Diandongosuchus Temporal range: Middle Triassic | |
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Skull in Beijing Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Order: | † Phytosauria |
Genus: | † Diandongosuchus Li et al., 2012 |
Type species | |
†Diandongosuchus fuyuanensis Li et al., 2012 |
Diandongosuchus is an extinct genus of archosauriform reptile, possibly a member of the Phytosauria, known from the Middle Triassic of China. The type species Diandongosuchus fuyuanensis was named in 2012 from the Zhuganpo Formation of Yunnan Province. It is a marine species that shows similarities with another Chinese Triassic species called Qianosuchus mixtus , although it has fewer adaptations toward marine life. It was originally classified as the basal-most member of the pseudosuchian clade Poposauroidea. [1] However, a subsequent study conducted by Stocker et al. (2016, 2017) indicated it to be the basalmost known phytosaur instead. [2] [3]
Diandongosuchus is known from a nearly complete articulated skeleton (ZMNH M8770) missing most of the tail. The total length of ZMNH M8770 is 97 centimetres (3.18 ft) and the estimated body length of the animal in life is around 155 centimetres (5.09 ft). The specimen is preserved on its right side, with the underside of the lower jaws and the trunk showing. It was prepared out of a limestone slab to reveal details on the left side of the skeleton, many of which are better preserved. The skull of Diandongosuchus is pointed, with oval-shaped eye sockets, antorbital and temporal openings. Distinctive features include a long premaxilla bone at the tip of the snout that extends backward past the nostril openings, a large ridge on the jugal bone that runs beneath the eye socket, and two supratemporal openings on the skull table that have prominent ridges surrounding them. The skull has similar proportions to that of Qianosuchus, and has the same number of teeth in the premaxilla. Like the terrestrial poposauroid Poposaurus, Diandongosuchus has a maxilla (upper jaw) bone that does not reach the border of the nostril opening. [1]
ZMNH M8770 has 25 vertebrae in the back and neck, two sacral vertebrae (as in most Triassic pseudosuchians), and seven of the forward-most tail vertebrae. The neck vertebrae are taller and narrower than they are in Qianosuchus. Most of the back vertebrae are obscured by overlying ribs. At the back of the trunk near the hips are bones belonging to small vertebrates such as fish - likely the stomach contents of the individual. Small overlapping osteoderms (bony scutes) overlay many of the vertebrae. Two rows run along the neck, back, and tail with about two osteoderms overlaying each vertebra. Small osteoderms also cover the limb bones. [1]
Some features of the limbs, pelvic and pectoral girdles are also diagnostic in Diandongosuchus, including a thick ischium bone in the hip, an opening of the coracoid bone in the pectoral girdle that is much larger than those of other archosaurs and is closed by the end of the scapula, and a fourth metatarsal bone in the foot that is longer than the other metatarsals. The scapula of Diandongosuchus is longer and narrower than that of Qianosuchus. The iliac blade of the hip is unusual in that it is narrow and projects far back from the rest of the hip. As in Qianosuchus, the femur of Diandongosuchus is slightly twisted, but the fibula is thinner and more curved. The astragalus and calcaneum bones of the ankle fit together like a ball-and-socket, a feature that confirms Diandongosuchus as a pseudosuchian. Some of the phalanges or toe bones are missing in ZMNH M8770, but the metatarsals are present and have unique proportions among Triassic archosaurs in which the fourth is longer than the third. [1]
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A phylogenetic analysis conducted by Li et al. (2012) in the original description of Diandongosuchus showed that it was the most basal member of a clade called Poposauroidea, which includes mostly terrestrial pseudosuchians such as the bipedal Poposaurus and the sail-backed Arizonasaurus . It was found to be closely related to Qianosuchus, an aquatic pseudosuchian that was the second most basal member of Poposauroidea. The data matrix of Li et al., a list of characteristics that was used in the analysis, was based on that of Nesbitt (2011), one of the most extensive on archosaurs. Because of this, many of the relationships found by Li et al. are the same as those found by Nesbitt. Below is a cladogram from the analysis: [1]
Archosauria |
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However, more recent studies have found it to be a basal phytosaur. [2] [3]
Diandongosuchus was found in a Ladinian-age marine limestone formation that has preserved many marine reptiles including thallatosaurs, nothosaurs, pistosaurs, and some protorosaurs. The closely related pseudosuchian Qianosuchus was found in a marine deposit about 50 kilometres (31 mi) northwest of the Diandongosuchus locality that is slightly older (Anisian in age) and possesses many features consistent with a marine lifestyle. However, Diandongosuchus shows no features that are clear adaptations to a marine lifestyle. Possible adaptations include nostrils that are positioned slightly farther back on the skull than most terrestrial pseudosuchians and a greater number of premaxillary teeth (a feature seen in possible semiaquatic archosaurs such as Chanaresuchus and spinosaurids). Fish bones within its stomach contents are additional evidence that it was a marine archosaur. Diandongosuchus may have had a similar lifestyle to modern marine crocodylians like the saltwater crocodile that live along coastlines yet are not fully marine. [1]
The fossil assemblage in which Diandongosuchus was found bears many similarities to that of European fossil localities such as Monte San Giorgio. Both include marine reptiles like thallatosaurs and nothosaurs and probably represented environments along the northern shorelines of the Tethys Ocean. No marine archosaurs like Diandongosuchus and Qianosuchus are known from Europe, although the pseudosuchian Ticinosuchus from Monte San Giorgio was probably adapted to life along the shorelines of the Tethys. In the analyses of Li et al. (2012) and Nesbitt (2011), Ticinosuchus is either the most basal member of a clade called Loricata which is the sister taxon of Poposauroidea, or the sister taxon of Paracrocodylomorpha which includes both Loricata and Poposauroidea. [1] Although Ticinosuchus and Diandongosuchus were initially believed to have been very closely related basal paracrocodylomorphs, this hypothesis is invalidated if Diandongosuchus is a phytosaur as other studies have shown. [3]
"Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians than to birds and other dinosaurs. First named in the 1940s, Rauisuchia was a name exclusive to Triassic archosaurs which were generally large, carnivorous, and quadrupedal with a pillar-erect hip posture, though exceptions exist for all of these traits. Rauisuchians, as a traditional taxonomic group, were considered distinct from other Triassic archosaur groups such as early dinosaurs, phytosaurs, aetosaurs, and crocodylomorphs.
Phytosaurs are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. and are sometimes referred to as parasuchians. Phytosauria, Parasuchia, Parasuchidae, and Phytosauridae have often been considered equivalent groupings containing the same species. Some recent studies have offered a more nuanced approach, defining Parasuchidae and Phytosauridae as nested clades within Phytosauria as a whole. Phytosaurs were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodilians in size, appearance, and lifestyle, as an example of convergence or parallel evolution. The name "phytosaur" means "plant reptile", as the first fossils of phytosaurs were mistakenly thought to belong to plant eaters.
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.
Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Despite Pseudosuchia meaning "false crocodiles", the name is a misnomer as true crocodilians are now defined as a subset of the group.
Poposaurus is an extinct genus of pseudosuchian archosaur from the Late Triassic of the southwestern United States. It belongs to the clade Poposauroidea, an unusual group of Triassic pseudosuchians that includes sail-backed, beaked, and aquatic forms. Fossils have been found in Wyoming, Utah, Arizona, and Texas. Except for the skull, most parts of the skeleton are known. The type species, P. gracilis, was described and named by Maurice Goldsmith Mehl in 1915. A second species, P. langstoni, was originally the type species of the genus Lythrosuchus. Since it was first described, Poposaurus has been variously classified as a dinosaur, a phytosaur, and a "rauisuchian".
Batrachotomus is a genus of prehistoric archosaur. Fossils of this animal have been found in southern Germany and dated from the Ladinian stage of the Middle Triassic period, around 242 to 237 million years ago. Batrachotomus was described by palaeontologist David J. Gower 22 years after its discovery.
Qianosuchus is an extinct genus of aquatic poposauroid archosaur from the middle Triassic (Anisian) Guanling Formation of Pan County, China. It is represented by two nearly complete skeletons and a crushed skull preserved in the limestone. Qianosuchus was at least 3 metres long, and had several skeletal adaptations which indicate a semi-marine lifestyle, similar to modern-day saltwater crocodiles. These adaptations have not been seen in any other archosaur from the Triassic.
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.
Heptasuchus is an extinct genus of loricatan pseudosuchian known from the Middle or Late Triassic upper Chugwater Group of Wyoming, United States. It contains a single species, Heptasuchus clarki, the first formally recognized "rauisuchian" or loricatan pseudosuchian from North America.
Mandasuchus is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956, a formal description was not published until 2018.
Parringtonia is an extinct genus of Triassic archosaur within the family Erpetosuchidae, known from the type species Parringtonia gracilis. It is known from a single specimen, NHMUK R8646, found from the Anisian-age Manda Formation of Tanzania. This specimen, like most archosaur material from the Manda Formation, is fragmentary, including only a maxilla and a few postcranial bones. They show similarities with those of another archosaur called Erpetosuchus, known from the Middle Triassic of Scotland and the eastern United States. The phylogenetic placement of Parringtonia and Erpetosuchus are uncertain; some studies placed them close to the group Crocodylomorpha, which includes all modern crocodylians and many extinct forms that diversified after the Triassic, but this relationship has more recently been questioned.
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.
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.
Loricata is a clade of archosaur reptiles that includes crocodilians and some of their Triassic relatives, such as Postosuchus and Prestosuchus. More specifically, Loricata includes Crocodylomorpha and most "rauisuchians", a paraphyletic grade of large terrestrial pseudosuchians which were alive in the Triassic period and ancestral to crocodylomorphs.
Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as Qianosuchus and Lotosaurus. It excludes most large predatory quadrupedal "rauisuchians" such as rauisuchids and "prestosuchids". Those reptiles are now allied with crocodylomorphs in a clade known as Loricata, which is the sister taxon to the poposauroids in the clade Paracrocodylomorpha. Although it was first formally defined in 2007, the name "Poposauroidea" has been used for many years. The group has been referred to as Poposauridae by some authors, although this name is often used more narrowly to refer to the family that includes Poposaurus and its close relatives.
Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomorpha was named by paleontologist J. Michael Parrish in 1993, although the group is now considered to encompass more reptiles than his original definition intended. The most recent definition of Paracrocodylomorpha, as defined by Sterling Nesbitt in 2011, is "the least inclusive clade containing Poposaurus and Crocodylus niloticus. Most groups of paracrocodylomorphs became extinct at the end of the Triassic period, with the exception of the crocodylomorphs, from which crocodylians such as crocodiles and alligators evolved in the latter part of the Mesozoic.
Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include Erpetosuchus. It includes the type species Erpetosuchus granti from the Late Triassic of Scotland, Erpetosuchus sp. from the Late Triassic of eastern United States and Parringtonia gracilis from the middle Middle Triassic of Tanzania; the group might also include Dyoplax arenaceus from the Late Triassic of Germany, Archeopelta arborensis and Pagosvenator candelariensis from Brazil and Tarjadia ruthae from Argentina.
Nundasuchus is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, Nundasuchus songeaensis, known from a single partially complete skeleton, including vertebrae, limb elements, osteoderms, and skull fragments.
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.
Aphanosauria is an extinct group of reptiles distantly related to dinosaurs. They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchia, includes modern crocodilians. Aphanosaurs possessed features from both groups, indicating that they are the oldest and most primitive known clade of avemetatarsalians, at least in terms of their position on the archosaur family tree. Other avemetatarsalians include the flying pterosaurs, small bipedal lagerpetids, herbivorous silesaurids, and the incredibly diverse dinosaurs, which survive to the present day in the form of birds. Aphanosauria is formally defined as the most inclusive clade containing Teleocrater rhadinus and Yarasuchus deccanensis but not Passer domesticus or Crocodylus niloticus. This group was first recognized during the description of Teleocrater. Although only known by a few genera, Aphanosaurs had a widespread distribution across Pangaea in the Middle Triassic. They were fairly slow quadrupedal long-necked carnivores, a biology more similar to basal archosaurs than to advanced avemetatarsalians such as pterosaurs, lagerpetids, and early dinosaurs. In addition, they seemingly possess 'crocodile-normal' ankles, showing that 'advanced mesotarsal' ankles were not basal to the whole clade of Avemetatarsalia. Nevertheless, they possessed elevated growth rates compared to their contemporaries, indicating that they grew quickly, more like birds than other modern reptiles. Despite superficially resembling lizards, the closest modern relatives of aphanosaurs are birds.