Azendohsauridae

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Azendohsauridae
Temporal range: Anisian to Norian
Shringasaurus indicus life restoration.jpg
Life restoration of Shringasaurus indicus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Allokotosauria
Family: Azendohsauridae
Nesbitt et al., 2015
Subgroups

Azendohsauridae is a family of allokotosaurian archosauromorphs that lived during the Middle to Late Triassic period, around 242-216 million years ago. The family was originally named solely for the eponymous Azendohsaurus , marking out its distinctiveness from other allokotosaurs, [1] but as of 2022 the family now includes four other genera: the basal genus Pamelaria , the large horned herbivore Shringasaurus , [2] and two carnivorous genera grouped into the subfamily-level subclade Malerisaurinae, Malerisaurus and Puercosuchus , and potentially also the dubious genus Otischalkia. [3] [4] Most fossils of azendohsaurids have a Gondwanan distribution, with multiple species known across Morocco and Madagascar in Africa as well as India, although fossils of malerisaurine azendohsaurids have also been found in the southwestern United States of North America.

Contents

Azendohsaurids are notable for the various dinosaur-like traits found in some species, including the sauropodomorph-like neck, jaws and teeth of Azendohsaurus, the ceratopsid-like horns of Shringasaurus, and theropod-like teeth of Puercosuchus. [1] [2] [4] These traits are all convergently evolved with later dinosaurs, and some similarities are so striking that it is difficult to distinguish isolated azendohsaurid teeth and jaw bones from those of dinosaurs. Indeed, Azendohsaurus itself was initially described as a herbivorous dinosaur until better remains of its skull and skeleton were found. [5]

Description

Azendohsaurids were robust quadrupeds with sprawled limbs, characterised by their long necks and proportionately small heads, and reached body sizes of up to 3 or 4 metres (9.8 or 13.1 ft) in length in the largest species. [2] Although initially characterised as herbivores based solely on Azendohsaurus, azendohsaurids had a diverse diet and lifestyles including large herbivores, insectivores, and carnivores. Consequently, although broadly similar in body form, their skulls varied from short and boxy with leaf-shaped teeth, to long and narrow with recurved, blade-like teeth. [1] [4] [6]

Azendohsaurids lack a prenarial process, the bony splint of the premaxilla that otherwise divides the two external nares in typical reptile skulls, giving them a single fused (confluent) opening for their nostrils. Another characteristic of azendohsaurids is that they often possess palatal teeth on the roof of their mouths similar in size and shape to those along the jaw margins, including serrations, compared to the typical tiny and conical palatal teeth of other reptiles. Such large and specialised palatal teeth are known definitively in Pamelaria, Azendohsaurus and Shringasaurus, [7] [8] while simpler but notably large palatal teeth with coarse serrations are found in Malerisaurus. [3] The palatal teeth of Puercosuchus, however, are simply described as "peg-like". [4]

All known azendohsaurids have long necks, with neck vertebrae very similar in shape and construction to those of early sauropodomorph dinosaurs, and held them raised above their shoulders. The shoulder girdles themselves are tall and very well developed, with long scapular blades. Azendohsaurids had a sprawling gait like other early archosauromorphs, although their shoulder joint faces back as well as out to the sides, suggesting they may have been able to hold their forelimbs closer to their body. Their bodies are deep, ranging from relatively narrow-bodied in Pamelaria to barrel-shaped in Azendohsaurus, and their tails are proportionally shorter and stockier compared to other archosauromorphs. [1] [4] [6]

Classification and evolution

Azendohsaurids are one of two or three families included in the clade Allokotosauria, a group of unusual Triassic non-archosaur archosauromorphs that also includes the families Trilophosauridae and possibly the gliding Kuehneosauridae. [1] [9] They have consistently been recognised as the sister taxon of trilophosaurids, initially united on shared yet differing herbivorous traits. However, as more azendohsaurids have been discovered and recognised, they demonstrate that the group was likely to be ancestrally carnivorous. [4]

A phylogenetic analysis performed by palaeontologist Sterling J. Nesbitt and colleagues in 2021 included all recognised azendohsaurids and other allokotosaurian taxa. Their results found Pamelaria to be the earliest branching azendohsaurid, with the remaining azendohsaurids divided into two subclades, one containing the herbivores Shringasaurus and two species of Azendohsaurus, and the Malerisaurinae containing both species of Malerisaurus and similar larger material (including the material that would be named Puercosuchus). The cladogram below depicts the simplified consensus result of their analysis, where the uncertain relationships within Malerisaurinae are presented as a polytomy: [3]

Allokotosauria


  1. Representing material referred to ' Otischalkia elderae ' and similar-sized material. [4]
  2. Taxon described as Puercosuchus in 2022. [4]

Biogeography

The oldest known azendohsaurids are Pamelaria and Shringasaurus, both from India and dated to the Anisian stage of the Middle Triassic. Azendohsaurus itself has been dated from the end of the Middle Triassic during the Ladinian (at least for A. madagaskarensis) into the Carnian stage of the earliest Late Triassic in Morocco and Madagascar. [8] Malerisaurines, including Malerisaurus and Puercosuchus, are exclusively known from the Late Triassic during an interval from the late Carnian into the early Norian of both India and North America, and are the latest-surviving azendohsaurids known. This is in spite of their relatively plesiomorphic (i.e. ancestral) features compared to other azendohsaurids, representing a relictual lineage of early-diverging carnivorous azendohsaurids that survived longer than their more derived herbivorous kin. Malerisaurines disappear from the fossil record in North America at or the near the end of the Adamanian teilzone (a local biostratigraphic unit in the southwestern United States) roughly 216 million years ago, marking the extinction of the azendohsaurids globally. [3] [4]

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<i>Azendohsaurus</i> Genus of herbivorous Triassic reptile

Azendohsaurus is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, Azendohsaurus laaroussii, was described and named by Jean-Michel Dutuit in 1972 based on partial jaw fragments and some teeth from Morocco. A second species from Madagascar, A. madagaskarensis, was first described in 2010 by John J. Flynn and colleagues from a multitude of specimens representing almost the entire skeleton. The generic name "Azendoh lizard" is for the village of Azendoh, a local village near where it was first discovered in the Atlas Mountains. It was a bulky quadruped that unlike other early archosauromorphs had a relatively short tail and robust limbs that were held in an odd mix of sprawled hind limbs and raised forelimbs. It had a long neck and a proportionately small head with remarkably sauropod-like jaws and teeth.

<span class="mw-page-title-main">Trilophosauridae</span> Extinct family of reptiles

Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is Trilophosaurus, a herbivore up to 2.5 metres long. It had a short, unusually heavily built skull, equipped with massive, broad flattened cheek teeth with sharp shearing surfaces for cutting up tough plant material. Teeth are absent from the premaxilla and front of the lower jaw, which in life were probably equipped with a horny beak.

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<i>Malerisaurus</i> Extinct genus of reptiles

Malerisaurus is an extinct genus of archosauromorph known from Telangana of India and Texas of the USA.

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Pamelaria is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, Pamelaria dolichotrachela, from the Middle Triassic of India. Pamelaria has sprawling legs, a long neck, and a pointed skull with nostrils positioned at the very tip of the snout. Among early archosauromorphs, Pamelaria is most similar to Prolacerta from the Early Triassic of South Africa and Antarctica. Both have been placed in the family Prolacertidae. Pamelaria, Prolacerta, and various other Permo-Triassic reptiles such as Protorosaurus and Tanystropheus have often been placed in a group of archosauromorphs called Protorosauria, which was regarded as one of the most basal group of archosauromorphs. However, more recent phylogenetic analyses indicate that Pamelaria and Prolacerta are more closely related to Archosauriformes than are Protorosaurus, Tanystropheus, and other protorosaurs, making Protorosauria a polyphyletic grouping.

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<span class="mw-page-title-main">Allokotosauria</span> Extinct clade of reptiles

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Shringasaurus is an extinct genus of archosauromorph reptile from the Middle Triassic (Anisian) of India. It is known from the type and only known species, S. indicus. Shringasaurus is known from the Denwa Formation in the state of Madhya Pradesh. Shringasaurus was an allokotosaur, a group of unusual herbivorous reptiles from the Triassic, and is most closely related to the smaller and better known Azendohsaurus in the family Azendohsauridae. Like some ceratopsid dinosaurs, Shringasaurus had two large horns over its eyes that faced up and forwards from its skull. Shringasaurus also bears convergent physical similarities to sauropodomorph dinosaurs, such as its long neck, its shoulders and forelimbs, and the shape of its teeth. Shringasaurus possibly occupied a similar ecological niche as a large browsing herbivore before such dinosaurs had evolved.

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<i>Puercosuchus</i> Extinct genus of reptile

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References

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