Gorgetosuchus Temporal range: Late Triassic | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Order: | † Aetosauria |
Family: | † Stagonolepididae |
Subfamily: | † Desmatosuchinae |
Genus: | † Gorgetosuchus Heckert et al., 2015 |
Type species | |
†Gorgetosuchus pekinensis Heckert et al., 2015 |
Gorgetosuchus is an extinct genus of aetosaur from the Late Triassic of the North Carolina, represented by the type species Gorgetosuchus pekinensis. It is mainly known from osteoderms, including the front half of an articulated carapace. [1] Gorgotesuchus is typically considered a basal desmatosuchin, [1] [2] though alternative interpretations exist. [3]
G. pekinensis was named and described by Heckert et al. (2015) on the basis of ten rows of bony plates called osteoderms, representing the front part of an armored carapace that would have covered the back of the animal. These plates were found embedded in sandstone and conglomerate boulders near a brick quarry in Chatham County, North Carolina, which likely originated from the Late Triassic Pekin Formation. (The Pekin Formation consists of interbedded red mudstones, siltstones, sandstones and conglomerates; it was the fine-grained mudstones and siltstones that the mining operation was targeting for brick-making, so coarser-grained blocks were removed from the pit). Gorgetosuchus was part of a diverse faunal community in a continental rift valley system during the Late Triassic, which also included traversodontids, phytosaurs, dicynodonts, and temnospondyls. Gorgetosuchus coexisted with two other aetosaurs, Lucasuchus and Coahomasuchus . [1]
Gorgetosuchus is unique among aetosaurs in having cervical (neck) osteoderms that nearly wrap around the entire neck and are strongly angled to give the neck a hexagonal shape in cross-section. The lateral (side) and paramedian (upper) osteoderms each bear prominent spines. Other aetosaurs such as Longosuchus also have neck spines, but only on the lateral osteoderms. The cervical osteoderms are wider than they are long, a feature that unites Gorgetosuchus with basal "aetosaurine" aetosaurs. However, several features of the osteoderms (such as a flange on the lateral cervical osteoderms that overlaps the paramedian osteoderms) link it with the desmatosuchines, a more derived group of aetosaurs.
Among aetosaurs, Gorgetosuchus closely resembles Lucasuchus and Longosuchus, both of which are desmatosuchines. A phylogenetic analysis conducted by Heckert et al. (2015) placed Gorgetosuchus close to these taxa as the most basal desmatosuchine, but the results were not statistically well supported. Below is a strict consensus tree from their analysis showing the placement of Gorgetosuchus: [1]
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Gorgetosuchus was also placed as the basal-most member of Desmatosuchini (traditional desmatosuchines) in an analysis by Parker (2016). [2] However, an analysis by Hoffman et al. (2018), based on an earlier analysis by Schoch & Desojo (2016), placed it within Typothoracinae, closely related to Typothorax and Redondasuchus . Typothoracines and desmatosuchins are known to have a large degree of homoplasy (convergent evolution) in certain osteoderm traits observed in Gorgetosuchus. On the other hand, other skeletal traits strongly diverge between the two groups, so the position of Gorgetosuchus would likely stabilize if more non-osteoderm bones were discovered. [3]
Aetosaurs are heavily armored reptiles belonging to the extinct order Aetosauria. They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs more closely related to crocodilians than to birds and other dinosaurs. All known aetosaurs are restricted to the Late Triassic, and in some strata from this time they are among the most abundant fossil vertebrates. They have small heads, upturned snouts, erect limbs, and a body ornamented with four rows of plate-like osteoderms. Aetosaur fossil remains are known from Europe, North and South America, parts of Africa, and India. Since their armoured plates are often preserved and are abundant in certain localities, aetosaurs serve as important Late Triassic tetrapod index fossils. Many aetosaurs had wide geographic ranges, but their stratigraphic ranges were relatively short. Therefore, the presence of particular aetosaurs can accurately date a site in which they are found.
Stagonolepis is an extinct genus of stagonolepidid aetosaur known from the Late Triassic Hassberge Formation of Germany, the Drawno Beds of Poland, and the Lossiemouth Sandstone of Scotland. Supposed fossils from North and South America have been placed into their own genera, Calyptosuchus and Aetosauroides, respectively.
Calyptosuchus is an extinct genus of aetosaur from the Late Triassic of North America. Like other aetosaurs, it was heavily armored and had a pig-like snout used to uproot plants.
Acaenasuchus is an extinct genus of pseudosuchian, endemic to what would be presently be known as Arizona during the Late Triassic, specifically during the Carnian and Norian stages of the Triassic. Acaenasuchus had a stratigraphic range of approximately 11.5 million years. Acaenasuchus is further categorized as one of the type fauna that belong to the Adamanian LVF, based on the fauna of the Blue Mesa Member of the Chinle Petrified Forest Formation of Arizona, where Acaenasuchus was initially discovered.
Longosuchus is an extinct genus of desmatosuchin aetosaur from the Late Triassic of North America. Reported fossils from Morocco are likely misidentified scutes of a paratypothoracin aetosaur. Longosuchus measured about 3 metres in length.
Typothorax is an extinct genus of typothoracine aetosaur that lived in the Late Triassic. Its remains have been found in North America. Two species are known: T. coccinarum, the type species, and T. antiquum.
Coahomasuchus is an extinct genus of aetosaurine aetosaur. Remains of the genus have been found from deposits in Texas and North Carolina that date to the Otischalkian faunachron of the Late Triassic. It was small for an aetosaur, being less than 1.5 metres long. The dorsal plates are distinctively flat and unflexed, and have a faint sub-parallel to radial ornamentation. The genus lacked spines or keels on these plates, features seen in many other aetosaurs. Coahomasuchus was very similar in appearance to the closely related Aetosaurus.
Lucasuchus is an extinct genus of aetosaur. Fossils have been found from the Bull Canyon Formation of the Dockum Group outcropping in the Revuelto Creek locality in Quay County, New Mexico. All specimens date back to the Norian stage of the Late Triassic. The genus was named in 1995 after the American paleontologist Spencer G. Lucas.
Paratypothorax is an extinct genus of aetosaur, known from a single species, Paratypothorax andressorum. It was a broadly distributed member of the group found in Germany, North America, and possibly parts of Gondwana. The best specimens come from Germany, though for more than a century they were mistakenly considered phytosaur armor. Paratypothorax was a large and wide-bodied typothoracine aetosaur, as well as the namesake of the tribe Paratypothoracisini.
Typothoracinae is a clade of aetosaurs within the subfamily Aetosaurinae. It was originally defined as a stem-based taxon including all aetosaurs closer to Typothorax than to Stagonolepis or Desmatosuchus. This definition was later expanded to specifically exclude Aetosaurus; as of 2016, Typothoracinae is defined as the least inclusive clade containing Typothorax and Paratypothorax, but not Aetosaurus,Stagonolepis, or Desmatosuchus. The clade was first named in 2007 under the spelling Typothoracisinae, after its namesake Typothorax. However, this spelling was based on incorrect taxonomic nomenclature, and the clade's name was corrected to Typothoracinae in 2016.
Aetosaurinae is one of the two main clades of aetosaurs, the other being Desmatosuchia. It is a stem-based taxon defined as all aetosaurs more closely related to Aetosaurus than Desmatosuchus. Aetosaurinae currently comprises Aetosaurus, similar forms such as Coahomasuchus and Stenomyti, and the widespread and successful aetosaur clade Typothoracinae.
Paratypothoracini is a clade of aetosaurs within the group Typothoracinae. It is a node-based taxon that includes Rioarribasuchus (=Heliocanthus), Paratypothorax, Tecovasuchus, and all descendants of their most recent common ancestor. The clade was first named in 2007 under the spelling Paraypothoracisini, after its namesake Paratypothorax. However, this spelling was based on incorrect taxonomic nomenclature, and the clade's name was corrected to Paratypothoracinae in 2016.
Desmatosuchinae is a major subfamily of aetosaurs within the clade Desmatosuchia. It is a stem-based taxon defined as all aetosaurs more closely related to Desmatosuchus than to Stagonolepis,Aetosaurus, or Paratypothorax.
Tarjadia is an extinct genus of erpetosuchid pseudosuchian, distantly related to modern crocodilians. It is known from a single species, T. ruthae, first described in 1998 from the Middle Triassic Chañares Formation in Argentina. Partial remains have been found from deposits that are Anisian-Ladinian in age. Long known mostly from osteoderms, vertebrae, and fragments of the skull, specimens described in 2017 provided much more anatomical details and showed that it was a fairly large predator. Tarjadia predates known species of aetosaurs and phytosaurs, two Late Triassic groups of crurotarsans with heavy plating, making it one of the first heavily armored archosaurs. Prior to 2017, most studies placed it outside Archosauria as a member of Doswelliidae, a family of heavily armored and crocodile-like archosauriforms. The 2017 specimens instead show that it belonged to the Erpetosuchidae.
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.
Aetobarbakinoides is an extinct genus of stagonolepidid aetosaur known from the Late Triassic of Rio Grande do Sul state, southern Brazil. Fossils have been found from the Santa Maria Supersequence of the late Carnian and early Norian stages, making Aetobarbakinoides one of the oldest aetosaurs. The type species, A. brasiliensis, was named in 2012, and is notable for being described primarily by features of the vertebrae; most other aetosaurs are diagnosed by features in bony plates called osteoderms, which are by far the most common material. Although placed in a basal phylogenetic position among aetosaurs, Aetobarbakinoides is closely related to both desmatosuchines and typothoracisines, two derived clades of aetosaurs.
Polesinesuchus is an extinct genus of stagonolepidid aetosaur known from the Late Triassic of southern Brazil. Fossils have been found from the Santa Maria Supersequence of the late Carnian and early Norian stages, making Polesinesuchus one of the oldest aetosaurs. It contains a single species, Polesinesuchus aurelioi, the fifth aetosaur species known from South America to date. Anatomical evidence suggests that Polesinesuchus likely represents a juvenile individual of the contemporary Aetosauroides.
Scutarx is an extinct genus of Aetosauriformes, most commonly regarded by its species name Scutarx deltatylus. Scutarx lived around 230 million years ago during the Carnian and Norian stage of the Late Triassic. Scutarx are “medium sized” paramedian osteoderms belonging to the clade Aetosauria, a heavily armored and more herbivorous cousin of crocodiles.
Apachesuchus is an extinct genus of aetosaur from the Late Triassic of New Mexico. It is only known from several paramedian osteoderms discovered in Quay County in eastern New Mexico. This area belongs to the late Norian-age Quay Member of the Redonda Formation. Unique among aetosaurs, its osteoderms are nearly completely smooth, without strong pits or grooves. The left dorsal paramedian has a relatively high width-to-length ration, suggesting that Apachesuchus is a wide-bodied aetosaur within the clade Typothoracinae.
Venkatasuchus is an extinct genus of aetosaur from the Late Triassic Dharmaram Formation of India. It was described in 2023 on the basis of a series of associated osteoderms that formed the paramedian and lateral armour. Based on the osteoderms the carapace of Venkatasuchus was disc-shaped and very wide, with curved, horn-like elements along its sides. Phylogenetic analysis indicates that Venkatasuchus belonged to the subfamily Typothoracinae and more specifically the clade Paratypothoracini, where it is recovered as the sister taxon to Kocurypelta. Venkatasuchus is among the few aetosaurs recovered from the region that would later become Gondwana and lends credence to the idea that late Triassic India represented a connective hub between Laurasian and Gondwanan fauna. The genus is monotypic, meaning it only includes a single species, Venkatasuchus armatum.