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Estemmenosuchidae is an extinct family of large, very early herbivorous therapsids that flourished during the Guadalupian period. They are distinguished by horn-like structures, probably for
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. Apart from the best known genus, ''
Estemmenosuchus ''Estemmenosuchus'' (meaning "crowned crocodile" in Greek) is an extinct genus of large, early omnivorous therapsid. It is believed and interpreted to have lived during the middle part of the Middle Permian around 267 million years ago. The two ...
'', the group is poorly known. To date, their fossils are known only from the Perm region of Russia (a region referred to by Russian paleontologists as the Cis-Urals).


Description

Estemmenosuchids are among the most distinctive of the Permian tetrapods. The high and massive skull is equipped with a number of horns projecting both upwards and outwards, which were probably used for intra-specific display. The incisors and canine teeth are large, but those at the side are reduced, with a serrated apex, and may have helped to break up plant material, although they were too small to be of much use. The body is large and bulky, indicating a large digestive tract for digesting volumes of plant food. The skull superficially resembles that of ''
Styracocephalus ''Styracocephalus platyrhynchus'' ('spike head') is an extinct species of tapinocephalian therapsids that lived during the Guadalupian epoch. ''Styracocephalus''s head ornament meant that it could be recognised from a distance. The most striki ...
'', but the "horns" are formed from different bones.


Evolutionary Relationships

Estemmenosuchids belong to the Dinocephalian group, a group of early, primitive, but diverse therapsids – often of large size – that are known only from the Middle Permian period. They are however far more primitive and unspecialised than the better known dinocephalians of the South African
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( Beaufort Group), and mostly lived somewhat earlier. They are also unusual in that, despite their primitive nature and early date of appearance, they show herbivorous adaptations. Because of this, there have been two main interpretations of their evolutionary relationships with other Dinocephalia. Hopson and Barghusen in 1986, who provided the first cladistic study of the Therapsida, coined the term Tapinocephalia for herbivorous dinocephalians, as opposed to the " Anteosauria" for the carnivorous forms. They suggested that Estemmenosuchids are very early/primitive members of the Tapinocephalia. However
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(1982) and Gillian King (1988) argue instead that the Estemmenosuchidae are the most
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Dinocephalia, being more primitive than both the Anteosauria and the Tapinocephalia.


Ecological succession

The Estemmenosuchids replaced the caseids as the dominant megaherbivores of the Wordian age (middle of the Middle Permian), before being themselves replaced by the Tapinocephalidae during the Capitanian age (late Middle Permian).


See also

* Permian tetrapods


References


General references

* Olsen, E. C. (1962). " Late Permian terrestrial vertebrates, USA and USSR." '' Transactions of the American Philosophical Society'' 52: 1–224. * Rubidge, B.S. & Sidor, C.A. (2001). "Evolutionary patterns among Permo-Triassic therapsids." '' Annual Review of Ecology and Systematics'' 32: 449–480.


External links


Palaeos
– detailed description

– an earlier page, which was incorporated into the Palaeos material (above) {{Taxonbar, from=Q839455 Tapinocephalians Guadalupian first appearances Guadalupian extinctions Prehistoric therapsid families