Allokotosauria

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Allokotosauria
Temporal range: MiddleLate Triassic, 247.2–201.3  Ma
Trilophosaurus buettneri - AMNH - DSC06265.JPG
Skeleton of Trilophosaurus buettneri
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Crocopoda
Clade: Allokotosauria
Nesbitt et al., 2015
Families

Allokotosauria is a clade of early archosauromorph reptiles from the Middle to Late Triassic known from Asia, Africa, North America and Europe. Allokotosauria was first described and named when a new monophyletic grouping of specialized herbivorous archosauromorphs was recovered by Sterling J. Nesbitt, John J. Flynn, Adam C. Pritchard, J. Michael Parrish, Lovasoa Ranivoharimanana and André R. Wyss in 2015. The name Allokotosauria is derived from Greek meaning "strange reptiles" in reference to unexpected grouping of early archosauromorph with a high disparity of features typically associated with herbivory.

Contents

History

Nesbitt et al. (2015) defined the group as a stem-based taxon containing Azendohsaurus madagaskarensis and Trilophosaurus buettneri and all taxa more closely related to them than to Tanystropheus longobardicus , Proterosuchus fergusi , Protorosaurus speneri or Rhynchosaurus articeps . Therefore, Allokotosauria includes the families Azendohsauridae and Trilophosauridae by definition, as well as the potentially more basal Pamelaria which is closer to them than to other early archosauromorphs. Pamelaria is the earliest known allokotosaur, dating to the Anisian of India. Azendohsauridae is currently represented by a single genus Azendohsaurus known from the Ladinian to Carnian of Africa, while trilophosaurids are mostly known from the Carnian to Norian stages of North America, England and potentially European Russia, [1] though one member of the latter group, Variodens inopinatus, is known from Rhaetian. [2] According to studies of Arctosaurus material from Cameron Island in Canada, the latter may have been an allokotosaurian because of the similarities with Azendohsaurus due to the presence of a posterior ridge from the centrum to the diapophyses which extends from the diapophysis all the way to the posterior ventrolateral corner of the centrum. This ridge overhangs a deep groove in the lateral surface of the centrum. [3]

Description and phylogeny

Allokotosauria is most notably characterized by wrinkled side surface of orbital border of the frontal bone, expanded and hooked quadrate bone head on the posterior side, and a prominent tubercle developed above to the glenoid fossa of the scapula, although there are other unambiguous traits that differentiate it from other early archosauromorphs. Below is a cladogram showing the phylogenetic relationships of Allokotosauria within Archosauromorpha as recovered by Nesbitt et al. (2015). [1] Ezcurra (2016) also recovered a highly supported Allokotosauria with the same topology (including only Pamelaria , Azendohsaurus madagaskarensis and Trilophosaurus buettneri in his analysis), but noted that Pamelaria is nearly as likely to represent a basal azendohsaurid instead. [4]

Archosauromorpha

Protorosaurus speneri

Tanystropheidae

Rhynchosauria

Crocopoda [4]
Allokotosauria

Pamelaria dolichotrachela

Trilophosauridae

Teraterpeton hrynewichorum

Trilophosaurus buettneri

Spinosuchus caseanus (= "Trilophosaurus" jacobsi)

Azendohsauridae

Azendohsaurus laaroussii

Azendohsaurus madagaskarensis

Prolacerta broomi

Archosauriformes

Sengupta et al. (2017) described a new azendohsaurid and recovered Pamelaria as an azendohsaurid. [5]

Archosauromorpha

Choristodera

Aenigmastropheus

Protorosaurus

Prolacertoides

Jesairosaurus

Tanystropheidae

Allokotosauria
Trilophosauridae

Teraterpeton hrynewichorum

Trilophosaurus buettneri

Spinosuchus caseanus (= "Trilophosaurus" jacobsi)

Azendohsauridae

Pamelaria dolichotrachela

Shringasaurus indicus

Azendohsaurus laaroussii

Azendohsaurus madagaskarensis

Rhynchosauria

Boreopricea

Prolacertidae

Tasmaniosaurus

Archosauriformes

Pritchard and Nesbitt (2017) found support for the inclusion of Kuehneosauridae within Allokotosauria. [6]

Related Research Articles

<span class="mw-page-title-main">Archosauromorpha</span> Infraclass of reptiles

Archosauromorpha is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs rather than lepidosaurs. Archosauromorphs first appeared during the late Middle Permian or Late Permian, though they became much more common and diverse during the Triassic period.

<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.

Spinosuchus is an extinct genus of trilophosaurid allokotosaur from the Late Triassic of Texas, southern United States. It has been assigned to a variety of groups over its history, from coelophysid dinosaur to pseudosuchian to uncertain theropod dinosaur and to Proterosuchidae. This uncertainty is not unusual, given that it was only known from a poorly preserved, wall-mounted, partial vertebral column of an animal that lived in a time of diverse, poorly known reptile groups. However, newly collected material and recent phylogenetic studies of early archosauromorphs suggest that it represents an advanced trilophosaurid very closely related to Trilophosaurus.

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

Trilophosaurus is a lizard-like trilophosaurid allokotosaur known from the Late Triassic of North America. It was a herbivore up to 2.5 m 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.

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

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

Sterling Nesbitt is an American paleontologist best known for his work on the origin and early evolutionary patterns of archosaurs. He is currently an associate professor at Virginia Tech in the Department of Geosciences.

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

Doswelliidae is an extinct family of carnivorous archosauriform reptiles that lived in North America and Europe during the Middle to Late Triassic period. Long represented solely by the heavily-armored reptile Doswellia, the family's composition has expanded since 2011, although two supposed South American doswelliids were later redescribed as erpetosuchids. Doswelliids were not true archosaurs, but they were close relatives and some studies have considered them among the most derived non-archosaurian archosauriforms. They may have also been related to the Proterochampsidae, a South American family of crocodile-like archosauriforms.

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

Teraterpeton is an extinct genus of trilophosaurid archosauromorphs. It is known from a partial skeleton from the Late Triassic Wolfville Formation of Nova Scotia, described in 2003. It has many unique features seen in no other related form, including an elongated, toothless snout and large openings for the nostrils. Because of this, Teraterpeton was originally placed in its own family, Teraterpetidae, related to Trilophosaurus. Newer studies generally place it within Trilophosauridae.

<span class="mw-page-title-main">Protorosauria</span> Extinct order of reptiles

Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian to the end of the Late Triassic of Asia, Europe and North America. It was named by the English anatomist and paleontologist Thomas Henry Huxley in 1871 as an order, originally to solely contain Protorosaurus. Other names which were once considered equivalent to Protorosauria include Prolacertiformes and Prolacertilia.

<span class="mw-page-title-main">Epipophyses</span> Bony projections of neck vertebrae in certain reptiles

Epipophyses are bony projections of the cervical vertebrae found in archosauromorphs, particularly dinosaurs. These paired processes sit above the postzygapophyses on the rear of the vertebral neural arch. Their morphology is variable and ranges from small, simple, hill-like elevations to large, complex, winglike projections. Epipophyses provided large attachment areas for several neck muscles; large epipophyses are therefore indicative of a strong neck musculature.

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

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.

The Yerrapalli Formation is a Triassic rock formation consisting primarily of mudstones that outcrops in the Pranhita–Godavari Basin in southeastern India. The Yerrapalli Formation preserves fossils of freshwater and terrestrial vertebrates as well as trace fossils of invertebrates. The tetrapod fauna includes temnospondyl amphibians, archosauromorph reptiles, and dicynodonts.

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

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, but as of 2022 the family now includes four other genera: the basal genus Pamelaria, the large horned herbivore Shringasaurus, and two carnivorous genera grouped into the subfamily-level subclade Malerisaurinae, Malerisaurus and Puercosuchus, and potentially also the dubious genus Otischalkia. 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.

<i>Ozimek volans</i> Extinct species of reptile

Ozimek is a genus of sharovipterygid archosauromorph reptile, known from Late Triassic deposits in Poland and closely related to the Kyrgyzstani Sharovipteryx. It contains one species, O. volans, named in 2016 by Jerzy Dzik and Tomasz Sulej. Like Sharovipteryx, Ozimek had long, slender limbs with the hindlimbs longer than the forelimbs; the hindlimbs likely supported gliding membranes as fossilized in Sharovipteryx. Another unusual characteristic was the shoulder girdle, where the massive coracoids formed a shield-like structure covering the bottom of the shoulder region that would have limited mobility. In other respects, such as its long neck, it was a typical member of the non-natural grouping Protorosauria. Phylogenetic analysis has indicated that it, possibly along with Sharovipteryx, may have been an unusual member of the protorosaur group Tanystropheidae, although further study of its anatomy is needed to resolve its precise relationships.

<i>Shringasaurus</i> Genus of herbivorous Triassic horned reptile

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.

Boreopricea is an extinct genus of archosauromorph reptile from the Early Triassic of arctic Russia. It is known from a fairly complete skeleton discovered in a borehole on Kolguyev Island, though damage to the specimen and loss of certain bones has complicated study of the genus. Boreopricea shared many similarities with various other archosauromorphs, making its classification controversial. Various studies have considered it a close relative of Prolacerta, tanystropheids, both, or neither. Boreopricea is unique among early archosauromorphs due to possessing contact between the jugal and squamosal bones at the rear half of the skull.

Isalo II, also known as the Makay Formation, is an informal Triassic geological unit in Madagascar.

Puercosuchus is an extinct genus of archosauromorph reptile from the Late Triassic (Norian) of what is now Arizona, North America. It includes only the type species P. traverorum, and was described and named in 2022. Puercosuchus is known mainly from two bonebeds in the Blue Mesa Member of the Chinle Formation, preserving the mixed remains of multiple individuals in each one representing almost the entire skeleton. It is a member of the Azendohsauridae, a clade of Triassic reptiles that was initially recognised by adaptations for herbivory. However, Puercosuchus and its close relatives in the subclade Malerisaurinae retained the carnivorous diet and body form ancestral to archosauromorphs. Unlike non-malerisaurine azendohsaurids, Puercosuchus had a long and shallow snout with sharp, blade-like teeth similar to those of carnivorous dinosaurs. Despite its seemingly ancestral morphology and ecology, Puercosuchus is the youngest known genus of azendohsaurid in the world. The discovery of Puercosuchus allowed palaeontologists to recognise similar bones and teeth that had been collected from Late Triassic southwestern North America in the past as belonging to it or similar animals, acting as a sort of "rosetta stone" for malerisaurine azendohsaurid anatomy.

Mambachiton is an extinct genus of basal avemetatarsalian from the Middle/Upper Triassic Makay Formation of Madagascar. The genus contains a single species, M. fiandohana, known from a partial skeleton with articulated osteoderms.

References

  1. 1 2 Nesbitt, S.J.; Flynn, J.J.; Pritchard, A.C.; Parrish, M.J.; Ranivoharimanana, L.; Wyss, A.R. (2015). "Postcranial osteology of Azendohsaurus madagaskarensis (?Middle to Upper Triassic, Isalo Group, Madagascar) and its systematic position among stem archosaur reptiles" (PDF). Bulletin of the American Museum of Natural History. 398: 1–126. doi:10.5531/sd.sp.15. hdl: 2246/6624 . ISSN   0003-0090.
  2. David I. Whiteside, FLS; Christopher J. Duffin, FLS (2017). "Late Triassic terrestrial microvertebrates from Charles Moore's "Microlestes" quarry, Holwell, Somerset, UK". Zoological Journal of the Linnean Society. 179 (3): 677–705. doi: 10.1111/zoj.12458 .
  3. Sues, Hans-Dieter (February 2017). "Arctosaurus osborni, a Late Triassic archosauromorph reptile from the Canadian Arctic Archipelago". Canadian Journal of Earth Sciences. 54 (2): 129–133. Bibcode:2017CaJES..54..129S. doi: 10.1139/cjes-2016-0159 .
  4. 1 2 Ezcurra, M.D. (2016). "The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms". PeerJ. 4: e1778. doi: 10.7717/peerj.1778 . PMC   4860341 . PMID   27162705.
  5. Sengupta, S.; Ezcurra, M.D.; Bandyopadhyay, S. (2017). "A new horned and long-necked herbivorous stem-archosaur from the Middle Triassic of India". Scientific Reports. 7 (1): 8366. Bibcode:2017NatSR...7.8366S. doi:10.1038/s41598-017-08658-8. PMC   5567049 . PMID   28827583.
  6. Pritchard, Adam C.; Nesbitt, Sterling J. (2017-10-11). "A bird-like skull in a Triassic diapsid reptile increases heterogeneity of the morphological and phylogenetic radiation of Diapsida". Royal Society Open Science. 4 (10): 170499. Bibcode:2017RSOS....470499P. doi:10.1098/rsos.170499. ISSN   2054-5703. PMC   5666248 . PMID   29134065.