by the powerful external adductor muscles during the bite on a large prey, as has been suggested in other macropredators with a thickened supraorbital region such as [[Rubidgeinae|rubidgeine]] [[gorgonopsia]]ns, [[mosasaur]]s, some [[thalattosuchia]]ns, [[sebecosuchia]]ns, [[rauisuchia]]ns and various large carnivorous [[dinosaur]]s.<ref name="kammerer2011"/><ref name="Angielczyk&Kammerer2018"/>
@@ Line 88: / Line 88: @@
 ===Ontogeny===
-Ashley Kruger and team in 2016 described a juvenile specimen of ''Anteosaurus'' (BP/1/7074), providing details into the ontogeny of this anteosaurid. Analyzed allometry between this specimen and others suggests that the cranial ontogeny of ''Anteosaurus'' was characterized by a rapid growth in the temporal region, a significant difference in the development of the postorbital bar and suborbital bar between juveniles and adults, as well as a notorious [[pachyostosis]] (bone thickening) during development, which ultimately modified the [[skull roof]] of adults. Consequently, pachyostosis was responsible for thickening important skull bones such as the frontal and postfrontal which were of great importance in the overall paleobiology and behavior of ''Anteosaurus''. Kruger and team noted that these differences, when compared, are extreme between juvenile and mature ''Anteosaurus'' individuals.<ref>{{cite journal|last1=Kruger|first1=A.|last2=Rubidge|first2=B. S.|last3=Abdala|first3=F.|date=2016|title=A juvenile specimen of Anteosaurus magnificus Watson, 1921 (Therapsida: Dinocephalia) from the South African Karoo, and its implications for understanding dinocephalian ontogeny|journal=Journal of Systematic Palaeontology|volume=16|issue=2|pages=139–158 |doi=10.1080/14772019.2016.1276106|s2cid=90346300 |url=https://fernando.losabdala.com/a-juvenile-specimen-of-anteosaurus-magnificus-watson-1921-therapsida-dinocephalia-from-the-south-african-karoo-and-its-implications-for-understanding-dinocephalian-ontogeny/}}</ref>
+Ashley Kruger and team in 2016 described a juvenile specimen of ''Anteosaurus'' (BP/1/7074), providing details into the ontogeny of this anteosaurid. Analyzed allometry between this specimen and others suggests that the cranial ontogeny of ''Anteosaurus'' was characterized by a rapid growth in the temporal region, a significant difference in the development of the postorbital bar and suborbital bar between juveniles and adults, as well as a notorious [[pachyostosis]] (bone thickening) during development, which ultimately modified the [[skull roof]] of adults. Consequently, pachyostosis was responsible for thickening important skull bones such as the frontal and postfrontal which were of great importance in the overall paleobiology and behavior of ''Anteosaurus''. Kruger and team noted that these differences, when compared, are extreme between juvenile and mature ''Anteosaurus'' individuals.<ref>{{cite journal|last1=Kruger|first1=A.|last2=Rubidge|first2=B. S.|last3=Abdala|first3=F.|date=2016|title=A juvenile specimen of Anteosaurus magnificus Watson, 1921 (Therapsida: Dinocephalia) from the South African Karoo, and its implications for understanding dinocephalian ontogeny|journal=Journal of Systematic Palaeontology|volume=16|issue=2|pages=139–158 |doi=10.1080/14772019.2016.1276106|s2cid=90346300 |url=https://fernando.losabdala.com/a-juvenile-specimen-of-anteosaurus-magnificus-watson-1921-therapsida-dinocephalia-from-the-south-african-karoo-and-its-implications-for-understanding-dinocephalian-ontogeny/|hdl=11336/67088|hdl-access=free}}</ref>
 In 2021 Mohd Shafi Bhat [[histologically]] studied several skeletal remains of specimens referred to ''Anteosaurus'', finding three growth stages. The first growth stage is characterized by the predominance of highly vascularized, uninterrupted fibrolamellar bone tissue in the inner bone cortex, which suggests rapid formation of new bone during early ontogeny. A second stage of growth in ''Anteosaurus'' is represented by periodic/seasonal interruptions in the bone formation, indicated by the deposition of [[lines of arrested growth]]. Third and last reported growth stage by the team features the development of lamellar bone tissue with rest lines in the peripheral part of the bone cortex, which indicates that ''Anteosaurus'' slowed down growth at advanced age.<ref name=Bhat2021>{{cite journal|last1=Bhat|first1=M. S.|last2=Shelton|first2=C. D.|last3=Chinsamy|first3=A.|date=2021|title=Inter-element variation in the bone histology of Anteosaurus (Dinocephalia, Anteosauridae) from the Tapinocephalus Assemblage Zone of the Karoo Basin of South Africa|journal=PeerJ|volume=9|pages=e12082|doi=10.7717/peerj.12082|doi-access=free|pmc=8434808|pmid=34589298}}</ref>
@@ Line 95: / Line 95: @@
 [[File:Anteosaurus & Moschognathus.png|thumb|left|Semi-transparent 3D reconstructions (based on X-ray micro-computed tomography) of the skulls of dinocephalians ''Anteosaurus magnificus'' (A, BP/1/7074) and ''Moschognathus whaitsi'' (B, AM4950) from the middle Permian of South Africa, aligned on the plane of their lateral semicircular canal. The black arrow indicates the tilting of the long axis of the skull compared to the plane of the lateral semicircular canal.]]
-Boonstra in 1954 indicated that the overall dentition of ''Anteosaurus''—characterized by prominent canines, elongated incisors, and relatively weak postcanines—reflects an specialized [[carnivore]], and that this anteosaurid did not rely on [[chewing]] and shearing when feeding, but rather it was well-adapted for tearing flesh chunks from [[prey]]. In addition, Boonstra noted that some of the flesh material was likely held and/or teared by the recurved palatal dentition.<ref name="boonstra1954a"/> Later in 1955, Boonstra indicated that anteosaurids had a crawling locomotion similar to [[crocodile]]s, based mostly on their hip joint and femur morphology, useful in a [[semiaquatic]] setting.<ref>{{cite journal |last1=Boonstra|first1=L. D.|date=1955|title=The girdles and limbs of South African dinocephalians|journal=Annals of the South African Museum|volume=42|pages=185−327}}</ref>
+Boonstra in 1954 indicated that the overall dentition of ''Anteosaurus''—characterized by prominent canines, elongated incisors, and relatively weak postcanines—reflects a specialized [[carnivore]], and that this anteosaurid did not rely on [[chewing]] and shearing when feeding, but rather it was well-adapted for tearing flesh chunks from [[prey]]. In addition, Boonstra noted that some of the flesh material was likely held and/or torn by the recurved palatal dentition.<ref name="boonstra1954a"/> Later in 1955, Boonstra indicated that anteosaurids had a crawling locomotion similar to [[crocodile]]s, based mostly on their hip joint and femur morphology, useful in a [[semiaquatic]] setting.<ref>{{cite journal |last1=Boonstra|first1=L. D.|date=1955|title=The girdles and limbs of South African dinocephalians|journal=Annals of the South African Museum|volume=42|pages=185−327}}</ref>
-In 2008 Mivah F. Ivakhnenko analyzed a vast majority of [[Permian]] therapsid skulls, and suggested that anteosaurs, such as ''Anteosaurus'', were strict semiaquatic [[piscivorous]] ([[fish]]-eater) synapsids, mostly similar to modern-day [[otter]]s.<ref>{{cite journal |last1=Ivakhnenko|first1=M. F.|date=2008|title=Cranial morphology and evolution of Permian Dinomorpha (Eotherapsida) of eastern Europe|journal=Paleontological Journal|volume=42|number=9|pages=859−995|doi=10.1134/S0031030108090013|bibcode=2008PalJ...42..859I |s2cid=85114195 }}</ref> Christian F. Kammerer in 2011 questioned this proposal, given that numerous anatomical traits of anteosaurs make this life-style unlikely. The typical dentition of piscivore animals include elongate, numerous, strongly recurved, and very sharp teeth in order to hold and kill fast-moving fish prey. In addition, the jaws of piscivores are commonly elongated and narrow for quick snatchs and minimize water resistance when shaking prey. Unlike these traits, the skull morphology of most anteosaurs—specifically anteosaurids—is extremely robust with deep jaws, and the teeth are bulbous and blunt, with only the canine being the recurved-most tooth. Kammerer instead indicated that anteosaurids like ''Anteosaurus'' likely preyed on large terrestrial dinocephalians, such as the gigantic [[titanosuchid]]s and [[tapinocephalid]]s. He also noted that anteosaurid teeth are mostly similar to that of large [[tyrannosaurids]] (postcanines robust bases, faceted surfaces, and obliquely angled serrations), whose dentition is interpreted as bone-crunching. Accordingly, bone-crunching may also have been employed by anteosaurids and an important component in their diet.<ref name="kammerer2011"/>
+In 2008 Mivah F. Ivakhnenko analyzed a vast majority of [[Permian]] therapsid skulls, and suggested that anteosaurs, such as ''Anteosaurus'', were strict semiaquatic [[piscivorous]] ([[fish]]-eater) synapsids, mostly similar to modern-day [[otter]]s.<ref>{{cite journal |last1=Ivakhnenko|first1=M. F.|date=2008|title=Cranial morphology and evolution of Permian Dinomorpha (Eotherapsida) of eastern Europe|journal=Paleontological Journal|volume=42|number=9|pages=859−995|doi=10.1134/S0031030108090013|bibcode=2008PalJ...42..859I |s2cid=85114195 }}</ref> Christian F. Kammerer in 2011 questioned this proposal, given that numerous anatomical traits of anteosaurs make this life-style unlikely. The typical dentition of piscivore animals include elongate, numerous, strongly recurved, and very sharp teeth in order to hold and kill fast-moving fish prey. In addition, the jaws of piscivores are commonly elongated and narrow for quick snatchs and minimal water resistance when shaking prey. Unlike these traits, the skull morphology of most anteosaurs—specifically anteosaurids—is extremely robust with deep jaws, and the teeth are bulbous and blunt, with only the canine being the recurved-most tooth. Kammerer instead indicated that anteosaurids like ''Anteosaurus'' likely preyed on large terrestrial dinocephalians, such as the gigantic [[titanosuchid]]s and [[tapinocephalid]]s. He also noted that anteosaurid teeth are mostly similar to that of large [[tyrannosaurids]] (postcanines robust bases, faceted surfaces, and obliquely angled serrations), whose dentition is interpreted as bone-crunching. Accordingly, bone-crunching may also have been employed by anteosaurids and an important component in their diet.<ref name="kammerer2011"/>
 [[File:Anteosaurus in landscape.jpg|thumb|right|Reconstruction of ''Anteosaurus'' in terrestrial paleoenvironment]]
-In 2020 Kévin Rey with colleagues analyzed stable oxygen isotope compositions of phosphate from teeth and bones from [[pareiasaurs]] and ''Anteosaurus'', in order to estimate their affinity for water dependence. Obtained results showed similar δ18Op values between pareiasaurs, ''Anteosaurus'', and therocephalians, with a wide range of extant terrestrial species, which indicated a terrestrial preference for these synapsids. However, it was noted that the δ18Op values were slightly lower in ''Anteosaurus'', casting doubt for this interpretation. Nevertheless, Rey with colleagues concluded that a larger sample size may result in a more robust conclusion for ''Anteosaurus''.<ref>{{cite journal|last1=Rey|first1=K.|last2=Day|first2=M. O.|last3=Amiot|first3=R.|last4=Fourel|first4=F.|last5=Luyt|first5=J.|last6=Van den Brandt|first6=M. J.|last7=Lécuyer|first7=C.|last8=Rubidge|first8=B. S.|date=2020|title=Oxygen isotopes and ecological inferences of Permian (Guadalupian) tetrapods from the main Karoo Basin of South Africa|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|volume=538|page=109485 |doi=10.1016/j.palaeo.2019.109485|bibcode=2020PPP...538j9485R |s2cid=214085715 |url=https://hal.archives-ouvertes.fr/hal-02991787/file/Rey%20et%20al.%20%282020a%29%20pre-proof.pdf }}</ref>
+In 2020 Kévin Rey with colleagues analyzed stable oxygen isotope compositions of phosphate from teeth and bones from [[pareiasaurs]] and ''Anteosaurus'', in order to estimate their affinity for water dependence. Obtained results showed similar δ18Op values between pareiasaurs, ''Anteosaurus'', and therocephalians, with a wide range of extant terrestrial species, which indicated a terrestrial preference for these synapsids. However, it was noted that the δ18Op values were slightly lower in ''Anteosaurus'', casting doubt for this interpretation. Nevertheless, Rey with colleagues concluded that a larger sample size may result in a more robust conclusion for ''Anteosaurus''.<ref>{{cite journal|last1=Rey|first1=K.|last2=Day|first2=M. O.|last3=Amiot|first3=R.|last4=Fourel|first4=F.|last5=Luyt|first5=J.|last6=Van den Brandt|first6=M. J.|last7=Lécuyer|first7=C.|last8=Rubidge|first8=B. S.|date=2020|title=Oxygen isotopes and ecological inferences of Permian (Guadalupian) tetrapods from the main Karoo Basin of South Africa|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|volume=538|page=109485 |doi=10.1016/j.palaeo.2019.109485|bibcode=2020PPP...53809485R |s2cid=214085715 |url=https://hal.archives-ouvertes.fr/hal-02991787/file/Rey%20et%20al.%20%282020a%29%20pre-proof.pdf }}</ref>
 Bhat and team in 2021 noted that most skeletal elements of ''Anteoaurus'' are characterized by relatively thickened bone walls, extensive secondary bone reconstruction and the complete infilling of the medullary cavity. Combined, these traits indicate that ''Anteosaurus'' was mostly adapted for a terrestrial life-style. However a radius and femur have open medullary cavities with struts of bony trabeculae. The team suggested that it is conceable that ''Anteosaurus'' may have also occasionally inhabited shallow and short-lived pools, in a similar manner to modern-day [[hippopotamuses]].<ref name=Bhat2021/>
@@ Line 114: / Line 114: @@
 The Middle Permian Abrahamskraal Formation is [[Biostratigraphy|biostratigraphically]] subdivided in two faunal zone : the [[Eodicynodon Assemblage Zone|''Eodicynodon'' Assemblage Zone]] which is the oldest one with an essentially [[Wordian]] age, and the [[Tapinocephalus Assemblage Zone]], which is mainly [[Capitanian]] in age. ''Anteosaurus'' belongs to the [[Tapinocephalus Assemblage Zone|''Tapinocephalus'' Assemblage Zone]] which is characterized by the abundance and the diversification of the [[dinocephalia]]ns [[Therapsida|therapsids]]. Since 2020, this zone is divided into two subzones : a lower ''[[Eosimops]]'' - ''[[Glanosuchus]]'' subzone and an upper ''[[Diictodon]]'' - ''[[Styracocephalus]]'' subzone, both of which contain ''Anteosaurus'' fossils.<ref name="Day&Rubidge2020"/> Like all other South African dinocephalians, ''Anteosaurus'' was presumed extinct at the top of the Abrahamskraal Formation. However, remains of ''Anteosaurus'' and two other dinocephalian genera (''[[Titanosuchus]]'' and ''[[Criocephalosaurus]]'') have been found in the basal portion of the Poortjie Member of the overlying Teekloof Formation. These discoveries greatly expanded both the stratigraphic range of these three dinocephalian genera and the upper limit of the ''Tapinocephalus'' Assemblage Zone that reaches the base of the Teekloof Formation.<ref name="day2015b">{{cite journal|last=Day|first=M.O.|author2=Güven, S.|author3=Abdala, F.|author4=Jirah, S.|author5=Rubidge, B.S.|author6=Almond, J.|title=Youngest dinocephalian fossils extend the Tapinocephalus Zone, Karoo Basin, South Africa.|journal=South African Journal of Science|year=2015|volume=111|issue=3–4|pages=1–5|doi=10.17159/sajs.2015/20140309|url=http://www.sajs.co.za/sites/default/files/publications/pdf/Day_Research%20Letter_0.pdf|access-date=2022-10-31|archive-date=2017-12-02|archive-url=https://web.archive.org/web/20171202210544/http://www.sajs.co.za/sites/default/files/publications/pdf/Day_Research%20Letter_0.pdf|url-status=dead}}</ref><ref name="Day&Rubidge2021"/> In the latter, the remains of these three dinocephalians were found in an interval of {{convert|30|m|0}} above a level dated to 260.259 ± 0.081 million years ago, representing the Upper Capitanian.<ref name="Day&Rubidge2021"/> Other [[radiometric dating]] have constrained the base of the ''Tapinocephalus'' Assemblage Zone (Leeuvlei Member in the middle part of the Abrahamskraal Formation) to be older than 264.382 ± 0.073 Ma and placed the boundary between the two subzones at 262.03 ± 0.15 Ma.<ref name="Day&al2022">{{cite journal |last1=Day|first1=M.O.|author2=Ramezani, J.|author3=Frazer, R.E.|author4=Rubidge, B.S.|year=2022|title=U-Pb zircon age constraints on the vertebrate assemblages and palaeomagnetic record of the Guadalupian Abrahamskraal Formation, Karoo Basin, South Africa|journal=Journal of African Earth Sciences|volume=186|issue=|page=104435|doi=10.1016/j.jafrearsci.2021.104435|bibcode=2022JAfES.18604435D |s2cid=245086992 }}</ref> The upper part of the Abrahamskraal Formation (top of the Karelskraal Member) gave an age of 260.226 ± 0.069 Ma which is consistent with the age of 260.259 ± 0.081 of the base of the Teekloof Formation.<ref name="Day&al2022"/> These datings show that the age of the ''Tapinocephalus'' Assemblage Zone extends from Late [[Wordian]] to Late [[Capitanian]] (based on Guadalupian radiometric ages obtained in 2020 from the type locality of the [[Guadalupe Mountains]] in west [[Texas]]).<ref name="Day&al2022"/><ref name="Wu&al2020">{{cite journal |last1=Wu|first1=Q.|author2=Ramezani, J.|author3=Zhang, H.|author4=Yuan, D-x|author5=Erwin, D.H.|author6=Henderson, C.M.|author7=Lambert, L.L.|author8=Zhang, Y-c|author9=Shen, S-z|year=2020|title=High-precision U-Pb age constraints on the Guadalupian in west Texas, USA|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|volume=548|issue=|page=109668|doi=10.1016/j.palaeo.2020.109668|s2cid=214342275 }}</ref>
-===Russia ?===
+===Russia?===
-The genus ''Anteosaurus'' is possibly present in [[Russia]] based on a fragmentary cranial remain found in the 19th century in the [[Tatarstan|Republic of Tatarstan]] ([[Alexeyevsky District, Republic of Tatarstan|Alexeyevsky District]]). This specimen, firstly interpreted as a snout boss of a [[dicynodontia|dicynodont]] (named ''Oudenodon rugosus''), was later correctly identified by [[Ivan Yefremov|Ivan Efremov]] as an angular boss of an anteosaurid. The shape of this boss clearly differs from those of others Russian anteosaurids, so this specimen was attributed to a new species of the genus ''Titanophoneus'' (and named ''Titanophoneus rugosus''). More recently, Christian Kammerer showed that the shape of this boss differs markedly from the lenticular bosses of the Russian anteosaurs ''T. potens'' and ''T. adamanteus''. In contrast the angular boss of ''T. rugosus'' is very similar to the ''Anteosaurus'' morphotype, so this specimen can be the first representative of the genus ''Anteosaurus'' in Russia. The dermal sculturing of the boss, with prominent furrows, is different from that observed in few well preserved ''A. magnificus'' specimens. According to Kammerer, as the range of variation in dermal sculturing between ''Anteosaurus'' individuals is no well known, it is more reasonable to consider provisionally ''Titanophoneus rugosus'' as a [[nomen dubium]] (maybe an ''Anteosaurus'' sp.). Only the discovery of more complete Russians specimens with the ''rugosus'' morphotype will clarify the relationship of this taxon with ''Anteosaurus''.<ref name="kammerer2011"/>
+The genus ''Anteosaurus'' is possibly present in [[Russia]] based on a fragmentary cranial remain found in the 19th century in the [[Tatarstan|Republic of Tatarstan]] ([[Alexeyevsky District, Republic of Tatarstan|Alexeyevsky District]]). This specimen, firstly interpreted as a snout boss of a [[dicynodontia|dicynodont]] (named ''Oudenodon rugosus''), was later correctly identified by [[Ivan Yefremov|Ivan Efremov]] as an angular boss of an anteosaurid. The shape of this boss clearly differs from those of others Russian anteosaurids, so this specimen was attributed to a new species of the genus ''Titanophoneus'' (and named ''Titanophoneus rugosus''). More recently, Christian Kammerer showed that the shape of this boss differs markedly from the lenticular bosses of the Russian anteosaurs ''T. potens'' and ''T. adamanteus''. In contrast the angular boss of ''T. rugosus'' is very similar to the ''Anteosaurus'' morphotype, so this specimen can be the first representative of the genus ''Anteosaurus'' in Russia. The dermal sculpturing of the boss, with prominent furrows, is different from that observed in few well preserved ''A. magnificus'' specimens. According to Kammerer, as the range of variation in dermal sculpturing between ''Anteosaurus'' individuals is no well known, it is more reasonable to consider provisionally ''Titanophoneus rugosus'' as a [[nomen dubium]] (maybe an ''Anteosaurus'' sp.). Only the discovery of more complete Russians specimens with the ''rugosus'' morphotype will clarify the relationship of this taxon with ''Anteosaurus''.<ref name="kammerer2011"/>
 ==Paleoenvironment==
@@ Line 121: / Line 121: @@
 [[File:Anteosaurus distribution.jpg|thumb|left|Map of Earth during Guadalupian time showing the paleogeographic distribution of the genus ''Anteosaurus''.]]
-At the time of ''Anteosaurus'', most of the landmasses were united in one supercontinent, [[Pangaea]]. It 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 Ocean|Tethys Sea]].<ref name="McLoughlin2001">{{cite journal|last1=McLoughlin|first1=S.|title=The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism|journal=Australian Journal of Botany|date=2001|volume=49|issue=3|pages=271–300|doi=10.1071/BT00023}}</ref> A long string of microcontinents, grouped under the name of [[Cimmeria (continent)|Cimmeria]], divided the Tethys in two : the Paleo-Tethys in the north, and the Neo-Tethys in the south.<ref name="Şengör1987">{{cite journal|last1=Şengör|first1=A.M.C.|title=Tectonics of the Tethysides: orogenic collage development in a collisional setting|journal=Annual Review of Earth and Planetary Sciences|date=1987|volume=15|issue=|pages=214–244|doi=10.1146/annurev.ea.15.050187.001241|bibcode=1987AREPS..15..213C }}</ref> The territory that would become the South African Karoo was located much further south than today, at the level of the [[60th parallel south]].<ref name="Bamford2016">{{cite book|last=Bamford|first=M.K.|year=2016|chapter=Fossil woods from the Upper Carboniferous to Lower Jurassic Karoo Basin and their environmental interpretation|pages=159–167|title=Origin and evolution of the Cape Mountains and Karoo Basin|editor1-last=Linol|editor1-first=B.|editor2=de Wit, M.J.|publisher=Regional Geology Reviews, Springer|location=Cham|isbn=978-3-319-40858-3}}</ref> Although located close to the [[Antarctic Circle]], the climate prevailing at this [[latitude]] during most of the Permian was [[Temperate climate|temperate]] with distinct seasons.<ref name="king1990">{{cite book|last=King|first=G.|year=1990|chapter=The environment of the ''Tapinocephalus'' zone|pages=107–111|title=The dicynodonts. A study in palaeobiology|editor1-last=King|editor1-first=G.|publisher=Chapman and Hall|location=London and New York|isbn=0-412-33080-6}}</ref><ref name="Rayner1996">{{cite journal|last1=Rayner|first1=R.J.|title=The palaeoclimate of the Karoo: evidence from plant fossils|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|date=1996|volume=119|issue=3–4|pages=385–394|doi=10.1016/0031-0182(95)00021-6|bibcode=1996PPP...119..385R }}</ref><ref name="Chumakov&Zharkov2003">{{cite journal|last1=Chumakov|first1=N.M.|author2=Zharkov, M.A.|title=Climate during the Permian-Triassic Biosphere Reorganizations. Article 2. Climate of the Late Permian and Early Triassic : General Inferences|journal=Stratigraphy and Geological Correlation|date=2003|volume=11|issue=4|pages=361–376}}</ref>  There are uncertainties about the temperatures that prevailed in South Africa during the Middle Permian. Previously, this region of the world had undergone significant [[Karoo Ice Age|glaciation]] during the Upper [[Carboniferous]].<ref name="Scheffler & al.2006">{{cite journal|last1=Scheffler|first1=K.|author2=Buehmann, D.|author3=Schwark, L.|title=Analysis of Late Palaeozoic glacial to potglacial sedimentary successions in South Africa by geochemical proxies- Response to climate evolution and sedimentary environment|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|date=2006|volume=240|issue=1–2|pages=184–203|doi=10.1016/j.palaeo.2006.03.059|bibcode=2006PPP...240..184S }}</ref> Subsequently, the Lower Permian had first seen the retreat of [[glacier]]s and the emergence of subpolar [[tundra]] and [[taiga]]-like vegetation (dominated by ''[[Botrychiopsis]]'' and ''[[Gangamopteris]]''),<ref name="Jasper&al2007">{{cite book|last=Jasper|first=A.|author2=Guerra-Sommer, M.|author3=Cazzulo-Klepzig, M.|author4=Iannuzzi, R.|year=2007|chapter=Biostratigraphic and paleoclimatic significance of ''Botrychiopsis'' fronds in the Gondwana realm|pages=379–388|title=Proceedings of the XVth International Congress on Carboniferous and Permian Stratigraphy, Utrecht, 10-16 August 2003|editor1-last=Wong|editor1-first=Th.E.|publisher=Edita KNAW|location=Amsterdam : Royal Netherlands Academy of Arts and Sciences|isbn=978-9069844794|url=http://sigep.cprm.gov.br/propostas/Afloramento_Quiteria_4_Jasper_etal2007.pdf}}</ref> then the introduction of warmer and wetter climatic conditions that allowed the development of the ''[[Mesosaurus]]'' fauna and the ''[[Glossopteris]]'' flora.<ref name="Scheffler & al.2006"/> The scientists who studied the climate of that time found very different results on the thermal ranges that existed in the ancient Karoo. At the end of the 1950s, [[Edna P. Plumstead|Edna Plumstead]] compared the Karoo to today's [[Siberia]] or [[Canada]], with a highly seasonal climate including very cold winters and temperate summers supporting the ''Glossopteris'' flora, which would have been restricted to sheltered basins.<ref name="Plumstead">{{cite book|last=Plumstead|first=E.|year=1957|title=Coal in southern Africa|editor1-last=Plumstead|editor1-first=E.|publisher=Witwatersrand University Press|location=Johannesburg}}</ref> Later, other studies, mainly based on [[climate model]]s, also suggested a cold temperate climate with high thermal amplitude between summer (+15 to +20&nbsp;°C) and winter (-20 to -25&nbsp;°C).<ref name="Kutzbach & Gallimore1989">{{cite journal|last1=Kutzbach|first1=J. E.|author2=Gallimore, R.G.|title=Pangean climates: megamonsoons of the megacontinent|journal=Journal of Geophysical Research|date=1989|volume=94|issue=D3|pages=3341–3357|doi=10.1029/JD094iD03p03341|bibcode=1989JGR....94.3341K }}</ref><ref name="Crowley & al.1989">{{cite journal|last1=Crowley|first1=T. J.|author2=Hyde, W.T.|author3=Short, D.A.|title=Seasonal cycle variations on the supercontinent of Pangaea|journal=Geology|date=1989|volume=17|issue=5|pages=457–460|doi=10.1130/0091-7613(1989)017<0457:SCVOTS>2.3.CO;2|bibcode=1989Geo....17..457C }}</ref> More recent studies also indicate a temperate climate, but with much less severe winters than those previously suggested. Keddy Yemane thus suggested that the vast river system and the many giant lakes present at the time throughout southern Africa must have significantly moderated the continentality of the Karoo climate during most of the Permian.<ref name="Yemane1993">{{cite journal|last1=Yemane|first1=K.|title=Contribution of the Late Permian palaeogeography in maintaining a temperate climate in Gondwana|journal=Nature|date=1993|volume=3651|issue=6407|pages=51–54|doi=10.1038/361051a0|bibcode=1993Natur.361...51Y |s2cid=4245546 }}</ref> [[Paleobotany|Paleobotanical]] studies focusing on the characteristic morphology of plant leaves and the [[Dendrochronology#growth rings|growth rings]] of [[fossil wood]]s also indicate a seasonal climate<ref name="Rayner1996"/><ref name="Bamford2016"/> with summer temperatures of up to 30&nbsp;°C and free-frost winters.<ref name="Rayner1996"/> According to Richard Rayner, the high southern latitudes experienced very hot and humid summers, with an average of 18 hours of light per day for more than four months during which precipitation was comparable to the annual amount falling in the present-day tropics. These conditions were extremely conducive to rapid growth in plants such as ''Glossopteris''.<ref name="Rayner1996"/> The habit in ''Glossopteris'' of losing its leaves at the beginning of the bad season would be linked to a shorter duration of daylight rather than the existence of very cold winter temperatures.<ref name="Rayner1996"/> From the [[Geochemistry|geochemical]] study of sediments from several Karoo sites, Kay Scheffler also obtains a temperate climate (with mean annual temperatures of about 15 to 20&nbsp;°C), with free-frost winter, but with an increase in aridity during the Middle Permian.<ref name="Scheffler & al.2006"/>
+At the time of ''Anteosaurus'', most of the landmasses were united in one supercontinent, [[Pangaea]]. It 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 Ocean|Tethys Sea]].<ref name="McLoughlin2001">{{cite journal|last1=McLoughlin|first1=S.|title=The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism|journal=Australian Journal of Botany|date=2001|volume=49|issue=3|pages=271–300|doi=10.1071/BT00023}}</ref> A long string of microcontinents, grouped under the name of [[Cimmeria (continent)|Cimmeria]], divided the Tethys in two : the Paleo-Tethys in the north, and the Neo-Tethys in the south.<ref name="Şengör1987">{{cite journal|last1=Şengör|first1=A.M.C.|title=Tectonics of the Tethysides: orogenic collage development in a collisional setting|journal=Annual Review of Earth and Planetary Sciences|date=1987|volume=15|issue=|pages=214–244|doi=10.1146/annurev.ea.15.050187.001241|bibcode=1987AREPS..15..213C }}</ref> The territory that would become the South African Karoo was located much further south than today, at the level of the [[60th parallel south]].<ref name="Bamford2016">{{cite book|last=Bamford|first=M.K.|year=2016|chapter=Fossil woods from the Upper Carboniferous to Lower Jurassic Karoo Basin and their environmental interpretation|pages=159–167|title=Origin and evolution of the Cape Mountains and Karoo Basin|editor1-last=Linol|editor1-first=B.|editor2=de Wit, M.J.|publisher=Regional Geology Reviews, Springer|location=Cham|isbn=978-3-319-40858-3}}</ref> Although located close to the [[Antarctic Circle]], the climate prevailing at this [[latitude]] during most of the Permian was [[Temperate climate|temperate]] with distinct seasons.<ref name="king1990">{{cite book|last=King|first=G.|year=1990|chapter=The environment of the ''Tapinocephalus'' zone|pages=107–111|title=The dicynodonts. A study in palaeobiology|editor1-last=King|editor1-first=G.|publisher=Chapman and Hall|location=London and New York|isbn=0-412-33080-6}}</ref><ref name="Rayner1996">{{cite journal|last1=Rayner|first1=R.J.|title=The palaeoclimate of the Karoo: evidence from plant fossils|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|date=1996|volume=119|issue=3–4|pages=385–394|doi=10.1016/0031-0182(95)00021-6|bibcode=1996PPP...119..385R }}</ref><ref name="Chumakov&Zharkov2003">{{cite journal|last1=Chumakov|first1=N.M.|author2=Zharkov, M.A.|title=Climate during the Permian-Triassic Biosphere Reorganizations. Article 2. Climate of the Late Permian and Early Triassic : General Inferences|journal=Stratigraphy and Geological Correlation|date=2003|volume=11|issue=4|pages=361–376}}</ref>  There are uncertainties about the temperatures that prevailed in South Africa during the Middle Permian. Previously, this region of the world had undergone significant [[Karoo Ice Age|glaciation]] during the Upper [[Carboniferous]].<ref name="Scheffler & al.2006">{{cite journal|last1=Scheffler|first1=K.|author2=Buehmann, D.|author3=Schwark, L.|title=Analysis of Late Palaeozoic glacial to potglacial sedimentary successions in South Africa by geochemical proxies- Response to climate evolution and sedimentary environment|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|date=2006|volume=240|issue=1–2|pages=184–203|doi=10.1016/j.palaeo.2006.03.059|bibcode=2006PPP...240..184S }}</ref> Subsequently, the Lower Permian had first seen the retreat of [[glacier]]s and the emergence of subpolar [[tundra]] and [[taiga]]-like vegetation (dominated by ''[[Botrychiopsis]]'' and ''[[Gangamopteris]]''),<ref name="Jasper&al2007">{{cite book|last=Jasper|first=A.|author2=Guerra-Sommer, M.|author3=Cazzulo-Klepzig, M.|author4=Iannuzzi, R.|year=2007|chapter=Biostratigraphic and paleoclimatic significance of ''Botrychiopsis'' fronds in the Gondwana realm|pages=379–388|title=Proceedings of the XVth International Congress on Carboniferous and Permian Stratigraphy, Utrecht, 10-16 August 2003|editor1-last=Wong|editor1-first=Th.E.|publisher=Edita KNAW|location=Amsterdam : Royal Netherlands Academy of Arts and Sciences|isbn=978-9069844794|url=http://sigep.cprm.gov.br/propostas/Afloramento_Quiteria_4_Jasper_etal2007.pdf|access-date=31 October 2022|archive-date=31 October 2022|archive-url=https://web.archive.org/web/20221031185031/http://sigep.cprm.gov.br/propostas/Afloramento_Quiteria_4_Jasper_etal2007.pdf|url-status=dead}}</ref> then the introduction of warmer and wetter climatic conditions that allowed the development of the ''[[Mesosaurus]]'' fauna and the ''[[Glossopteris]]'' flora.<ref name="Scheffler & al.2006"/> The scientists who studied the climate of that time found very different results on the thermal ranges that existed in the ancient Karoo. At the end of the 1950s, [[Edna P. Plumstead|Edna Plumstead]] compared the Karoo to today's [[Siberia]] or [[Canada]], with a highly seasonal climate including very cold winters and temperate summers supporting the ''Glossopteris'' flora, which would have been restricted to sheltered basins.<ref name="Plumstead">{{cite book|last=Plumstead|first=E.|year=1957|title=Coal in southern Africa|editor1-last=Plumstead|editor1-first=E.|publisher=Witwatersrand University Press|location=Johannesburg}}</ref> Later, other studies, mainly based on [[climate model]]s, also suggested a cold temperate climate with high thermal amplitude between summer (+15 to +20&nbsp;°C) and winter (-20 to -25&nbsp;°C).<ref name="Kutzbach & Gallimore1989">{{cite journal|last1=Kutzbach|first1=J. E.|author2=Gallimore, R.G.|title=Pangean climates: megamonsoons of the megacontinent|journal=Journal of Geophysical Research|date=1989|volume=94|issue=D3|pages=3341–3357|doi=10.1029/JD094iD03p03341|bibcode=1989JGR....94.3341K }}</ref><ref name="Crowley & al.1989">{{cite journal|last1=Crowley|first1=T. J.|author2=Hyde, W.T.|author3=Short, D.A.|title=Seasonal cycle variations on the supercontinent of Pangaea|journal=Geology|date=1989|volume=17|issue=5|pages=457–460|doi=10.1130/0091-7613(1989)017<0457:SCVOTS>2.3.CO;2|bibcode=1989Geo....17..457C }}</ref> More recent studies also indicate a temperate climate, but with much less severe winters than those previously suggested. Keddy Yemane thus suggested that the vast river system and the many giant lakes present at the time throughout southern Africa must have significantly moderated the continentality of the Karoo climate during most of the Permian.<ref name="Yemane1993">{{cite journal|last1=Yemane|first1=K.|title=Contribution of the Late Permian palaeogeography in maintaining a temperate climate in Gondwana|journal=Nature|date=1993|volume=3651|issue=6407|pages=51–54|doi=10.1038/361051a0|bibcode=1993Natur.361...51Y |s2cid=4245546 }}</ref> [[Paleobotany|Paleobotanical]] studies focusing on the characteristic morphology of plant leaves and the [[Dendrochronology#growth rings|growth rings]] of [[fossil wood]]s also indicate a seasonal climate<ref name="Rayner1996"/><ref name="Bamford2016"/> with summer temperatures of up to 30&nbsp;°C and free-frost winters.<ref name="Rayner1996"/> According to Richard Rayner, the high southern latitudes experienced very hot and humid summers, with an average of 18 hours of light per day for more than four months during which precipitation was comparable to the annual amount falling in the present-day tropics. These conditions were extremely conducive to rapid growth in plants such as ''Glossopteris''.<ref name="Rayner1996"/> The habit in ''Glossopteris'' of losing its leaves at the beginning of the bad season would be linked to a shorter duration of daylight rather than the existence of very cold winter temperatures.<ref name="Rayner1996"/> From the [[Geochemistry|geochemical]] study of sediments from several Karoo sites, Kay Scheffler also obtains a temperate climate (with mean annual temperatures of about 15 to 20&nbsp;°C), with free-frost winter, but with an increase in aridity during the Middle Permian.<ref name="Scheffler & al.2006"/>
 ===Paleoecology===
@@ Line 128: / Line 128: @@
 The sediments of the Abrahamskraal Formation consists of a succession of [[sandstone]]s, and versicolor [[siltstone]]s and [[Mudrock#Mudstone|mudstones]], deposited by large rivers that flowed from south to north from the Gondwanide mountain range. These large rivers of variable sinuosity drained a vast alluvial plain that sloped gently down to the northeast toward the Ecca sea (a former landlocked sea), while in receding phase.<ref name="turner1978">{{cite book|last=Turner|first=B.R.|year=1978|chapter=Sedimentary patterns of uranium mineralization in the Beaufort Group of the southern Karoo (Gondwana) Basin, South Africa|pages=831–848|title=Fluvial Sedimentology|editor1-last=Miall|editor1-first=A.D.|publisher=Canadian Association of Petroleum Geologists, Memoir 5}}</ref><ref name="smith1990">{{cite journal|last1=Smith|first1=R.M.H.|title=A review of the stratigraphy and sedimentary environments of the Karoo basin of South Africa|journal=Journal of African Earth Sciences|date=1990|volume=10|issue=1|pages=117–137|doi=10.1016/0899-5362(90)90050-O |bibcode=1990JAfES..10..117S }}</ref><ref name="jirah2014">{{cite journal |last1=Jirah|first1=S.|author2=Rubidge, B.S.|year=2014|title=Refined stratigraphy of the Middle Permian Abrahamskraal Formation (Beaufort Group) in the southern Karoo Basin|journal=Journal of African Earth Sciences|volume=100|issue=|pages=121–135|doi=10.1016/j.jafrearsci.2014.06.014|bibcode=2014JAfES.100..121J }}</ref><ref name="Cole&al.2016">{{cite journal|last1=Cole|first1=D.I.|author2=Johnson, M.R.|author3=Day, M.O.|year=2016|title=Lithostratigraphy of the Abrahamskraal Formation (Karoo Supergroup), South Africa|journal=South African Journal of Geology|volume=119|issue=2|pages=415–424|doi=10.2113/gssajg.119.2.415|bibcode=2016SAJG..119..415C }}</ref> The landscape was composed of marshy land, interrupted by rivers, lakes, woods and forests. Many fossil traces (footprints, [[ripple marks]], [[mudcrack]]s) indicate that swampy areas, which were the most extensive habitat, were frequently exposed to the open air and should not often be deeply flooded.<ref name="king1990"/> The vegetation was dominated by the [[deciduous]] [[Pteridospermatophyta|pteridosperm]] ''[[Glossopteris]]'', which formed woodlands and large forests concentrated along the streams and on the uplands.<ref name="king1990"/><ref name="Rayner1996"/> Large [[Equisetum|horsetails]] ({{convert|2|-|3|m|ft|abbr=on}} high), such as ''[[Schizoneura]]'' and ''[[Paraschizoneura]]'', formed bamboo-like stands that grew in and around swamps. Herbaceous horsetails (''[[Phyllotheca]]'') and [[fern]]s carpeted the undergrowth and small [[Lycopodiophyta|lycopods]] occupied the wetter areas.<ref name="king1990"/><ref name="Rayner1996"/>
-Aquatic fauna included the lamellibranch ''[[Palaeomutela]]'', the [[palaeonisciformes]] fishes ''[[Atherstonia]]'', ''[[Bethesdaichthys]]'', ''[[Blourugia]]'', ''[[Namaichthys]]'' and ''[[Westlepis]]'', and large freshwater predators, the [[Temnospondyli|temnospondyl]] [[amphibian]]s ''[[Rhinesuchoides]]'' and ''[[Rhinesuchus]]''.<ref name="damiani2003">{{cite journal|last1=Damiani|first1=R.J.|author2=Rubidge, B.S.|title=A review of the South African temnospondyl amphibian record|journal=Palaeontologia Africana|date=2003|volume=39|issue=|pages=21–36|url=http://wiredspace.wits.ac.za/handle/10539/13304}}</ref><ref name="Marsicano&al.2017">{{cite journal|last=Marsicano|first=C.A.|author2=Latimer, E.|author3=Rubidge, B.|author4=Smith, R.M.H.|title=The Rhinesuchidae and early history of the Stereospondyli (Amphibia: Temnospondyli) at the end of the Palaeozoic|journal=Journal of Vertebrate Paleontology|year=2017|volume=181|issue=2|pages=1–28|doi=10.1093/zoolinnean/zlw032}}</ref> The terrestrial fauna was particularly diverse and dominated by the therapsids.{{refn|group=nb|Several taxa from the lower part of the ''Tapinocephalus'' zone, such as the biarmosuchian ''[[Pachydectes]]'', the dicynodont ''[[Lanthanostegus]]'', and the gorgonopsian ''[[Phorcys dubei|Phorcys]]'', have been found in strata located several hundred meters below the oldest known specimens of ''Anteosaurus''. This particular fauna could represent a new zone or sub-zone not yet recognized as such and located between the ''Eodicynodon'' and ''Tapinocephalus'' Assemblage Zones (Day & Rubidge 2020, Rubidge & al. 2021).}} ''Anteosaurus'' occupied the top of the food chain there. It shared its environment with many other carnivorous [[tetrapod]]s. Other large predatory animals included the lion-sized [[Lycosuchidae|Lycosuchid]] [[therocephalia]]ns ''[[Lycosuchus]]'' and ''[[Simorhinella]]'',<ref name="abdala2014">{{cite journal|last1=Abdala|first1=F.|author2=Kammerer, C.F.|author3=Day, M.O.|author4=Rubidge, B.S.|title=Adult morphology of the therocephalian ''Simorhinella baini'' from the middle Permian of South Africa and the taxonomy, paleobiogeography, and temporal distribution of the Lycosuchidae|journal=Journal of Paleontology|date=2014|volume=88|issue=6|pages=1139–1153|doi=10.1666/13-186|bibcode=2014JPal...88.1139A |s2cid=129323281 }}</ref> and the [[Scylacosauridae|Scylacosaurid]] therocephalian ''[[Glanosuchus]]''.<ref name="abdala2008">{{cite journal |last1=Abdala|first1=F.|author2=Rubidge, B.S.|author3=van den Heever, J.|year=2008|title=The oldest Therocephalians (Therapsida, Eutheriodontia) and the early diversification of Therapsida|journal=Palaeontology|volume=51|issue=4|pages=1011–1024|doi=10.1111/j.1475-4983.2008.00784.x|bibcode=2008Palgy..51.1011A |s2cid=129791548 |doi-access=free}}</ref> Medium-sized carnivorous were represented by the basal biarmosuchian ''[[Hipposaurus]]'',<ref name=Boonstra1969/> the more derived biarmosuchian ''[[Bullacephalus]]'',<ref name=rubidge2003>{{cite journal |last=Rubidge |first=B.S. |author2=Kitching, J.W. |year=2003 |title=A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa |journal=Palaeontology |volume=46 |issue=1 |pages=199–210 |doi=10.1111/1475-4983.00294 |bibcode=2003Palgy..46..199R |s2cid=129213631 |url=http://onlinelibrary.wiley.com/doi/10.1111/1475-4983.00294/pdf}}</ref> the scylacosaurids ''[[Ictidosaurus]]'',<ref name="abdala2008"/> ''[[Scylacosaurus]]'',<ref name="smith2012">{{cite book|last=Smith|first=R.|author2=Rubidge, B.|author3=van de Walt, M.|year=2012|chapter=Therapsid biodiversity patterns and paleoenvironments of the Karoo Basin, South Africa.|pages=31–62|title=Forerunners of Mammals : Radiation histology biology.|editor1-last=Chinsamy-Turan|editor1-first=A.|publisher=Indiana University Press.|location=Bloomington and Indianapolis|isbn=978-0-253-35697-0}}</ref> and ''[[Pristerognathus]]'',<ref name="Day&Rubidge2020"/> and the small and basal [[gorgonopsia]]n ''[[Eriphostoma]]''.<ref name="kammerer2013">{{cite book|last=Kammerer|first=C.F.|year=2013|chapter=A redescription of ''Eriphostoma microdon'' Broom, 1911 (Therapsida, Gorgonopsia) from the ''Tapinocephalus'' Assemblage Zone of South Africa and a review of Middle Permian gorgonopsians|pages=171–184|title=The early evolutionary history of the Synapsida|editor1-last=Kammerer|editor1-first=C.F.|editor2= Angielszyk, K.D.|editor3=Fröbisch, J.|publisher=Dordrecht: Springer|isbn=978-9400768406}}</ref><ref name="kammerer2015">{{cite journal|last1=Kammerer|first1=C.F.|author2=Smith, R.M.H.|author3=Day, M.O.|author4=Rubidge, B.S.|title=New information on the morphology and stratigraphic range of the mid-Permian gorgonopsian ''Eriphostoma microdon'' Broom, 1911|journal=Papers in Palaeontology|date=2015|volume=1|issue=2|pages=201–221|doi=10.1002/spp2.1012|s2cid=128762256 }}</ref> The small predator guild (mainly insectivorous forms) included the therocephalians ''[[Alopecodon]]'',<ref name="Day&Rubidge2020"/> and ''[[Pardosuchus]]'',<ref name="Day&Rubidge2020"/> the small monitor-like [[Varanopidae|varanopids]] ''[[Elliotsmithia]]'',<ref name="reisz1998">{{cite journal|last1=Reisz|first1=R.R.|author2=Dilkes, D.W.|author3=Berman, D.S.|title=Anatomy and relationships of Elliotsmithia longiceps Broom, a small synapsid (Eupelycosauria, Varanopseidae) from the Late Permian of South Africa|journal=Journal of Vertebrate Paleontology|date=1998|volume=18|issue=3|pages=602–611|doi=10.1080/02724634.1998.10011087 |bibcode=1998JVPal..18..602R }}</ref><ref name="smith2012"/> ''[[Heleosaurus]]'',<ref name="botha-brink2007">{{cite journal|last1=Botha-Brink|first1=J.|author2=Modesto, S.P.|title=A mixed-age classed "pelycosaur" aggregation from South Africa: earliest evidence of parental care in amniotes?|journal=Proceedings of the Royal Society B: Biological Sciences|date=2007|volume=274|issue=1627|pages=2829–2834|doi=10.1098/rspb.2007.0803|pmid=17848370 |pmc=2288685 }}</ref><ref name="reisz2007">{{cite journal|last1=Reisz|first1=R.R.|author2=Modesto, S.P.|title=Heleosaurus sholtzi from the Permian of South Africa: A Varanopid Synapsid, Not a Diapsid Reptile|journal=Journal of Vertebrate Paleontology|date=2007|volume=27|issue=3|pages=734–739|doi=10.1671/0272-4634(2007)27[734:HSFTPO]2.0.CO;2|s2cid=86246959 }}</ref><ref name="botha-brink2009">{{cite journal|last1=Botha-Brink|first1=J.|author2=Modesto, S.P.|title=Anatomy and relationships of the Middle Permian Varanopid Heleosaurus scholtzi based on a social aggregation from the Karoo basin of South Africa|journal=Journal of Vertebrate Paleontology|date=2009|volume=29|issue=2|pages=389–400|doi=10.1671/039.029.0209|bibcode=2009JVPal..29..389B |s2cid=84703110 }}</ref> and ''[[Microvaranops]]'',<ref name="spindler&al2018">{{cite journal|last=Spindler|first=F.|author2=Werneburg, R.|author3=Schneider, J.W.|author4=Luthardt, L.|author5=Annacker, V.|author6=Rößler, R.|title=First arboreal 'pelycosaur' (Synapsida : Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny|journal=PalZ|year=2018|volume=92|issue=2|pages=315–364|doi=10.1007/s12542-018-0405-9|bibcode=2018PalZ...92..315S |s2cid=133846070 }}</ref> the [[Millerettidae|millerettid]] ''[[Broomia]]'',<ref name="Day&Rubidge2020"/> the [[Procolophonomorpha|procolophonomorph]] ''[[Australothyris]]'',<ref name=Modesto&al.2009>{{Cite journal|last=Modesto|first=S.P.|author2=Scott, D.M.|author3=Reisz, R.R.|year=2009|title=A new parareptile with temporal fenestration from the Middle Permian of South Africa|journal=Canadian Journal of Earth Sciences|volume=46|issue=1|pages=9–20|doi=10.1139/E09-001|bibcode=2009CaJES..46....9M }}</ref> and the lizard-like ''[[Eunotosaurus]]''<ref name=rubidge1999>{{Cite journal|last=Rubidge|first=B.S.|author2=Modesto, S.|author3=Sidor, C.|author4=Welman, J.|year=1999|title=''Eunotosaurus africanus'' from the Ecca–Beaufort contact in Northern Cape Province, South Africa — implications for Karoo Basin development|journal=South African Journal of Science|volume=95|issue=|pages=553–555|url=http://iris.nyit.edu/nycom/Faculty/Profiles/Sidor/Rubidge_etal1999.pdf|access-date=2022-10-31|archive-date=2011-07-16|archive-url=https://web.archive.org/web/20110716133817/http://iris.nyit.edu/nycom/Faculty/Profiles/Sidor/Rubidge_etal1999.pdf|url-status=dead}}</ref> of uncertain affinities (variously considered as a [[Parareptilia|parareptile]],<ref name="modesto2000">{{cite journal |last1=Modesto|first1=S.P.|year=2000|title=''Eunotosaurus africanus'' and the Gondwanan ancestry of anapsid reptiles|journal=Palaeontologia Africana|volume=36|issue=|pages=15–20}}</ref> a [[Pantestudines|pantestudine]]<ref name="bever2015">{{cite journal |last1=Bever|first1=G.S.|author2=Lyson, T.R.|author3=Field, D.J.|author4=Bhullar, B-A.S.|year=2015|title=Evolutionary origin of the turtle skull|journal=Nature|volume=525|issue=7568|pages=239–242|doi=10.1038/nature14900|pmid=26331544 |bibcode=2015Natur.525..239B |s2cid=4401555 }}</ref><ref name="lyson2016">{{cite journal |last1=Lyson|first1=T.R|author2=Rubidge, B.S.|author3=Scheyer, T.M.|author4=de Queiroz, K.|author5=Schachner, E.R.|author6=Smith, R.M.H.|author7=Botha-Brink, J.|author8=Bever, G.S.|year=2016|title=Fossorial origin of the turtle shell|journal=Current Biology|volume=26|issue=14|pages=1887–1894|doi=10.1016/j.cub.2016.05.020|pmid=27426515 |s2cid=3935231 |doi-access=free}}</ref> or a [[Caseidae|caseid]] synapsid<ref name="Lichtig&Lucas2021">{{cite journal|last1=Lichtig|first1=A.J.|author2=Lucas, S.G.|title=''Chinlechelys'' from the Upper Triassic of New Mexico, USA, and the oigin of turtles|journal=Palaeontologia Electronica|date=2021|volume=24|issue=1|pages=a13|doi=10.26879/886|s2cid=233454789 |doi-access=free}}</ref>).
+Aquatic fauna included the lamellibranch ''[[Palaeomutela]]'', the [[palaeonisciformes]] fishes ''[[Atherstonia]]'', ''[[Bethesdaichthys]]'', ''[[Blourugia]]'', ''[[Namaichthys]]'' and ''[[Westlepis]]'', and large freshwater predators, the [[Temnospondyli|temnospondyl]] [[amphibian]]s ''[[Rhinesuchoides]]'' and ''[[Rhinesuchus]]''.<ref name="damiani2003">{{cite journal|last1=Damiani|first1=R.J.|author2=Rubidge, B.S.|title=A review of the South African temnospondyl amphibian record|journal=Palaeontologia Africana|date=2003|volume=39|issue=|pages=21–36|url=http://wiredspace.wits.ac.za/handle/10539/13304}}</ref><ref name="Marsicano&al.2017">{{cite journal|last=Marsicano|first=C.A.|author2=Latimer, E.|author3=Rubidge, B.|author4=Smith, R.M.H.|title=The Rhinesuchidae and early history of the Stereospondyli (Amphibia: Temnospondyli) at the end of the Palaeozoic|journal=Journal of Vertebrate Paleontology|year=2017|volume=181|issue=2|pages=1–28|doi=10.1093/zoolinnean/zlw032|hdl=11336/105150|hdl-access=free}}</ref> The terrestrial fauna was particularly diverse and dominated by the therapsids.{{refn|group=nb|Several taxa from the lower part of the ''Tapinocephalus'' zone, such as the biarmosuchian ''[[Pachydectes]]'', the dicynodont ''[[Lanthanostegus]]'', and the gorgonopsian ''[[Phorcys dubei|Phorcys]]'', have been found in strata located several hundred meters below the oldest known specimens of ''Anteosaurus''. This particular fauna could represent a new zone or sub-zone not yet recognized as such and located between the ''Eodicynodon'' and ''Tapinocephalus'' Assemblage Zones (Day & Rubidge 2020, Rubidge & al. 2021).}} ''Anteosaurus'' occupied the top of the food chain there. It shared its environment with many other carnivorous [[tetrapod]]s. Other large predatory animals included the lion-sized [[Lycosuchidae|Lycosuchid]] [[therocephalia]]ns ''[[Lycosuchus]]'' and ''[[Simorhinella]]'',<ref name="abdala2014">{{cite journal|last1=Abdala|first1=F.|author2=Kammerer, C.F.|author3=Day, M.O.|author4=Rubidge, B.S.|title=Adult morphology of the therocephalian ''Simorhinella baini'' from the middle Permian of South Africa and the taxonomy, paleobiogeography, and temporal distribution of the Lycosuchidae|journal=Journal of Paleontology|date=2014|volume=88|issue=6|pages=1139–1153|doi=10.1666/13-186|bibcode=2014JPal...88.1139A |s2cid=129323281 }}</ref> and the [[Scylacosauridae|Scylacosaurid]] therocephalian ''[[Glanosuchus]]''.<ref name="abdala2008">{{cite journal |last1=Abdala|first1=F.|author2=Rubidge, B.S.|author3=van den Heever, J.|year=2008|title=The oldest Therocephalians (Therapsida, Eutheriodontia) and the early diversification of Therapsida|journal=Palaeontology|volume=51|issue=4|pages=1011–1024|doi=10.1111/j.1475-4983.2008.00784.x|bibcode=2008Palgy..51.1011A |s2cid=129791548 |doi-access=free}}</ref> Medium-sized carnivorous were represented by the basal biarmosuchian ''[[Hipposaurus]]'',<ref name=Boonstra1969/> the more derived biarmosuchian ''[[Bullacephalus]]'',<ref name=rubidge2003>{{cite journal |last=Rubidge |first=B.S. |author2=Kitching, J.W. |year=2003 |title=A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa |journal=Palaeontology |volume=46 |issue=1 |pages=199–210 |doi=10.1111/1475-4983.00294 |bibcode=2003Palgy..46..199R |s2cid=129213631 |url=http://onlinelibrary.wiley.com/doi/10.1111/1475-4983.00294/pdf}}</ref> the scylacosaurids ''[[Ictidosaurus]]'',<ref name="abdala2008"/> ''[[Scylacosaurus]]'',<ref name="smith2012">{{cite book|last=Smith|first=R.|author2=Rubidge, B.|author3=van de Walt, M.|year=2012|chapter=Therapsid biodiversity patterns and paleoenvironments of the Karoo Basin, South Africa.|pages=31–62|title=Forerunners of Mammals : Radiation histology biology.|editor1-last=Chinsamy-Turan|editor1-first=A.|publisher=Indiana University Press.|location=Bloomington and Indianapolis|isbn=978-0-253-35697-0}}</ref> and ''[[Pristerognathus]]'',<ref name="Day&Rubidge2020"/> and the small and basal [[gorgonopsia]]n ''[[Eriphostoma]]''.<ref name="kammerer2013">{{cite book|last=Kammerer|first=C.F.|year=2013|chapter=A redescription of ''Eriphostoma microdon'' Broom, 1911 (Therapsida, Gorgonopsia) from the ''Tapinocephalus'' Assemblage Zone of South Africa and a review of Middle Permian gorgonopsians|pages=171–184|title=The early evolutionary history of the Synapsida|editor1-last=Kammerer|editor1-first=C.F.|editor2= Angielszyk, K.D.|editor3=Fröbisch, J.|publisher=Dordrecht: Springer|isbn=978-9400768406}}</ref><ref name="kammerer2015">{{cite journal|last1=Kammerer|first1=C.F.|author2=Smith, R.M.H.|author3=Day, M.O.|author4=Rubidge, B.S.|title=New information on the morphology and stratigraphic range of the mid-Permian gorgonopsian ''Eriphostoma microdon'' Broom, 1911|journal=Papers in Palaeontology|date=2015|volume=1|issue=2|pages=201–221|doi=10.1002/spp2.1012|bibcode=2015PPal....1..201K |s2cid=128762256 }}</ref> The small predator guild (mainly insectivorous forms) included the therocephalians ''[[Alopecodon]]'',<ref name="Day&Rubidge2020"/> and ''[[Pardosuchus]]'',<ref name="Day&Rubidge2020"/> the small monitor-like [[Varanopidae|varanopids]] ''[[Elliotsmithia]]'',<ref name="reisz1998">{{cite journal|last1=Reisz|first1=R.R.|author2=Dilkes, D.W.|author3=Berman, D.S.|title=Anatomy and relationships of ''Elliotsmithia longiceps'' Broom, a small synapsid (Eupelycosauria, Varanopseidae) from the Late Permian of South Africa|journal=Journal of Vertebrate Paleontology|date=1998|volume=18|issue=3|pages=602–611|doi=10.1080/02724634.1998.10011087 |bibcode=1998JVPal..18..602R }}</ref><ref name="smith2012"/> ''[[Heleosaurus]]'',<ref name="botha-brink2007">{{cite journal|last1=Botha-Brink|first1=J.|author2=Modesto, S.P.|title=A mixed-age classed "pelycosaur" aggregation from South Africa: earliest evidence of parental care in amniotes?|journal=Proceedings of the Royal Society B: Biological Sciences|date=2007|volume=274|issue=1627|pages=2829–2834|doi=10.1098/rspb.2007.0803|pmid=17848370 |pmc=2288685 }}</ref><ref name="reisz2007">{{cite journal|last1=Reisz|first1=R.R.|author2=Modesto, S.P.|title=''Heleosaurus sholtzi'' from the Permian of South Africa: A Varanopid Synapsid, Not a Diapsid Reptile|journal=Journal of Vertebrate Paleontology|date=2007|volume=27|issue=3|pages=734–739|doi=10.1671/0272-4634(2007)27[734:HSFTPO]2.0.CO;2|s2cid=86246959 }}</ref><ref name="botha-brink2009">{{cite journal|last1=Botha-Brink|first1=J.|author2=Modesto, S.P.|title=Anatomy and relationships of the Middle Permian Varanopid ''Heleosaurus scholtzi'' based on a social aggregation from the Karoo basin of South Africa|journal=Journal of Vertebrate Paleontology|date=2009|volume=29|issue=2|pages=389–400|doi=10.1671/039.029.0209|bibcode=2009JVPal..29..389B |s2cid=84703110 }}</ref> and ''[[Microvaranops]]'',<ref name="spindler&al2018">{{cite journal|last=Spindler|first=F.|author2=Werneburg, R.|author3=Schneider, J.W.|author4=Luthardt, L.|author5=Annacker, V.|author6=Rößler, R.|title=First arboreal 'pelycosaur' (Synapsida : Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny|journal=PalZ|year=2018|volume=92|issue=2|pages=315–364|doi=10.1007/s12542-018-0405-9|bibcode=2018PalZ...92..315S |s2cid=133846070 }}</ref> the [[Millerettidae|millerettid]] ''[[Broomia]]'',<ref name="Day&Rubidge2020"/> the [[Procolophonomorpha|procolophonomorph]] ''[[Australothyris]]'',<ref name=Modesto&al.2009>{{Cite journal|last=Modesto|first=S.P.|author2=Scott, D.M.|author3=Reisz, R.R.|year=2009|title=A new parareptile with temporal fenestration from the Middle Permian of South Africa|journal=Canadian Journal of Earth Sciences|volume=46|issue=1|pages=9–20|doi=10.1139/E09-001|bibcode=2009CaJES..46....9M }}</ref> and the lizard-like ''[[Eunotosaurus]]''<ref name=rubidge1999>{{Cite journal|last=Rubidge|first=B.S.|author2=Modesto, S.|author3=Sidor, C.|author4=Welman, J.|year=1999|title=''Eunotosaurus africanus'' from the Ecca–Beaufort contact in Northern Cape Province, South Africa — implications for Karoo Basin development|journal=South African Journal of Science|volume=95|issue=|pages=553–555|url=http://iris.nyit.edu/nycom/Faculty/Profiles/Sidor/Rubidge_etal1999.pdf|access-date=2022-10-31|archive-date=2011-07-16|archive-url=https://web.archive.org/web/20110716133817/http://iris.nyit.edu/nycom/Faculty/Profiles/Sidor/Rubidge_etal1999.pdf|url-status=dead}}</ref> of uncertain affinities (variously considered as a [[Parareptilia|parareptile]],<ref name="modesto2000">{{cite journal |last1=Modesto|first1=S.P.|year=2000|title=''Eunotosaurus africanus'' and the Gondwanan ancestry of anapsid reptiles|journal=Palaeontologia Africana|volume=36|issue=|pages=15–20}}</ref> a [[Pantestudines|pantestudine]]<ref name="bever2015">{{cite journal |last1=Bever|first1=G.S.|author2=Lyson, T.R.|author3=Field, D.J.|author4=Bhullar, B-A.S.|year=2015|title=Evolutionary origin of the turtle skull|journal=Nature|volume=525|issue=7568|pages=239–242|doi=10.1038/nature14900|pmid=26331544 |bibcode=2015Natur.525..239B |s2cid=4401555 }}</ref><ref name="lyson2016">{{cite journal |last1=Lyson|first1=T.R|author2=Rubidge, B.S.|author3=Scheyer, T.M.|author4=de Queiroz, K.|author5=Schachner, E.R.|author6=Smith, R.M.H.|author7=Botha-Brink, J.|author8=Bever, G.S.|year=2016|title=Fossorial origin of the turtle shell|journal=Current Biology|volume=26|issue=14|pages=1887–1894|doi=10.1016/j.cub.2016.05.020|pmid=27426515 |s2cid=3935231 |doi-access=free|bibcode=2016CBio...26.1887L }}</ref> or a [[Caseidae|caseid]] synapsid<ref name="Lichtig&Lucas2021">{{cite journal|last1=Lichtig|first1=A.J.|author2=Lucas, S.G.|title=''Chinlechelys'' from the Upper Triassic of New Mexico, USA, and the oigin of turtles|journal=Palaeontologia Electronica|date=2021|volume=24|issue=1|pages=a13|doi=10.26879/886|s2cid=233454789 |doi-access=free}}</ref>).
 [[File:Moschops11DB.jpg|thumb|right|''[[Moschops|Moschops capensis]]'', a large [[Tapinocephalidae]] that coexisted with ''Anteosaurus''.]]
-Herbivorous were also numerous and diversified. Large-sized vegetarians were mainly represented by numerous dinocephalians including the tapinocephalids ''[[Agnosaurus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020">{{Cite thesis |degree=Ph.D. |title= Taxonomic Revision of the Short-Snouted Tapinocephalid Dinocephalia (Amniota-Therapsida) The Key to Understanding Middle Permian Tetrapod Biodiversity |url= https://wiredspace.wits.ac.za/handle/10539/31602 |last=Neumann |first=S. |year=2020 |publisher= University of the Witwatersrand |location=Johannesburg |pages=1–411}}</ref> ''[[Criocephalosaurus]]'',<ref name="broom1928">{{cite journal |last1=Broom|first1=R.|year=1928|title=On ''Tapinocephalus'' and two other dinocephalians|journal=Annals of the South African Museum|volume=22|issue=|pages=427–438}}</ref><ref name="kammerer2002">{{cite journal |last1=Kammerer|first1=C.|author2=Sidor, C.A.|year=2002|title=Replacement names for the therapsid genera ''Criocephalus'' Boom 1928 and ''Oliviera'' Brink 1965|journal=Palaeontologia Africana|volume=38|issue=|pages=71–72|url=http://wiredspace.wits.ac.za/handle/10539/16356}}</ref><ref name="day2015b"/><ref name="Neumann2020"/> ''[[Mormosaurus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020"/> ''[[Moschognathus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020"/> ''[[Moschops]]'',<ref name="gregory1926">{{cite journal|last1=Gregory|first1=W.K.|year=1926|title=The skeleton of ''Moschops capensis'' Broom, a Dinocephalian Reptile from the Permian of South Africa|journal=Bulletin of the American Museum of Natural History|volume=56|issue=3|pages=179–251|hdl=2246/1323 |url=http://digitallibrary.amnh.org/handle/2246/1323}}</ref><ref name="boonstra1957">{{cite journal|last1=Boonstra|first1=L.D.|year=1957|title=The moschopid skulls in the South African Museum|journal=Annals of the South African Museum|volume=44|issue=|pages=15–38|url=http://www.biodiversitylibrary.org/page/40862867#page/33/mode/1up}}</ref><ref name="boonstra1963">{{cite journal |last1=Boonstra|first1=L.D.|year=1963|title=Diversity within the South African Dinocephalia|journal=South African Journal of Science|volume=59|issue=|pages=196–207}}</ref><ref name="Neumann2020"/> ''[[Riebeeckosaurus]]'',<ref name="boonstra1957"/><ref name="güven2013">{{cite journal |last1=Güven|first1=S.|author2=Rubidge, B.S.|author3=Abdala, F.|year=2013|title=Cranial morphology and taxonomy of South African Tapinocephalidae (Therapsida, Dinocephalia): the case of ''Avenantia'' and ''Riebeeckosaurus''|journal=Palaeontologia Africana|volume=48|issue=|pages=24–33|url=http://wiredspace.wits.ac.za/handle/10539/13310}}</ref> ''[[Struthiocephalus]]''<ref name="boonstra1953b">{{cite journal |last1=Boonstra|first1=L.D.|year=1953|title=The cranial morphology and taxonomy of the tapinocephalid genus ''Struthiocephalus''|journal=Annals of the South African Museum|volume=42|issue=|pages=32–53|url=http://direct.biostor.org/reference/111646}}</ref><ref name="brink1959">{{cite journal |last1=Brink|first1=A.S.|year=1959|title=''Struthiocephalus kitchingi'' sp. nov.|journal=Palaeontologia Africana|volume=5|issue=|pages=39–56|url=http://wiredspace.wits.ac.za/jspui/bitstream/10539/16102/1/1958.v5.BRINK.STRUTHIOCEPHALUS%20KITCHINGI.pdf}}</ref><ref name="boonstra1963"/><ref name="boonstra1965">{{cite journal |last1=Boonstra|first1=L.D.|year=1965|title=The skull of ''Struthiocephalus kitchingi''|journal=Annals of the South African Museum|volume=48|issue=14|pages=251–265|url=http://www.biodiversitylibrary.org/page/40731535#page/3/mode/1up}}</ref> ''[[Struthionops]]'',<ref name="Day&Rubidge2020"/> and ''[[Tapinocephalus]]'',<ref name="broom1928"/><ref name="boonstra1956">{{cite journal |last1=Boonstra|first1=L.D.|year=1956|title=The skull of ''Tapinocephalus'' and its near relatives|journal=Annals of the South African Museum|volume=42|issue=|pages=137–169|url=http://www.biodiversitylibrary.org/page/40864159#page/171/mode/1up}}</ref><ref name="boonstra1963"/><ref name="Neumann2020"/> the Styracocephalid ''[[Styracocephalus]]'',<ref name="haughton1929">{{cite journal |last1=Haughton|first1=S.H.|year=1929|title=On some new therapsid genera|journal=Annals of the South African Museum|volume=28|issue=|pages=55–78|url=http://www.biodiversitylibrary.org/item/126862#page/85/mode/1up}}</ref><ref name="rubidge1997">{{cite journal |last1=Rubidge|first1=B.S.|author2=van den Heever, J.A.|year=1997|title=Morphology and systematic position of the dinocephalian ''Styracocephalus platyrhynchus''|journal=Lethaia|volume=30|issue=2|pages=157–168|doi=10.1111/j.1502-3931.1997.tb00457.x|bibcode=1997Letha..30..157R }}</ref><ref name="Fraser-King&al.2019">{{cite journal |last1=Fraser-King|first1=S.W.|author2=Benoit, J.|author3=Day, M.O.|author4=Rubidge, B.S.|year=2019|title=Cranial morphology and phylogenetic relationship of the enigmatic dinocephalian ''Styracocephalus platyrhynchus'' from the Karoo Supergroup, South Africa|journal=Palaeontologia Africana|volume=54|issue=|pages=14–29|hdl=10539/28128 |url=https://hdl.handle.net/10539/28128}}</ref> and the huge [[Titanosuchidae|titanosuchids]] ''[[Jonkeria]]'' and ''[[Titanosuchus]]''.<ref name="broom1929">{{cite journal |last1=Broom|first1=R.|year=1929|title=On the carnivorous mammal-like reptiles of the family Titanosuchidae|journal=Annals of the Transvaal Museum|volume=13|issue=|pages=9–36}}</ref><ref name="boonstra1953a"/><ref name="smith2012"/><ref name="Day&Rubidge2020"/> Other large herbivores that were not synapsids included the large [[bradysauria]]n [[pareiasaur]]s represented by ''[[Bradysaurus]]'', ''[[Embrithosaurus]]'' and ''[[Nochelesaurus]]'', whose dentition very different from that of herbivorous dinocephalians indicates that the two groups occupied clearly distinct [[ecological niche]]s.<ref name="lee1997">{{cite journal |last1=Lee|first1=M.S.Y|year=1997|title=A taxonomic revision of pareiasaurian reptiles: implications for Permian terrestrial palaeoecology|journal=Modern Geology|volume=21|issue=1|pages=231–298|doi=10.1111/j.1096-3642.1997.tb01279.x|s2cid=84459755 }}</ref><ref name="VandenBrandt&al2020">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Abdala, F.|author3=Rubidge, B.S.|year=2020|title=Cranial morphology and phylogenetic relationships of the Middle Permian pareiasaur ''Embrithosaurus schwarzi'' from the Karoo Basin of South Africa|journal=Zoological Journal of the Linnean Society|volume=188|issue=|pages=202–241|doi=10.1093/zoolinnean/zlz064}}</ref><ref name="VandenBrandt&al2021a">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Rubidge, B.S.|author3=Benoit, J.|author4=Abdala, F.|year=2021|title=Cranial morphology of the middle Permian pareiasaur ''Nochelesaurus alexandri'' from the Karoo Basin of South Africa|journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh|volume=112|issue=1|pages=29–49|doi=10.1017/S1755691021000049|bibcode=2021EESTR.112...29V |s2cid=233839915 |doi-access=free}}</ref><ref name="VandenBrandt&al2021b">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Benoit, J.|author3=Abdala, F.|author4=Rubidge, B.S.|year=2021|title=Postcranial morphology of the South African middle Permian pareiasaurs from the Karoo Basin of South Africa|journal=Palaeontologia Africana|volume=55|issue=|pages=1–91|url=https://hdl.handle.net/10539/3190}}</ref><ref name="VandenBrandt&al2022">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Abdala, F.|author3=Benoit, J.|author4=Day, M.O.|author5=Groenewald, D.P.|author6=Rubidge, B.S.|year=2022|title=Taxonomy, phylogeny and stratigraphical ranges of middle Permian pareiasaurs from the Karoo Basin of South Africa|journal=Journal of Systematic Palaeontology|volume=19|issue=19|pages=1367–1393|doi=10.1080/14772019.2022.2035440|s2cid=247889546 |url=https://ddd.uab.cat/record/259563 }}</ref> The small to medium-sized forms included basal anomodonts (the non-dicynodonts ''[[Anomocephalus]]'',<ref name="modesto1999">{{cite journal|last1=Modesto|first1=S.P.|author2=Rubidge, B.S.|author3=Welman, J.|title=The most basal anomodont therapsid and the primacy of Gondwana in the evolution of the anomodonts|journal=Proceedings of the Royal Society B: Biological Sciences|date=1999|volume=266|issue=1417|pages=331–337|doi=10.1098/rspb.1999.0642|pmc=1689688 }}</ref><ref name="cisneros2015">{{cite journal|last1=Cisneros|first1=Juan Carlos|author2=Abdala, Fernando|author3=Jashashvili, Tea|author4=de Oliveira Bueno, Ana|author5=Dentzien-Dias, Paula|title=''Tiarajudens eccentricus'' and ''Anomocephalus africanus'', two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana|journal=Royal Society Open Science|date=2015|volume=2|issue=150090|page=150090 |doi=10.1098/rsos.150090|pmid=26587266 |pmc=4632579 |bibcode=2015RSOS....250090C |s2cid=25503025 |url=http://rsos.royalsocietypublishing.org/content/2/7/150090}}</ref> ''[[Galechirus]]'', ''[[Galeops]]'' and ''[[Galepus]]''<ref name="brinkman1981">{{cite journal |last1=Brinkman|first1=D.|year=1981|title=The structure and relationships of the dromasaurs (Reptila: Therapsida)|journal=Breviora|volume=465|issue=|pages=1–34|url=http://www.biodiversitylibrary.org/page/4287724#page/37/mode/1up}}</ref><ref name="smith2012"/><ref name="Day&Rubidge2020"/>) and numerous dicynodonts (''[[Brachyprosopus]]'',<ref name="angielczyk2016">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|author3=Day, M.O.|author4=Lin, F.|title=A reevaluation of ''Brachyprosopus broomi'' and ''Chelydontops altidentalis'', dicynodonts (Therapsida, Anomodontia) from the middle Permian ''Tapinocephalus'' Assemblage Zone of the Karoo Basin, South Africa|journal=Journal of Vertebrate Paleontology|date=2016|volume=36|issue=2|pages=e1078342 |doi=10.1080/02724634.2016.1078342|bibcode=2016JVPal..36E8342A |s2cid=130520407 }}</ref> ''[[Colobodectes]]'',<ref name="modesto2003a">{{cite journal|last1=Modesto|first1=S.|author2=Rubidge, B.|author3=Visser, I.|author4=Welman, J.|title=A new basal dicynodont from the Upper Permian of South Africa|journal=Palaeontology|date=2003|volume=46|issue=Part 1|pages=211–223|doi=10.1111/1475-4983.00295|bibcode=2003Palgy..46..211M |s2cid=128620351 |doi-access=free}}</ref><ref name="angielczyk2009">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|title=The Permian Dicynodont ''Colobodectes cluveri'' (Therapsida, Anomodontia), with notes on Its Ontogeny and Stratigraphic Range in the Karoo Basin, South Africa|journal=Journal of Vertebrate Paleontology|date=2009|volume=29|issue=4|pages=1162–1173|doi=10.1671/039.029.0431|bibcode=2009JVPal..29.1162A |s2cid=128771787 }}</ref> ''[[Pristerodon]]'',<ref name="smith2012"/> and the [[Pylaecephalidae|Pylaecephalids]] ''[[Diictodon]]'',<ref name="king1993">{{cite journal|last1=King|first1=G.M.|title=How many species of ''Diictodon'' were there ?|journal=Annals of the South African Museum|date=1993|volume=102|issue=|pages=303–325|url=http://biostor.org/reference/110293}}</ref><ref name="smith2012"/> ''[[Eosimops]]'',<ref name="angielczyk2012">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|title=Skeletal morphology, phylogenetic relationships and stratigraphic range of ''Eosimops newtoni'' Broom, 1921, a pylaecephalid dicynodont (Therapsida, Anomodontia) from the Middle Permian of South Africa|journal=Journal of Systematic Palaeontology|date=2012|volume=11|issue=2|pages=191–231|doi=10.1080/14772019.2011.623723|s2cid=129393393 }}</ref> ''[[Prosictodon]]'',<ref name="angielczyk2010">{{Cite journal|author=Kenneth D. Angielczyk; Bruce S. Rubidge|year=2010|title=A new pylaecephalid dicynodont (Therapsida, Anomodontia) from the Tapinocephalus Assemblage Zone, Karoo Basin, Middle Permian of South Africa|journal=Journal of Vertebrate Paleontology|volume=30|issue=5|pages=1396–1409|doi=10.1080/02724634.2010.501447|bibcode=2010JVPal..30.1396A |s2cid=129846697 }}</ref> and ''[[Robertia]]'',<ref name="smith2012"/>).
+Herbivorous were also numerous and diversified. Large-sized vegetarians were mainly represented by numerous dinocephalians including the tapinocephalids ''[[Agnosaurus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020">{{Cite thesis |degree=Ph.D. |title= Taxonomic Revision of the Short-Snouted Tapinocephalid Dinocephalia (Amniota-Therapsida) The Key to Understanding Middle Permian Tetrapod Biodiversity |url= https://wiredspace.wits.ac.za/handle/10539/31602 |last=Neumann |first=S. |year=2020 |publisher= University of the Witwatersrand |location=Johannesburg |pages=1–411}}</ref> ''[[Criocephalosaurus]]'',<ref name="broom1928">{{cite journal |last1=Broom|first1=R.|year=1928|title=On ''Tapinocephalus'' and two other dinocephalians|journal=Annals of the South African Museum|volume=22|issue=|pages=427–438}}</ref><ref name="kammerer2002">{{cite journal |last1=Kammerer|first1=C.|author2=Sidor, C.A.|year=2002|title=Replacement names for the therapsid genera ''Criocephalus'' Boom 1928 and ''Oliviera'' Brink 1965|journal=Palaeontologia Africana|volume=38|issue=|pages=71–72|url=http://wiredspace.wits.ac.za/handle/10539/16356}}</ref><ref name="day2015b"/><ref name="Neumann2020"/> ''[[Mormosaurus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020"/> ''[[Moschognathus]]'',<ref name="Day&Rubidge2020"/><ref name="Neumann2020"/> ''[[Moschops]]'',<ref name="gregory1926">{{cite journal|last1=Gregory|first1=W.K.|year=1926|title=The skeleton of ''Moschops capensis'' Broom, a Dinocephalian Reptile from the Permian of South Africa|journal=Bulletin of the American Museum of Natural History|volume=56|issue=3|pages=179–251|hdl=2246/1323 |url=http://digitallibrary.amnh.org/handle/2246/1323}}</ref><ref name="boonstra1957">{{cite journal|last1=Boonstra|first1=L.D.|year=1957|title=The moschopid skulls in the South African Museum|journal=Annals of the South African Museum|volume=44|issue=|pages=15–38|url=http://www.biodiversitylibrary.org/page/40862867#page/33/mode/1up}}</ref><ref name="boonstra1963">{{cite journal |last1=Boonstra|first1=L.D.|year=1963|title=Diversity within the South African Dinocephalia|journal=South African Journal of Science|volume=59|issue=|pages=196–207}}</ref><ref name="Neumann2020"/> ''[[Riebeeckosaurus]]'',<ref name="boonstra1957"/><ref name="güven2013">{{cite journal |last1=Güven|first1=S.|author2=Rubidge, B.S.|author3=Abdala, F.|year=2013|title=Cranial morphology and taxonomy of South African Tapinocephalidae (Therapsida, Dinocephalia): the case of ''Avenantia'' and ''Riebeeckosaurus''|journal=Palaeontologia Africana|volume=48|issue=|pages=24–33|url=http://wiredspace.wits.ac.za/handle/10539/13310}}</ref> ''[[Struthiocephalus]]''<ref name="boonstra1953b">{{cite journal|last1=Boonstra|first1=L.D.|year=1953|title=The cranial morphology and taxonomy of the tapinocephalid genus ''Struthiocephalus''|journal=Annals of the South African Museum|volume=42|issue=|pages=32–53|url=http://direct.biostor.org/reference/111646|access-date=2022-10-31|archive-date=2022-10-31|archive-url=https://web.archive.org/web/20221031190533/http://direct.biostor.org/reference/111646|url-status=dead}}</ref><ref name="brink1959">{{cite journal|last1=Brink|first1=A.S.|year=1959|title=''Struthiocephalus kitchingi'' sp. nov.|journal=Palaeontologia Africana|volume=5|issue=|pages=39–56|url=https://wiredspace.wits.ac.za/items/54fda047-347a-420c-bcb5-550f52582c40 |hdl=10539/16102 }}</ref><ref name="boonstra1963"/><ref name="boonstra1965">{{cite journal |last1=Boonstra|first1=L.D.|year=1965|title=The skull of ''Struthiocephalus kitchingi''|journal=Annals of the South African Museum|volume=48|issue=14|pages=251–265|url=http://www.biodiversitylibrary.org/page/40731535#page/3/mode/1up}}</ref> ''[[Struthionops]]'',<ref name="Day&Rubidge2020"/> and ''[[Tapinocephalus]]'',<ref name="broom1928"/><ref name="boonstra1956">{{cite journal |last1=Boonstra|first1=L.D.|year=1956|title=The skull of ''Tapinocephalus'' and its near relatives|journal=Annals of the South African Museum|volume=42|issue=|pages=137–169|url=http://www.biodiversitylibrary.org/page/40864159#page/171/mode/1up}}</ref><ref name="boonstra1963"/><ref name="Neumann2020"/> the Styracocephalid ''[[Styracocephalus]]'',<ref name="haughton1929">{{cite journal |last1=Haughton|first1=S.H.|year=1929|title=On some new therapsid genera|journal=Annals of the South African Museum|volume=28|issue=|pages=55–78|url=http://www.biodiversitylibrary.org/item/126862#page/85/mode/1up}}</ref><ref name="rubidge1997">{{cite journal |last1=Rubidge|first1=B.S.|author2=van den Heever, J.A.|year=1997|title=Morphology and systematic position of the dinocephalian ''Styracocephalus platyrhynchus''|journal=Lethaia|volume=30|issue=2|pages=157–168|doi=10.1111/j.1502-3931.1997.tb00457.x|bibcode=1997Letha..30..157R }}</ref><ref name="Fraser-King&al.2019">{{cite journal |last1=Fraser-King|first1=S.W.|author2=Benoit, J.|author3=Day, M.O.|author4=Rubidge, B.S.|year=2019|title=Cranial morphology and phylogenetic relationship of the enigmatic dinocephalian ''Styracocephalus platyrhynchus'' from the Karoo Supergroup, South Africa|journal=Palaeontologia Africana|volume=54|issue=|pages=14–29|hdl=10539/28128 |url=https://hdl.handle.net/10539/28128}}</ref> and the huge [[Titanosuchidae|titanosuchids]] ''[[Jonkeria]]'' and ''[[Titanosuchus]]''.<ref name="broom1929">{{cite journal |last1=Broom|first1=R.|year=1929|title=On the carnivorous mammal-like reptiles of the family Titanosuchidae|journal=Annals of the Transvaal Museum|volume=13|issue=|pages=9–36}}</ref><ref name="boonstra1953a"/><ref name="smith2012"/><ref name="Day&Rubidge2020"/> Other large herbivores that were not synapsids included the large [[bradysauria]]n [[pareiasaur]]s represented by ''[[Bradysaurus]]'', ''[[Embrithosaurus]]'' and ''[[Nochelesaurus]]'', whose dentition very different from that of herbivorous dinocephalians indicates that the two groups occupied clearly distinct [[ecological niche]]s.<ref name="lee1997">{{cite journal |last1=Lee|first1=M.S.Y|year=1997|title=A taxonomic revision of pareiasaurian reptiles: implications for Permian terrestrial palaeoecology|journal=Modern Geology|volume=21|issue=1|pages=231–298|doi=10.1111/j.1096-3642.1997.tb01279.x|s2cid=84459755 }}</ref><ref name="VandenBrandt&al2020">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Abdala, F.|author3=Rubidge, B.S.|year=2020|title=Cranial morphology and phylogenetic relationships of the Middle Permian pareiasaur ''Embrithosaurus schwarzi'' from the Karoo Basin of South Africa|journal=Zoological Journal of the Linnean Society|volume=188|issue=|pages=202–241|doi=10.1093/zoolinnean/zlz064}}</ref><ref name="VandenBrandt&al2021a">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Rubidge, B.S.|author3=Benoit, J.|author4=Abdala, F.|year=2021|title=Cranial morphology of the middle Permian pareiasaur ''Nochelesaurus alexandri'' from the Karoo Basin of South Africa|journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh|volume=112|issue=1|pages=29–49|doi=10.1017/S1755691021000049|bibcode=2021EESTR.112...29V |s2cid=233839915 |doi-access=free}}</ref><ref name="VandenBrandt&al2021b">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Benoit, J.|author3=Abdala, F.|author4=Rubidge, B.S.|year=2021|title=Postcranial morphology of the South African middle Permian pareiasaurs from the Karoo Basin of South Africa|journal=Palaeontologia Africana|volume=55|issue=|pages=1–91|url=https://hdl.handle.net/10539/3190}}</ref><ref name="VandenBrandt&al2022">{{cite journal |last1=Van den Brandt|first1=M.J.|author2=Abdala, F.|author3=Benoit, J.|author4=Day, M.O.|author5=Groenewald, D.P.|author6=Rubidge, B.S.|year=2022|title=Taxonomy, phylogeny and stratigraphical ranges of middle Permian pareiasaurs from the Karoo Basin of South Africa|journal=Journal of Systematic Palaeontology|volume=19|issue=19|pages=1367–1393|doi=10.1080/14772019.2022.2035440|s2cid=247889546 |url=https://ddd.uab.cat/record/259563 }}</ref> The small to medium-sized forms included basal anomodonts (the non-dicynodonts ''[[Anomocephalus]]'',<ref name="modesto1999">{{cite journal|last1=Modesto|first1=S.P.|author2=Rubidge, B.S.|author3=Welman, J.|title=The most basal anomodont therapsid and the primacy of Gondwana in the evolution of the anomodonts|journal=Proceedings of the Royal Society B: Biological Sciences|date=1999|volume=266|issue=1417|pages=331–337|doi=10.1098/rspb.1999.0642|pmc=1689688 }}</ref><ref name="cisneros2015">{{cite journal|last1=Cisneros|first1=Juan Carlos|author2=Abdala, Fernando|author3=Jashashvili, Tea|author4=de Oliveira Bueno, Ana|author5=Dentzien-Dias, Paula|title=''Tiarajudens eccentricus'' and ''Anomocephalus africanus'', two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana|journal=Royal Society Open Science|date=2015|volume=2|issue=150090|page=150090 |doi=10.1098/rsos.150090|pmid=26587266 |pmc=4632579 |bibcode=2015RSOS....250090C |s2cid=25503025 |url=http://rsos.royalsocietypublishing.org/content/2/7/150090}}</ref> ''[[Galechirus]]'', ''[[Galeops]]'' and ''[[Galepus]]''<ref name="brinkman1981">{{cite journal |last1=Brinkman|first1=D.|year=1981|title=The structure and relationships of the dromasaurs (Reptila: Therapsida)|journal=Breviora|volume=465|issue=|pages=1–34|url=http://www.biodiversitylibrary.org/page/4287724#page/37/mode/1up}}</ref><ref name="smith2012"/><ref name="Day&Rubidge2020"/>) and numerous dicynodonts (''[[Brachyprosopus]]'',<ref name="angielczyk2016">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|author3=Day, M.O.|author4=Lin, F.|title=A reevaluation of ''Brachyprosopus broomi'' and ''Chelydontops altidentalis'', dicynodonts (Therapsida, Anomodontia) from the middle Permian ''Tapinocephalus'' Assemblage Zone of the Karoo Basin, South Africa|journal=Journal of Vertebrate Paleontology|date=2016|volume=36|issue=2|pages=e1078342 |doi=10.1080/02724634.2016.1078342|bibcode=2016JVPal..36E8342A |s2cid=130520407 }}</ref> ''[[Colobodectes]]'',<ref name="modesto2003a">{{cite journal|last1=Modesto|first1=S.|author2=Rubidge, B.|author3=Visser, I.|author4=Welman, J.|title=A new basal dicynodont from the Upper Permian of South Africa|journal=Palaeontology|date=2003|volume=46|issue=Part 1|pages=211–223|doi=10.1111/1475-4983.00295|bibcode=2003Palgy..46..211M |s2cid=128620351 |doi-access=free}}</ref><ref name="angielczyk2009">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|title=The Permian Dicynodont ''Colobodectes cluveri'' (Therapsida, Anomodontia), with notes on Its Ontogeny and Stratigraphic Range in the Karoo Basin, South Africa|journal=Journal of Vertebrate Paleontology|date=2009|volume=29|issue=4|pages=1162–1173|doi=10.1671/039.029.0431|bibcode=2009JVPal..29.1162A |s2cid=128771787 }}</ref> ''[[Pristerodon]]'',<ref name="smith2012"/> and the [[Pylaecephalidae|Pylaecephalids]] ''[[Diictodon]]'',<ref name="king1993">{{cite journal|last1=King|first1=G.M.|title=How many species of ''Diictodon'' were there ?|journal=Annals of the South African Museum|date=1993|volume=102|issue=|pages=303–325|url=http://biostor.org/reference/110293}}</ref><ref name="smith2012"/> ''[[Eosimops]]'',<ref name="angielczyk2012">{{cite journal|last1=Angielczyk|first1=K.D.|author2=Rubidge, B.S.|title=Skeletal morphology, phylogenetic relationships and stratigraphic range of ''Eosimops newtoni'' Broom, 1921, a pylaecephalid dicynodont (Therapsida, Anomodontia) from the Middle Permian of South Africa|journal=Journal of Systematic Palaeontology|date=2012|volume=11|issue=2|pages=191–231|doi=10.1080/14772019.2011.623723|s2cid=129393393 }}</ref> ''[[Prosictodon]]'',<ref name="angielczyk2010">{{Cite journal |first1=Kenneth D. |last1=Angielczyk |author2=Bruce S. Rubidge|year=2010|title=A new pylaecephalid dicynodont (Therapsida, Anomodontia) from the ''Tapinocephalus'' Assemblage Zone, Karoo Basin, Middle Permian of South Africa|journal=Journal of Vertebrate Paleontology|volume=30|issue=5|pages=1396–1409|doi=10.1080/02724634.2010.501447|bibcode=2010JVPal..30.1396A |s2cid=129846697 }}</ref> and ''[[Robertia]]'',<ref name="smith2012"/>).
 == Classification and phylogeny ==
-Named by Watson in 1921, ''Anteosaurus'' was longtime classified as a ‘Titanosuchian Deinocephalian’, and it is only in 1954 that Boonstra separated the Titanosuchians in two families : Jonkeridae (a [[Synonym (taxonomy)#Zoology|junior synonym]] of Titanosuchidae) and Anteosauridae.<ref name="watson1921">{{cite journal|last=Watson|first=D.M.S.|title=The Bases of Classification of the Theriodontia|journal=Proceedings of the Zoological Society of London|year=1921|volume=91|issue=|pages=35–98|doi=10.1111/j.1096-3642.1921.tb03250.x |url=http://www.biodiversitylibrary.org/item/98474#page/124/mode/1up}}</ref><ref name="boonstra1954a"/> At about the same time, Efremov erected the family [[Brithopodidae]] in which he includes the fragmentary ''[[Brithopus]]'' and the better known forms ''Syodon'' and ''Titanophoneus''.<ref name="efremov1954">{{cite journal|last=Efremov|first=I.A.|title=[The terrestrial vertebrate fauna from the Permian copper sandstones of the western Fore-Urals]|journal=Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSSR|year=1954|volume=54|issue=|pages=1-416 [in Russian]}}</ref> Much later, Hopson and Barghusen argued that Brithopodidae should be discontinued and that the Russian taxa ''Syodon'', ''Titanophoneus'' and ''[[Doliosauriscus]]'' should be placed with ''Anteosaurus'' in [[Anteosauridae]]. These authors placed also Anteosauridae in the new group Anteosauria for distinguished them of the other major dinocephalian group the [[Tapinocephalia]] in which they included the titanosuchids and the tapinocephalids. They also created the taxa Anteosaurinae, containing ''Anteosaurus'' and the Russian forms ''Titanophoneus'' and ''Doliosauriscus'', and the Anteosaurini containing only the giant forms ''Anteosaurus'' and ''Doliosauriscus''.<ref name="hopson1986">{{cite book|last=Hopson|first=J.A.|author2=Barghusen, H.R.|year=1986|chapter=An analysis of therapsid relationships|pages=83–106|title=The Ecology and Biology of the Mammal-Like Reptiles|editor1-last=Hotton III|editor1-first=N.|editor2-last=MacLean|editor2-first=P.D.|editor3-last=Roth|editor3-first=J.J.|editor4-last=Roth|editor4-first=E.C.|publisher=Smithsonian Institution Press|location=Washington, D.C.|isbn=978-0874745245}}</ref> Gilian King retained the incorrectly spelled ‘Brithopidae’ (including the subfamilies ‘Brithopinae’ and Anteosaurinae) and placed both Brithopidae and Titanosuchidae (including Titanosuchinae and Tapinocephalinae) in the superfamily Anteosauroidea.<ref name="king1988">{{cite book|last=King|first=G.|year=1988|title=Anomodontia (Handbuch der Paläoherpetologie, 17C)|url=|location=Stuttgart|publisher=Gustav Fischer|pages=1–174|isbn=978-0895742506}}</ref> Later Ivakhnenko considered Brithopodidae as invalid and united Anteosauridae and Deuterosauridae (only known by the Russian ''[[Deuterosaurus]]'') in the superfamily Deuterosauroidea.<ref name="ivakhnenko1995">{{cite journal|last=Ivakhnenko|first=M.F.|title=Primitive Late Permian dinocephalian-titanosuchids of Eastern Europe|journal=Paleontological Journal|year=1995|volume=29|issue=|pages=120–129}}</ref><ref name="ivakhnenko2003">{{cite journal|last=Ivakhnenko|first=M.F.|title=Eotherapsids from the East European Placket (Late Permian)|journal=Paleontological Journal|year=2003|volume=37|issue=|pages=339–465}}</ref> More recently Kammerer in its systematic revision of the anteosaurs (in which ''Doliosauriscus'' become a junior synonym of ''Titanophoneus'') demonstrated that the [[wastebasket taxon|wastebasket]] genus ''Brithopus'' is a ''[[nomen dubium]]'' composed both of remains of indeterminate [[Estemmenosuchidae|estemmenosuchid]]-like tapinocephalian and indeterminate anteosaurian, so invalidating the Brithopodidae. He proposed also the first [[Phylogenetics|phylogenetic analysis]] including all anteosaurid taxa. This and other modern phylogenetic analysis of anteosaurs recovers a [[Monophyly|monophyletic]] Anteosauridae containing two major [[clade]]s, [[Syodontinae]] and [[Anteosaurinae]]. In the Kammerer analysis, the Chinese ''[[Sinophoneus]]'' is the most basal anteosaurine and the sister-group of an unresolved trichotomy including ''Titanophoneus potens'', ''T. adamanteus'' and ''Anteosaurus''.<ref name="kammerer2011"/>
+Named by Watson in 1921, ''Anteosaurus'' was longtime classified as a 'Titanosuchian Deinocephalian', and it is only in 1954 that Boonstra separated the Titanosuchians in two families: Jonkeridae (a [[Synonym (taxonomy)#Zoology|junior synonym]] of Titanosuchidae) and Anteosauridae.<ref name="watson1921">{{cite journal|last=Watson|first=D.M.S.|title=The Bases of Classification of the Theriodontia|journal=Proceedings of the Zoological Society of London|year=1921|volume=91|issue=|pages=35–98|doi=10.1111/j.1096-3642.1921.tb03250.x |url=http://www.biodiversitylibrary.org/item/98474#page/124/mode/1up}}</ref><ref name="boonstra1954a"/> At about the same time, Efremov erected the family [[Brithopodidae]] in which he includes the fragmentary ''[[Brithopus]]'' and the better known forms ''Syodon'' and ''Titanophoneus''.<ref name="efremov1954">{{cite journal|last=Efremov|first=I.A.|title=[The terrestrial vertebrate fauna from the Permian copper sandstones of the western Fore-Urals]|journal=Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSSR|year=1954|volume=54|issue=|pages=1-416 [in Russian]}}</ref> Much later, Hopson and Barghusen argued that Brithopodidae should be discontinued and that the Russian taxa ''Syodon'', ''Titanophoneus'' and ''[[Doliosauriscus]]'' should be placed with ''Anteosaurus'' in [[Anteosauridae]]. These authors placed also Anteosauridae in the new group Anteosauria for distinguished them of the other major dinocephalian group the [[Tapinocephalia]] in which they included the titanosuchids and the tapinocephalids. They also created the taxa Anteosaurinae, containing ''Anteosaurus'' and the Russian forms ''Titanophoneus'' and ''Doliosauriscus'', and the Anteosaurini containing only the giant forms ''Anteosaurus'' and ''Doliosauriscus''.<ref name="hopson1986">{{cite book|last=Hopson|first=J.A.|author2=Barghusen, H.R.|year=1986|chapter=An analysis of therapsid relationships|pages=83–106|title=The Ecology and Biology of the Mammal-Like Reptiles|editor1-last=Hotton III|editor1-first=N.|editor2-last=MacLean|editor2-first=P.D.|editor3-last=Roth|editor3-first=J.J.|editor4-last=Roth|editor4-first=E.C.|publisher=Smithsonian Institution Press|location=Washington, D.C.|isbn=978-0874745245}}</ref> Gilian King retained the incorrectly spelled 'Brithopidae' (including the subfamilies 'Brithopinae' and Anteosaurinae) and placed both Brithopidae and Titanosuchidae (including Titanosuchinae and Tapinocephalinae) in the superfamily Anteosauroidea.<ref name="king1988">{{cite book|last=King|first=G.|year=1988|title=Anomodontia (Handbuch der Paläoherpetologie, 17C)|url=|location=Stuttgart|publisher=Gustav Fischer|pages=1–174|isbn=978-0895742506}}</ref> Later Ivakhnenko considered Brithopodidae as invalid and united Anteosauridae and Deuterosauridae (only known by the Russian ''[[Deuterosaurus]]'') in the superfamily Deuterosauroidea.<ref name="ivakhnenko1995">{{cite journal|last=Ivakhnenko|first=M.F.|title=Primitive Late Permian dinocephalian-titanosuchids of Eastern Europe|journal=Paleontological Journal|year=1995|volume=29|issue=|pages=120–129}}</ref><ref name="ivakhnenko2003">{{cite journal|last=Ivakhnenko|first=M.F.|title=Eotherapsids from the East European Placket (Late Permian)|journal=Paleontological Journal|year=2003|volume=37|issue=|pages=339–465}}</ref> More recently Kammerer in its systematic revision of the anteosaurs (in which ''Doliosauriscus'' become a junior synonym of ''Titanophoneus'') demonstrated that the [[wastebasket taxon|wastebasket]] genus ''Brithopus'' is a ''[[nomen dubium]]'' composed both of remains of indeterminate [[Estemmenosuchidae|estemmenosuchid]]-like tapinocephalian and indeterminate anteosaurian, so invalidating the Brithopodidae. He proposed also the first [[Phylogenetics|phylogenetic analysis]] including all anteosaurid taxa. This and other modern phylogenetic analysis of anteosaurs recovers a [[Monophyly|monophyletic]] Anteosauridae containing two major [[clade]]s, [[Syodontinae]] and [[Anteosaurinae]]. In the Kammerer analysis, the Chinese ''[[Sinophoneus]]'' is the most basal anteosaurine and the sister-group of an unresolved trichotomy including ''Titanophoneus potens'', ''T. adamanteus'' and ''Anteosaurus''.<ref name="kammerer2011"/>
 Below the cladogramm of Kammerer published in 2011 :
@@ Line 283: / Line 283: @@
 ===Genus synonymy===
-As defined by [[Lieuwe Dirk Boonstra]], ''Anteosaurus'' is “a genus of anteosaurids in which the postfrontal forms a boss of variable size overhanging the dorso-posterior border of the orbit.” On this basis he synonymised six of the seven genera named from the [[Tapinocephalus Assemblage Zone|''Tapinocephalus'' zone]]: ''Eccasaurus'', ''Anteosaurus'', ''Titanognathus'', ''Dinosuchus'', ''Micranteosaurus'', and ''Pseudanteosaurus''. Of these, he says, ''Dinosuchus'' and ''Titanognathus'' can safely be considered synonyms of ''Anteosaurus''. ''Eccasaurus'', with a holotype of which the cranial material consists of only few typical anteosaurid incisors, appears to be only determinable as to family. The skull fragment forming the holotype of ''Pseudanteosaurus'' can best be considered as an immature specimen of ''Anteosaurus''. ''Micranteosaurus'', the holotype of which contains a small snout, was previously considered a new genus on account of its small size but is better be interpreted as a young specimen of ''Anteosaurus''. And likewise, the large number of species attributed to the genus ''Anteosaurus'' can also be considered synonyms. Boonstra still considers as valid the genus ''Paranteosaurus'', which is defined as a genus of anteosaurids in which the postfrontal is not developed to form a boss. This is probably an example of individual variation and hence another synonym of ''Anteosaurus''.<ref name=Boonstra1969>{{cite journal|last1=Boonstra|first1=L. D.|date=1969|title=The Fauna of the Tapinocephalus Zone (Beaufort Beds of the Karoo)|journal=Annals of the South African Museum|volume=56|number=1|pages=1−73|url=https://www.biodiversitylibrary.org/page/40722876#page/3/mode/1up}}</ref>
+As defined by [[Lieuwe Dirk Boonstra]], ''Anteosaurus'' is "a genus of anteosaurids in which the postfrontal forms a boss of variable size overhanging the dorso-posterior border of the orbit." On this basis he synonymised six of the seven genera named from the [[Tapinocephalus Assemblage Zone|''Tapinocephalus'' zone]]: ''Eccasaurus'', ''Anteosaurus'', ''Titanognathus'', ''Dinosuchus'', ''Micranteosaurus'', and ''Pseudanteosaurus''. Of these, he says, ''Dinosuchus'' and ''Titanognathus'' can safely be considered synonyms of ''Anteosaurus''. ''Eccasaurus'', with a holotype of which the cranial material consists of only few typical anteosaurid incisors, appears to be only determinable as to family. The skull fragment forming the holotype of ''Pseudanteosaurus'' can best be considered as an immature specimen of ''Anteosaurus''. ''Micranteosaurus'', the holotype of which contains a small snout, was previously considered a new genus on account of its small size but is better be interpreted as a young specimen of ''Anteosaurus''. And likewise, the large number of species attributed to the genus ''Anteosaurus'' can also be considered synonyms. Boonstra still considers as valid the genus ''Paranteosaurus'', which is defined as a genus of anteosaurids in which the postfrontal is not developed to form a boss. This is probably an example of individual variation and hence another synonym of ''Anteosaurus''.<ref name=Boonstra1969>{{cite journal|last1=Boonstra|first1=L. D.|date=1969|title=The Fauna of the Tapinocephalus Zone (Beaufort Beds of the Karoo)|journal=Annals of the South African Museum|volume=56|number=1|pages=1−73|url=https://www.biodiversitylibrary.org/page/40722876#page/3/mode/1up}}</ref>
 ===Species synonymy===
-[[File:Annals of the South African Museum - Annale van die Suid-Afrikaanse Museum (1953) (18422948111).jpg|thumb|left|Skull of ''A. magnificus'' (SAM-PK-11577) in latero-dorsal view. This large skull (74 cm long) has a very weak frontal boss and very large temporal fenestrae. It was formerly assigned to the species ''A. vorsteri''. This specimen was discovered near Beaufort West in the Western Cape province.]][[File:Annals of the South African Museum = Annale van die Suid-Afrikaanse Museum (1953) (17798603714).jpg|thumb|right|Skull of ''A. magnificus'' (SAM-PK-11293), this specimen was the [[holotype]] of ''A. major''. It is one of the largest skull of ''A. magnificus'' with a length of over 80 cm. The skull is crushed dorsoventraly and the snout has not been preserved. It is here restored with plaster, so the exact size of the skull remains uncertain. This specimen comes from [[Beaufort West]] area in [[Western Cape]] province.<ref name="boonstra1953a"/><ref name="boonstra1954a"/><ref name="kammerer2011"/>]]
 ''Anteosaurus'' was once known by a large number of species, but the current thinking on this is that they merely represent different growth stages of the same type species, ''A. magnificus''.<ref name=Boonstra1969/><ref name="kammerer2011"/>
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 [[Category:Fossil taxa described in 1921]]
 [[Category:Taxa named by D. M. S. Watson]]
-[[Category:Guadalupian life]]
 [[Category:Capitanian life]]

Latest revision as of 09:38, 11 November 2024

Etymology

Description

Size

Skull

Dentition

Postcranial skeleton

Paleobiology

Skull variations and agonistic behaviour

Ontogeny

Habitat preference and diet

Geographic and stratigraphic range

South Africa

Russia?

Paleoenvironment

Paleogeography and paleoclimate

Paleoecology

Classification and phylogeny

Genus synonymy

Species synonymy

Possible synonyms

Archaeosuchus

Eccasaurus

See also

Notes

References

External links