Aetosaurinae Temporal range: Late Triassic | |
---|---|
A bonebed of Aetosaurus | |
Paratypothorax , a typothoracine | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Pseudosuchia |
Order: | † Aetosauria |
Family: | † Stagonolepididae |
Subfamily: | † Aetosaurinae Heckert and Lucas, 2000 |
Clades | |
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. [1]
Aetosaurinae was originally named in 2000, as a subfamily solely including Aetosaurus, which was assumed to be the earliest-diverging aetosaur. [2] In 2007, it was extended to include the subfamily Typothoracinae as well as various basal aetosaurs which were not clearly within Desmatosuchinae. These proposed non-typothoracine aetosaurines included Coahomasuchus, Neoaetosauroides , Aetosauroides , Stagonolepis robertsoni, and "Stagonolepis" ( Calyptosuchus ) wellesi. As a subfamily containing practically all non-desmatosuchine aetosaurs, Aetosaurinae was poorly-supported. It was diagnosed by a singly synapomorphy: the dorsal eminence of each paramedian osteoderm was offset medially. [3] [4]
A phylogenetic study in 2012 found that this widest usage of Aetosaurinae was paraphyletic. Aetosaurus was recovered as the basal-most stagonolepidid, with other non-typothoracisine aetosaurines placed as successively more derived taxa leading up to a clade containing Desmatosuchinae and Typothoracisinae. In this phylogeny, all aetosaurines except for Aetosaurus were more closely related to Desmatosuchus than they are to Aetosaurus. As a result, the clade Aetosaurinae would only include Aetosaurus. [5] [6]
More recent data and phylogenies have brought Aetosaurinae back into usage, albeit in a more restricted form. A 2016 study once again considered Typothoracinae to be closer to Aetosaurus than to desmatosuchines. Therefore, Typothoracinae would lie within Aetosaurinae, making Aetosaurinae a valid and non-monotypic clade once more. [1] Newly described or redescribed genera such as Stenomyti and Coahomasuchus have also been incorporated into Aetosaurinae. [1] [7] Aetosaurinae is the sister taxon to Desmatosuchia, a new clade including desmatosuchines and their relatives, such as Stagonolepis. [1]
Relative to desmatosuchians, aetosaurines have proportionally shorter and more lightly built skulls, as well as several other distinctive traits. The orbit is large and circular, while the supratemporal fenestrae are fairly small. This also means that the parietals are wider than the frontals on the skull roof, at least in early-diverging genera such as Aetosaurus and Stenomyti. The nares are similar in size or smaller than the orbit. The snout tapers to a point, without the "shovel-shaped" premaxilla extensions found in Stagonolepis and Desmatosuchus . [1] [8]
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.
Aetosaurus is an extinct genus of pseudosuchian reptile belonging to the order Aetosauria. It is generally considered to be the most primitive aetosaur. Three species are currently recognized: A. ferratus, the type species from Germany and Italy; A. crassicauda from Germany; and A. arcuatus from eastern North America. Additional specimens referred to Aetosaurus have been found in the Chinle Group of the southwestern United States, and the Fleming Fjord Formation of Greenland. Specimens of Aetosaurus occur in Norian-age strata.
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.
Aetosauroides is an extinct genus of aetosaur from the Late Triassic of South America. It is one of four aetosaurs known from South America, the others being Neoaetosauroides, Chilenosuchus and Aetobarbakinoides. Three species have been named: the type species A. scagliai, A. subsulcatus and A. inhamandensis. Fossils have been found in the Cancha de Bochas Member of the Ischigualasto Formation in the Ischigualasto-Villa Unión Basin in northwestern Argentina and the Santa Maria Formation in the Paraná Basin in southeastern Brazil. The strata date to the late Carnian and early Norian stages, making Aetosauroides one of the oldest aetosaurs.
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.
Rioarribasuchus is a genus of aetosaur. Fossils have been found from the Chinle Formation in Arizona and New Mexico that date back to the upper Late Carnian stage of the Late Triassic.
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.
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.
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.
Sierritasuchus is an extinct genus of aetosaur in the subfamily Desmatosuchinae. It is known from a small holotype skeleton from the Late Triassic Tecovas Formation of Texas. This skeleton was discovered in 1939 and was originally assigned to the genus Desmatosuchus. It was placed in its own genus in 2008 after having been in the collections of the University of Michigan Museum of Paleontology, with the type species being S. macalpini. The generic name refers to Sierrita de la Cruz Creek where the holotype was found, and the specific name refers to Archie MacAlpin, who discovered the skeleton. Based on the histology of the scutes of the holotype, the individual was a subadult that was not fully grown.
Suchia is a clade of archosaurs containing the majority of pseudosuchians. It was defined as the least inclusive clade containing Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, and Crocodylus niloticus by Nesbitt (2011). Generally the only pseudosuchian group which is omitted from Suchia is the family Ornithosuchidae, although at least one analysis classifies ornithosuchids as close relatives of erpetosuchids and aetosaurs. Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place.
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.
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. Gorgotesuchus is typically considered a basal desmatosuchin, though alternative interpretations exist.
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.