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Jump to: ,
Parent taxa:
(check the following menu and phylogeny - the taxon in bold refers to the topic on this page)


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Anteosauria


The head of the small generalised anteosaur ((mail uk viagra,,,), from the Middle Permian of equatorial east-central Pangea. Illustration © [ Pedro Andrade


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Anteosaurs are a group of generally large, primitive (early "mammal-like reptiles"), with huge canines and incisors and short limbs, that are known from the Middle of South Africa, Russia, and China. The largest grew to very large size, with skulls 50 to 80 cm long, and were the largest predators of their time. They died out at the end of the Middle Permian, possibly as a result of the extinct of the herbivorous on which they fed.

The Anteosauria are destinguished from the Tapinocephalia by a number of features, such as very large , cheek teeth with bulbous , and an upturning of the , so that the front of mouth curves strongly upwards. The limbs are short and the skull long, narrow, and heavy. The temporal opening is much larger than in the early , indicating larger jaw muscles and a stronger bite. There is a tendency especially in more advanced forms such as towards thickening of the bones of the top of the skull, indicating head-butting behaviour. The tail is very long in at least some genera. Precise details of the of the group are still not well known.

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Taxonomy Very Speculative Phylogeny

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  • Family ?
  • Family (= ?)
    • Subfamily
    • Subfamily
    • Subfamily
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        |-- (Deuterosaurinae/idea)
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        |  |--
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:

  • Phylum:
    • Class: (traditional) / (Benton 2000, 2004)
      • Order:
        • Suborder:
          • Infraorder/Superfamily: Anteosauroidea

Author: , 1962

Stratigraphic range: to

Distribution:

Lifestyle: and and

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The family "Brithopodidae" was erected by Efremov in 1954 to include the carnivorous therapsids of the Russian Tatarian (Late Permian) Copper Sandstones and other Cis-Uralian localities, including both those of the Cis-Uralian Dinocephalian Complexes and the Isheevian Dinocephalian Complex. In the same year, Boonstra created the family Anteosauridae for the giant carnivorous dinocephalians of the South African Lower Beaufort (Tapinocephalus Zone).

The group was originally defined as a by in 1962 to include the and

In his 1962 monograph on American and Russian Late Permian reptiles, Olson (1962) places the distinct herbivorous "brithopodid" Deuterosaurus in a family of its own. For several decades these three families were considered distinct if closely related.

and in provided the first study of the Therapsida. They used the term "Anteosauria" and synonymised the families Brithopodidae and Anteosauridae. They sugggested the following schema:

             ,----------------------------------
             |
ANTEOSAURIA--|                 ,----------------
(=Anteosaur- |                 |
  idae)      `----|                
                               |                ,--
                               `----|
                                                `--

(see Hopson & Barghusen 1986 p.91)

in a 1988 monograph on the (including the Dinocephalia - however the view that the Dinocephalia are a subset of the Anomodontia is no longer held) as part of 's ongoing Encyclopedia of Paleoherpetology series of volumes, uses a more traditional Linnaean arrangement, but includes the herbivorous forms under the superfamily Anteosauroidea as well. She suggested a subfamily distinction, although rejecting the status of the Deuterosauridae:

Superfamily Anteosauroidea BOONSTRA 1962
    Family Brithopidae BOONSTRA 1972
      Subfamily Brithopodinae EFREMOV 1954
      Subfamily Anteosaurinae 1954
    Family Titanosuchidae BOONSTRA 1972
      Subfamily Titanosuchinae BROOM 1903
      Subfamily Tapinocephalinae LYDEKKER 1890

Note that the "Titanosuchidae" here is equivalent to the "".

The current cladistic-based tendency is of course to do away with these Linnean taxa altogether. Hence our non-committal title, Anteosauria.

Finally, as part of a much larger therapsid , and in 2001 offer the following:

  ,----------------- 
  |
--|    ,------------   
  |    |
  |    |------------ 
  `----|                
       |      ,----- 
       `------|
              `----- 

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The early Russian (Boonstra 1972) and Chinese (Rubidge & Sidor 2001) anteosaurs are generally considered the most primitive of the Dinocephalia, and hence of the (in which case they may constitute a assemblage), although it has also been suggested (Kemp, 1982, King 1988) that the are more . They have features in common with (Carroll 1988) and (Chudinov 1965), and, with the Tapinocephalia, are part of the first major evolutionary radiation of the Therapsida (Rubidge & Sidor 2001). So far little work has been done on detailed relationships between the various .

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The following can be considered partly or totally synonymous:

  • Superfamily: DEUTEROSAUROIDEA Seeley 1894,
  • Family: BRITHOPIDIDAE Efremov, 1954
  • Superfamily: ANTEOSAUROIDEA Boonstra 1954, Boonstra 1962
  • Order: EOTHERIODONTIA Olson 1962 (= Bairmosuchia + Anteosauria)
  • Family: BRITHOPIDAE Boonstra 1972, King 1988
  • Plesion: ANTEOSAURIA Hopson & Barghusen 1986

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The following section of this page needs to be . At its simplest, wikifying can just mean adding links to other pages via the edit tab. For help in editing Palaeos org pages, see the page

A full understanding of anteosaur phylogeny has to await greater scrutiny of the available remains. Much of the Russian material is fragmentary, and a lot of this will no doubt turn out to be synonymous with the better known forms. It was originally thought that the earliest anteosaur was the very fragmentary American Eosyodon Olson, of Roadian age. It now seems that all of the proto-therapsids Olson described in his monograph are chimeras - mostly misinterpretations of caseid pelycosaurs [C. Kammerer, pers. comm.]. That leaves the Russian Ocher fauna from Ezhovo (probably early Wordian age) as the oldest known anteosaurs. Two species are known from this period. Significantly, Archaeosyodon is known only from the skull, and Chthomaloporus only from postcrania. It is not unlikely that these two thus represent the same animal. The skull is typical of a fairly primitive anteosaur and indicates a reasonably large animal - perhaps a large predator intermediate in size between the dog-sized Biarmosuchus and the huge Eotitanosuchus, both of which were contemporary. The postcrania however include a pelvic girdle quite unique from that of other anteosaurs, indicating that this type represents a side-branch rather than a direct ancestor.

By the early Capitanian (late Kazanian - Bashkirian subzone), perhaps several million years later, the gigantic eotitanosuchid super-predators are gone, along with most of their estemmenosuchid herbivore prey (perhaps the victims of a minor mass-extinction) and only a few smallish anteosaurs remain. They seem to have diverged into at least two distinct lineages - the primitive carnivorous types (Syodon and Brithopus - Brithopididae) and the more specialized herbivores (Deuterosaurus). The latter perhaps filled the ecological role vacated by the estemmenosuchids.

A million or so years later, all the animals had grown in size, evolving into bigger and bigger types. A big species of Brithopus was joined by a giant Admetophoneus (subfamily Anteosaurinae), and the deuterosaurs had also increased in size, and were now existing alongside the big advanced tapinocephalid herbivores. These animals are by and large a lot bigger than those that existed during Bashkirian time.

By the time of the forms that have been found at Isheevo, the continuing basal primitive types like Syodon and Brithopus, and the advanced giant carnivorous forms like Titanophoneus and Doliosauriscus, were flourishing. The deuterosaurs were gone, perhaps replaced by the tapinocephalids. This seems to be the golden age of anteosaur diversity, and indeed of the dinocephalians in general. The South African Anteosaurus (Tapinocephalus zone), a representative of the titanophonine- doliosaur lineage (subfamily Anteosaurinae), seems to have existed at this time and also slightly later.

It is interesting that whereas in the South African temperate zone fauna there is only one anteosaurian predator (the giant Anteosaurus), in the tropical Isheevo fauna (yes, at the time Russia was located in the tropics!) there are four. Perhaps there were more ecological niches in the tropics for these animals. Perhaps some of the species will turn out to be synonyms or juveniles of other forms. Perhaps the smaller Russian anteosaurs like Syodon filled the same ecological niche as the similarly sized Titanosuchus did in the temperate regions. Much still remains to be known about these amazing animals.

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The following section of this page needs to be . At its simplest, wikifying can just mean adding links to other pages via the edit tab. For help in editing Palaeos org pages, see the page

mail uk viagra

The basic technical diagnosis of the Anteosauria (Brithopodidae), which includes the three subfamilies, is as follows: A group of primitive carnivorous dinocephalians. The skull is high and narrow. The palatal teeth - the teeth on the roof of the mouth, a feature of many Paleozoic tetrapods - are enlarged and usually confined to a kidney-shaped cluster on each palatine (near the outer tooth row). Any palatal teeth on the pterygoid flange are reduced. The marginal teeth (teeth in the "normal" position) are of three types: anterior (front or incisor), canine, and cheek teeth. Here we have the further differentiation of tooth types along the line to the mammalian condition. The incisors (the teeth at the front) each have a small ledge or "heel", a distinctive dinocephalian feature that only the estemmenosuchids lack. The canine is very large in all anteosaurs, even the herbivorous ones. There are usually about ten cheek teeth (although the number is very variable, depending on age and tooth replacement), possessing bulbous crowns. The teeth may decrease in size from front to rear. There is a "step" or upturning in the premaxilla bone of the upper jaw, so the front of mouth curves strongly upwards. A somewhat similar "kink" (possibly a mobile joint) can be seen among some archosaurs such as proterosuchids and coleophysid dinosaurs. The postorbital bone and bar behind the eye are large, modified to produce an area for the attachment of the superficial adductor jaw muscles on the outer surface of skull. There is a tendency towards pachyostosis, the thickening of the roof of the head that is a common dinocephalian characteristic, perhaps indicative of "head-butting" intra-specific behavior (rivalry for mates and/or territory). This takes the form of a tuberosity around the pineal opening, in advanced forms as a bony "boss" (almost like a low mound). There is a large canal for pineal organ ("third eye", probably tied in with the animal's diurnal and seasonal cycles). The shoulder girdle is fairly light, with a narrow interclavicle, clavicle, and scapular blade. The femur (thigh bone) is slender and curved. These were, in spite of their size, probably quite agile animals.

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Characters: skull long, high and narrow [BS00]; skull pachyostosis common, especially around orbital rims & parietals [BS00]; premaxilla with "step" or upturning, so the front of mouth curves strongly upwards; pineal foramen large, with pronounced tuberosity; temporal fenestra well open dorsally [BS00]; palatal teeth enlarged and usually confined to kidney-shaped cluster on palatines; postorbital bone and bar large, modified to area for external attachment of superficial adductors; any teeth on pterygoid flange reduced; marginal teeth include incisors, canines, and cheek teeth; incisors with small ledge or "heel" [BS00]; canines very large [BS00]; usually about ten cheek teeth; (rather variable, dependent on age and tooth replacement); cheek teeth with bulbous crowns [BS00]; teeth may decrease in size distally; shoulder girdle is fairly light, with a narrow interclavicle, clavicle, and scapular blade; femur slender and curved; mostly carnivorous.

References: Battail & Surkov (2000) [BS00]; Rubidge & Sidor (2001) [RS01].

Note: possibly paraphyletic [RS01]. ATW020219.

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  • Battail, B & MV Surkov (2000), Mammal-like reptiles from Russia in MJ Benton, MA Shishkin, DM Unwin & EN Kurochkin (eds.), The Age of Dinosaurs in Russia and Mongolia, Cambridge Univ. Press, pp 86-119.
  • , 1972, Discard the names Theriodontia and Anomodontia: a new classification of the Therapsida. 59:315-338.
  • , 1988, Vertebrate Paleontology and Evolution, WH Freeman & Co.
  • 1965, "New Facts about the Fauna of the Upper Permian of the USSR", , 73:117-30
  • and , 1986, An analysis of therapsid relationships in , , and , The Ecology and Biology of Mammal-like Reptiles, , pp. 83-106
  • , 1988, "Anomodontia" Part 17 C, Encyclopedia of Paleoherpetology, , Stuttgart and New York,
  • & 2001, Evolutionary patterns among Permo-Triassic therapsids. 32: 449-480.

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  • - includes brief but readable introduction of Anteosaurs and other Dinbocephalia
  • - list of genera in the form of a

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MAK000902 ATW020219 (characters) MAK060407

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