Estemmenosuchidae

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Estemmenosuchidae
Temporal range: Wordian
Estemmenosuchus mirabilis.jpg
Life restoration of Estemmenosuchus mirabilis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Suborder: Dinocephalia
Infraorder: Tapinocephalia (?)
Family: Estemmenosuchidae
Tchudinov, 1960

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 display or agonistic behavior. Apart from the best known genus, Estemmenosuchus , 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).

Contents

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 , 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 Karoo (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. [1] They suggested that Estemmenosuchids are very early/primitive members of the Tapinocephalia.

However Thomas Kemp (1982) and Gillian King (1988) argue instead that the Estemmenosuchidae are the most basal Dinocephalia, being more primitive than both the Anteosauria and the Tapinocephalia. [2] [3]

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

Related Research Articles

<span class="mw-page-title-main">Therapsid</span> Clade of tetrapods including mammals

A therapsid is a member of the clade Therapsida which is a major group of eupelycosaurian synapsids that includes mammals and their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, as opposed to the sprawling posture of many reptiles and salamanders.

<span class="mw-page-title-main">Dinocephalia</span> Extinct clade of stem-mammals

Dinocephalians are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. Dinocephalians included herbivorous, carnivorous, and omnivorous forms. Many species had thickened skulls with many knobs and bony projections. Dinocephalians were the first non-mammalian therapsids to be scientifically described and their fossils are known from Russia, China, Brazil, South Africa, Zimbabwe, and Tanzania.

<span class="mw-page-title-main">Biarmosuchia</span> Extinct suborder of therapsids

Biarmosuchia is an extinct clade of non-mammalian synapsids from the Permian. Biarmosuchians are the most basal group of the therapsids. They were moderately-sized, lightly built carnivores, intermediate in form between basal sphenacodont "pelycosaurs" and more advanced therapsids. Biarmosuchians were rare components of Permian ecosystems, and the majority of species belong to the clade Burnetiamorpha, which are characterized by elaborate cranial ornamentation.

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

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

<span class="mw-page-title-main">Anteosauridae</span> Extinct family of therapsids

Anteosauridae is an extinct family of large carnivorous dinocephalian therapsids that are known from the Middle Permian of Asia, Africa, and South America.These animals were by far the largest predators of the Permian period, with skulls reaching 80 cm in length in adult individuals, far larger than the biggest gorgonopsian.

<span class="mw-page-title-main">Therocephalia</span> Extinct order of therapsids

Therocephalia is an extinct clade of eutheriodont therapsids from the Permian and Triassic periods. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their teeth, suggest that they were carnivores. Like other non-mammalian synapsids, therocephalians were once described as "mammal-like reptiles". Therocephalia is the group most closely related to the cynodonts, which gave rise to the mammals, and this relationship takes evidence in a variety of skeletal features. Indeed, it had been proposed that cynodonts may have evolved from therocephalians and so that therocephalians as recognised are paraphyletic in relation to cynodonts.

<span class="mw-page-title-main">Theriodontia</span> Clade of therapsids

The theriodonts are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts.

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

Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian stage, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. Measuring 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb), Anteosaurus was the largest known carnivorous non-mammalian synapsid and the largest terrestrial predator of the Permian period. Occupying the top of the food chain in the Middle Permian, its skull, jaws and teeth show adaptations to capture large prey like the giants titanosuchids and tapinocephalids dinocephalians and large pareiasaurs.

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

Struthiocephalus is an extinct genus of dinocephalian therapsids from the Permian of South Africa. It was a large animal, reaching 288 kg (635 lb) in body mass.

<i>Brithopus</i> Extinct genus of mammal ancestors

Brithopus is an extinct genus of dinocephalian therapsids. It contains a single species, Brithopus priscus, known from fragmentary remains found in the Copper Sandstones near Isheevo, Russia.

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

Australosyodon is an extinct genus of dinocephalian therapsids from the middle Permian of South Africa. The first fossil was discovered in the 1980s near the village of Prince Albert Road in the Karoo region of South Africa.

<i>Styracocephalus</i>

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

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

Venyukovia is an extinct genus of venyukovioid therapsid, a basal anomodont from the Middle Permian of Russia. The type and sole species, V. prima, is known only by a partial lower jaw with teeth. Venyukovia has often been incorrectly spelt as 'Venjukovia' in English literature. This stems from a spelling error made by Russian palaeontologist Ivan Efremov in 1940, who mistakenly replaced the 'y' with a 'j', which subsequently permeated through therapsid literature before the mistake was caught and corrected. Venyukovia is the namesake for the Venyukovioidea, a group of small Russian basal anomodonts also including the closely related Otsheria, Suminia, Parasuminia and Ulemica, although it itself is also one of the poorest known. Like other venyukovioids, it had large projecting incisor-like teeth at the front and lacked canines, although the remaining teeth are simple compared to some other venyukovioids, but may resemble those of Otsheria.

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

Niaftasuchus is an extinct genus of therapsids. Its type and only named species is Niaftasuchus zekkeli.

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

Raranimus is an extinct genus of therapsids of the Middle Permian. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the Qingtoushan Formation, outcropping in the Qilian Mountains of Gansu, China. The genus is the most basal known member of the clade Therapsida, to which the later Mammalia belong.

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

Herpetoskylax is an extinct genus of biarmosuchians which existed in South Africa. The type species is Herpetoskylax hopsoni. It lived in the Late Permian Period.

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

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

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

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

<span class="mw-page-title-main">Eutheriodontia</span> Clade of therapsids

Eutheriodontia is a clade of therapsids which appear during the Middle Permian and which includes therocephalians and cynodonts, this latter group including mammals and related forms.

<span class="mw-page-title-main">Phthinosuchia</span> Extinct infraorder of mammals

Phthinosuchia is an extinct group of therapsids including two poorly known species, Phthinosuchus discors and Phthinosaurus borrisiaki, from the Middle Permian of Russia. Phthinthosuchus is known a partial crushed skull and Phthinosaurus is known from an isolated lower jaw. The two species have traditionally been grouped together based on their shared primitive characteristics, but more recent studies have proposed that they are more distantly related. Phthinosuchus is either a carnivorous gorgonopsian relative or an anteosaurian dinocephalian while Phthinosaurus is either a herbivorous rhopalodont dinocephalian or a therocephalian.

References

  1. Hopson, J. A.; Barghusen, H. (1986). "An analysis of therapsid relationships". In N. Hotton, III; P. D. MacLean; J. J. Roth; E. C. Roth (eds.). The Ecology and Biology of Mammal-like Reptiles. Smithsonian Institution Press. pp. 83–106.
  2. Kemp, T. S. (1982). Mammal-like Reptiles and the Origin of Mammals. New York: Academic Press. pp. 363pp.
  3. King, G. M. (1988). "Anomodontia". Encyclopedia of Paleoherpetology. Vol. Part 17 C. Stuttgart and New York: Gutsav Fischer Verlag.

General references