Yonghesuchus Temporal range: Ladinian, | |
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Skull | |
Life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Family: | † Gracilisuchidae |
Genus: | † Yonghesuchus Wu et al., 2001 |
Type species | |
†Yonghesuchus sangbiensis Wu et al., 2001 |
Yonghesuchus is an extinct genus of Late Triassic archosaur reptile. Remains have been found from the early Late Triassic Tongchuan Formation in Shanxi, China. It is named after Yonghe County, the county where fossils were found. Currently only one species, Y. sangbiensis, is known. The specific name refers to Sangbi Creek, as fossils were found in one of its banks. [1]
Yonghesuchus is known from two skulls, one with an attached mandible (the holotype) and one with articulated cervical vertebrae. The holotype skull, known as IVPP V 12378, has been deformed by compression during preservation so that it has become dorsoventrally flattened. The paratype skull, known as IVPP V 12379, has also experienced damage as a result of its preservation. [1]
The premaxilla, the bone at the front of the snout, projects past the front teeth to form a small pointed tip at the end of the snout. Behind the premaxilla is the maxilla, which contains a small depression that forms part of the antorbital fossa. This depression distinguishes Yonghesuchus from related archosauriforms such as Turfanosuchus . As in other archosauriforms such as Turfanosuchus and Euparkeria , the palate is covered in very small teeth called denticles. Behind the palate, the basisphenoid bone (which forms the floor of the braincase) is long and narrows toward the front. This is another distinguishing characteristic of Yonghesuchus, as other early archosauriforms have shorter and wider basisphenoid bones. Moreover, the entrance of the internal carotid artery, which passes through a foramen in the basisphenoid to supply blood to the brain, is in a different position than related genera. Its position is more similar to that of Dorosuchus (a euparkeriid) and more derived archosauriforms. [1]
In the lower jaw, the dentary bone has two projections at its posterior end where it attaches to the mandible, the higher one being markedly longer than the lower one. The high projection comprises much of the upper margin of the mandibular fenestra, an opening along the side of the jaw. The mandibular fenestra is longer and narrower than those of related archosauriforms such as Turfanosuchus, Euparkeria, and Ornithosuchus . At the back of the jaw is the retroarticular region, which extends backward from the jaw joint. On the dorsal surface of this region on the articular bone is a prominent ridge that is not seen in other archosauriforms. There is a wing-like projection on the medial, or inner, surface of the articular which is called the medial process. This process is also seen in sphenosuchian crocodylomorphs and rauisuchians. [1]
Yonghesuchus, like Turfanosuchus, has small, compressed, recurved premaxillary teeth in the front of the upper jaw. The maxillary teeth are larger, more compressed, and serrated. The largest of these are the fourth and fifth maxillary teeth (the ninth and tenth teeth from the tip of the jaw). Teeth behind these get progressively smaller and end below the orbit, or eye socket. [1]
The cervical vertebrae, which are only known in the paratype, are amphicoelous, meaning that they are concave at both ends. These vertebrae bear small ribs that are similar to other archosauriformes, including crocodyliforms. [1]
Yonghesuchus was not considered to be a crown-group archosaur, but rather a closely related advanced archosauriform. One feature that excluded it from Archosauria was its palatal teeth, which are not found in any archosaur (but are known to vary in other clades [2] ). Yonghesuchus was considered to be more closely related to archosaurs than the related family Proterochampsidae based on the position of the foramen for the carotid artery on the basisphenoid bone, which in Yonghesuchus was presumably more similar to that seen in archosaurs. The position of the foramen has also been an indication that Yonghesuchus is more closely related to archosaurs than Turfanosuchus, which has a foramen in a position that is plesiomorphic in archosauriformes and similar to proterochampsids. [1]
Xu et al. (2001), the first to describe Yonghesuchus, suggested that among non-archosaurian archosauriformes, Yonghesuchus was the most closely related, followed by Proterochampsidae, Turfanosuchus, and Euparkeria. [1] Xu et al. supported this claim with the geological position of these taxa, which are successively older from Yonghesuchus (early Late Triassic) to Euparkeria (late Early Triassic). A later phylogenetic analysis on basal archosauriforms by Dilkes and Sues (2009) placed proterochampsids in a more basal position than Turfanosuchus and Yonghesuchus, which were considered successive sister taxa to Archosauria. [3]
In most recent analyses, Yonghesuchus has been recovered as in a well supported clade, the Gracilisuchidae within the Pseudosuchia, along with Gracilisuchus and Turfanosuchus. [4]
Archosauriformes is a clade of diapsid reptiles encompassing archosaurs and some of their close relatives. It was defined by Jacques Gauthier (1994) as the clade stemming from the last common ancestor of Proterosuchidae and Archosauria. Phil Senter (2005) defined it as the most exclusive clade containing Proterosuchus and Archosauria. Gauthier as part of the Phylonyms (2020) defined the clade as the last common ancestor and all descendants of Gallus, Alligator, and Proterosuchus. Archosauriforms are a branch of archosauromorphs which originated in the Late Permian and persist to the present day as the two surviving archosaur groups: crocodilians and birds.
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.
Euparkeria is an extinct genus of archosauriform reptile from the Triassic of South Africa. Euparkeria is close to the ancestry of Archosauria, the reptile group that includes crocodilians, pterosaurs, and dinosaurs.
Gracilisuchus is an extinct genus of tiny pseudosuchian from the Late Triassic of Argentina. It contains a single species, G. stipanicicorum, which is placed in the clade Suchia, close to the ancestry of crocodylomorphs. Both the genus and the species were first described by Alfred Romer in 1972.
Tasmaniosaurus is an extinct genus of archosauromorph reptile known from the Knocklofty Formation of West Hobart, Tasmania, Australia. The type species is T. triassicus. This genus is notable not only due to being one of the most complete Australian Triassic reptiles known, but also due to being a very close relative of Archosauriformes. Once believed to be a proterosuchid, this taxon is now believed to have been intermediate between advanced non-archosauriform archosauromorphs such as Prolacerta, and basal archosauriforms such as Proterosuchus. Features traditionally used to define Archosauria and later Archosauriformes, such as the presence of an antorbital fenestra and serrated teeth, are now known to have evolved prior to those groups due to their presence in Tasmaniosaurus.
Proterochampsidae is a family of proterochampsian archosauriforms. Proterochampsids may have filled an ecological niche similar to modern crocodiles, and had a general crocodile-like appearance. They lived in what is now South America in the Middle and Late Triassic.
Mesosuchus is an extinct genus of basal rhynchosaur from early Middle Triassic deposits of Eastern Cape, South Africa. It is known from the holotype SAM 5882, a partial skeleton, and from the paratypes SAM 6046, SAM 6536, SAM 7416 and SAM 7701 from the Aliwal North Euparkeria site. Mesosuchus is quite small, spanning around 30 cm in length. Mesosuchus was discovered and named by D. M. S. Watson in 1912.
Proterochampsa is an extinct genus of proterochampsid archosauriform from the Late Triassic of South America. The genus is the namesake of the family Proterochampsidae, and the broader clade Proterochampsia. Like other proterochampsids, Proterochampsa are quadruped tetrapods superficially similar in appearance to modern crocodiles, although the two groups are not closely related. Proterochampsids can be distinguished from other related archosauriformes by characters such as a dorsoventrally flattened, triangular skull with a long, narrow snout at the anterior end and that expands transversally at the posterior end, asymmetric feet, and a lack of postfrontal bones in the skull, with the nares located near the midline. Proterochampsa is additionally defined by characters of dermal sculpturing consisting of nodular protuberances on the skull, antorbital fenestrae facing dorsally, and a restricted antorbital fossa on the maxilla. The genus comprises two known species: Proterochampsa barrionuevoi and Proterochampsa nodosa. P. barrionuevoi specimens have been discovered in the Ischigualasto Formation in northwestern Argentina, while P. nodosa specimens have been found in the Santa Maria supersequence in southeastern Brazil. The two species are distinct in several characters, including that P. nodosa has larger, more well-developed nodular protuberances, a more gradually narrowing snout, and a higher occiput than P. barrionuevoi. Of the two, P. nodosa is thought to have less derived features than P. barrionuevoi.
Turfanosuchus is a genus of archosauriform reptile, likely a gracilisuchid archosaur, which lived during the Middle Triassic (Anisian) of northwestern China. The type species, T. dabanensis, was described by C.C. Young in 1973, based on a partially complete but disarticulated fossil skeleton found in the Kelamayi Formation of the Turfan Basin.
Euparkeriidae is an extinct family of small carnivorous archosauriforms which lived from the Early Triassic to the Middle Triassic (Anisian). While most other early archosauriforms walked on four limbs, euparkeriids were probably facultative bipeds that had the ability to walk on their hind limbs at times. The most well known member of Euparkeriidae is the species Euparkeria capensis, which was named by paleontologist Robert Broom from the Karoo Basin of South Africa in 1913 and is known from several nearly complete skeletons. The family name was first proposed by German paleontologist Friedrich von Huene in 1920; Huene classified euparkeriids as members of Pseudosuchia, a traditional name for crocodilian-line archosaurs from the Triassic. However, phylogenetic analyses performed in the 21st century place Euparkeriidae as a group of Archosauriformes, a position outside Pseudosuchia and close to the ancestry of both crocodile-line archosaurs and bird-line archosaurs. However, they are probably not direct ancestors of archosaurs.
Vancleavea is a genus of extinct, armoured, non-archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is V. campi, named by Robert Long & Phillip A Murry in 1995. At that time, the genus was only known from fragmentary bones including osteoderms and vertebrae. However, since then many more fossils have been found, including a pair of nearly complete skeletons discovered in 2002. These finds have shown that members of the genus were bizarre semiaquatic reptiles. Vancleavea individuals had short snouts with large, fang-like teeth, and long bodies with small limbs. They were completely covered with bony plates known as osteoderms, which came in several different varieties distributed around the body. Phylogenetic analyses by professional paleontologists have shown that Vancleavea was an archosauriform, part of the lineage of reptiles that would lead to archosaurs such as dinosaurs and crocodilians. Vancleavea lacks certain traits which are present in most other archosauriforms, most notably the antorbital, mandibular and supratemporal fenestrae, which are weight-saving holes in the skulls of other taxa. However, other features clearly support its archosauriform identity, including a lack of intercentra, the presence of osteoderms, an ossified laterosphenoid, and several adaptations of the femur and ankle bones. In 2016, a new genus of archosauriform, Litorosuchus, was described. This genus resembled both Vancleavea and more typical archosauriforms in different respects, allowing Litorosuchus to act as a transitional fossil linking Vancleavea to less aberrant archosauriforms.
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.
Rhadinosuchus is an extinct genus of proterochampsian archosauriform reptile from the Late Triassic. It is known only from the type species Rhadinosuchus gracilis, reposited in Munich, Germany. The fossil includes an incomplete skull and fragments of post-cranial material. Hosffstetter (1955), Kuhn (1966), Reig (1970) and Bonaparte (1971) hypothesized it to be synonymous with Cerritosaurus, but other characteristics suggest it is closer to Chanaresuchus and Gualosuchus, while it is certainly different from Proterochampsa and Barberenachampsa. The small size indicates it is a young animal, making it hard to classify.
Youngosuchus is an extinct genus of archosaur from the Middle Triassic of China. The type species is Y. sinensis. Y. sinensis was first described in 1973 as a new species of the erythrosuchid Vjushkovia. In 1985, it was reassigned as its own genus of rauisuchid. A 1992 study supported the original classification of Youngosuchus sinensis as an erythrosuchid, but more recent studies classify it as a "rauisuchian"-grade loricatan archosaur completely unrelated to Vjushkovia, which is most likely a synonym of Garjainia.
Proterochampsia is a clade of early archosauriform reptiles from the Triassic period. It includes the Proterochampsidae and probably also the Doswelliidae. Nesbitt (2011) defines Proterochampsia as a stem-based taxon that includes Proterochampsa barrionuevoi and all forms more closely related to it than Euparkeria capensis, Erythrosuchus africanus, Passer domesticus, or Crocodylus niloticus. Therefore, the inclusion of Doswelliidae in it is dependent upon whether Doswellia and Proterochampsa form a monophyletic group to the exclusion of Archosauria and other related groups.
Jesairosaurus is an extinct genus of early archosauromorph reptile known from the Illizi Province of Algeria. It is known from a single species, Jesairosaurus lehmani. Although a potential relative of the long-necked tanystropheids, this lightly-built reptile could instead be characterized by its relatively short neck as well as various skull features.
Asperoris is an extinct genus of archosauriform reptile known from the Middle Triassic Manda Beds of southwestern Tanzania. It is the first archosauriform known from the Manda Beds that is not an archosaur. However, its relationships with other non-archosaurian archosauriforms are uncertain. It was first named by Sterling J. Nesbitt, Richard J. Butler and David J. Gower in 2013 and the type species is Asperoris mnyama. Asperoris means "rough face" in Latin, referring to the distinctive rough texture of its skull bones.
Litorosuchus is a genus of armored, semiaquatic archosauriform reptile from the Middle Triassic of China, closely related to the morphologically similar Vancleavea. It contains one species, L. somnii.
Rugarhynchos is an extinct genus of doswelliid archosauriform from the Late Triassic of New Mexico. The only known species is Rugarhynchos sixmilensis. It was originally described as a species of Doswellia in 2012, before receiving its own genus in 2020. Rugarhynchos was a close relative of Doswellia and shared several features with it, such as the absence of an infratemporal fenestra and heavily textured skull bones. However, it could also be distinguished by many unique characteristics, such as a thick diagonal ridge on the side of the snout, blunt spikes on its osteoderms, and a complex suture between the quadratojugal, squamosal, and jugal. Non-metric multidimensional scaling and tooth morphology suggest that Rugarhynchos had a general skull anatomy convergent with some crocodyliforms, spinosaurids, and phytosaurs. However, its snout was somewhat less elongated than those other reptiles.
Polymorphodon is an extinct genus of archosauriform reptile from the Middle Triassic of Germany. The only known species is Polymorphodon adorfi, discovered in Lower Keuper deposits at a quarry in Eschenau, Germany. Polymorphodon is notable for its heterodont dentition, with long and conical premaxillary teeth followed by thin maxillary teeth with large serrations. Maxillary teeth near the back of the mouth are short and leaf-shaped, similar to some living and extinct reptiles with a herbivorous or omnivorous diet. This may suggest that Polymorphodon had some reliance on plants in its diet, a rarity among basal archosauriforms, most of which are carnivores.