| Erythrosuchids Temporal range: Early – Middle Triassic, | |
|---|---|
 | |
| Skull of Erythrosuchus . | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Archosauromorpha |
| Clade: | Archosauriformes |
| Family: | † Erythrosuchidae Watson, 1917 |
| Genera | |
see below | |
Erythrosuchidae (meaning "red crocodiles" in Greek) are a family of large basal archosauriform carnivores that lived from the later Early Triassic (Olenekian) to the early Middle Triassic (Anisian).
The family Erythrosuchidae was named by David Meredith Seares Watson in 1917. [1]
They were the apex predators of their day, with lengths of 2.5 m (8 ft 2 in) to almost 5 m (16 ft). [2] Their fossil remains are known to date from South Africa (Beaufort Group of the Karoo Basin), China, India and European Russia, from the Early to Middle Triassic.
Erythrosuchids were unusually large and robust archosauromorphs. Several features set them apart from other archosauriformes and are also seen in later, more derived archosaurs. For example, they lack teeth on the palate, which are found in other early archosauriformes, such as Doswellia and euparkeriids. In erythrosuchids, the centra (central parts of vertebrae) are deeply indented on either side, differing considerably from the usual cylindrical shape of the centra in early archosauriformes, but similar to later archosaurs. [3]
The heads of erythrosuchids are generally disproportionately large and deep. In all erythrosuchids, the lower margin of the premaxilla, the bone at the tip of the upper jaw, is lower than the lower margin of the maxilla, the bone behind the premaxilla. [2] This forms a characteristic "step" that makes erythrosuchids easily distinguishable from all other early archosauriformes, which have smooth jaw margins that are either straight or gradually curved. [3]
Erythrosuchids are notable for being the first archosauriforms to have a triradiate pelvic girdle with three projecting areas formed from three bones: an ilium and an elongated pubis and ischium. [4] Although it is small, the fourth trochanter, a ridge on the femur that serves as a muscle attachment in archosaurs, first appears in erythrosuchids. The triradiate pelvis and fourth trochanter are both features which indicate that erythrosuchids had an erect stance similar to later archosaurs. More basal archosauriforms such as proterosuchids lacked these features and probably had a more sprawling posture. [5]
Erythrosuchids were formerly classified as thecodonts of the suborder Proterosuchia. This classification is no longer used by paleontologists, who now employ a cladistic approach. In this, erythrosuchids constitute an Archosauriformes clade that is an outgroup to the Archosauria proper. The presence of certain archosaurian features, such as the triradiate pelvic girdle, the fourth trochanter, and the third metatarsal longer than the fourth, indicate that erythrosuchids are closer to the true archosaurs than the Proterosuchidae, which lack these features. Thus the Erythrosuchidae occupy a transitional evolutionary position between the most primitive archisauriformes and more advanced Triassic archosaurs.
| Genus [6] | Status | Age | Location | Description | Images |
|---|---|---|---|---|---|
| Valid | Middle Triassic | India | |||
| Valid | Middle Triassic | European Russia |  | ||
| Valid | Early -Middle-Triassic | South Africa |  | ||
| Valid | Early Triassic | European Russia and South Africa |  | ||
| Valid | Early Triassic | China | |||
| Valid | Middle Triassic | China |  | ||
| Valid | Middle Triassic | European Russia | |||
| Valid | Middle Triassic | European Russia | |||
Cladogram from Parrish (1992): [3]
| Archosauriformes |
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Below is a cladogram from Ezcurra (2016) that reexamined all historical members of the "Proterosuchia" (a polyphyletic historical group including proterosuchids and erythrosuchids). The placement of fragmentary taxa that had to be removed to increase tree resolution is indicated by dashed lines (in the most derived position that they can be confidently assigned to). Taxa that are nomina dubia are indicated by the note "dubium". Bold terminal taxa are collapsed. Ezcurra (2016) recovered a monophyletic Erythrosuchidae, although the relationship of the Long Reef form (SAM P41754) and Uralosaurus remain unknown within the family. [6]
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The large contemporary kannemeyeriid dicynodonts doubtless constituted much of erythrosuchids' prey. However, the first erythrosuchids appear in the fossil record slightly earlier than the kannemeyeriids, so it must be assumed that they fed on other animals as well.
Archosauria is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term, which includes birds. Extinct archosaurs include non-avian dinosaurs, pterosaurs and extinct relatives of crocodilians. Modern paleontologists define Archosauria as a crown group that includes the most recent common ancestor of living birds and crocodilians, and all of its descendants. The base of Archosauria splits into two clades: Pseudosuchia, which includes crocodilians and their extinct relatives; and Avemetatarsalia, which includes birds and their extinct relatives.
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. 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.
The fourth trochanter is a shared characteristic common to archosaurs. It is a knob-like feature on the posterior-medial side of the middle of the femur shaft that serves as a muscle attachment, mainly for the musculus caudofemoralis longus, the main retractor tail muscle that pulls the thighbone to the rear.
Proterosuchidae is an early family of basal archosauriforms whose fossils are known from the Late Permian and the Early Triassic. The highest diversity of genera is known from European Russia, but fossils are also known from South Africa, India, China, Australia, Brazil and possibly Argentina. The name comes from Greek πρότερο- ("first") and σοῦχος ("crocodile").
Shansisuchus is an extinct genus of archosauriform reptile belonging to the family Erythrosuchidae that lived during the Middle Triassic in what is now China. The first fossils of Shansisuchus were discovered from the Ermaying Formation of Shanxi (Shansi) province in 1964 by Chinese paleontologist Yang Zhongjian. Like other erythrosuchids, Shansisuchus was a large-bodied carnivore with a large, deep skull. Shansisuchus is unique among early archosauriforms in having a hole in its skull called a subnarial fenestra.
Garjainia is an extinct genus of erythrosuchid archosauriform reptile from the Olenekian of Russia and South Africa. It was approximately 1.5–2 metres (4.9–6.6 ft) long. It contained two species, Garjainia prima from the Yarengian/Yarkenskian Supergorizont of Russia, and Garjainia madiba from the Burgersdorp Formation of South Africa. "Vjuskovia triplicostata", a name assigned to some erythrosuchid fossils from Russia, has been synonymized with Garjainia prima.
Proterosuchus is an extinct genus of archosauriform reptiles that lived during the Early Triassic. It contains three valid species: the type species P. fergusi and the referred species P. alexanderi and P. goweri. All three species lived in what is now South Africa. The genus was named in 1903 by the South African paleontologist Robert Broom. The genus Chasmatosaurus is a junior synonym of Proterosuchus.
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.
Archosaurus is an extinct genus of carnivorous proterosuchid archosauriform reptile. Its fossils are dated to the latest Permian of Russia and Poland, it is one of the earliest known archosauriforms. The type and only species is Archosaurus rossicus, known from several fragmentary specimens which cumulatively represent parts of the skull and cervical vertebrae. It would have been 3 metres (9.8 ft) long when fully grown.
Yarasuchus is an extinct genus of avemetatarsalian archosaur that lived during the Anisian stage of the Middle Triassic of India. The genus was named and described in 2005 from a collection of disarticulated but fairly complete fossil material found from the Middle Triassic Yerrapalli Formation. The material is thought to be from two individuals, possibly three, with one being much more complete and articulated than the other. The type and only species is Y. deccanensis. Yarasuchus was a quadruped roughly 2–2.5 metres (6.6–8.2 ft) long, with an elongated neck and tall spines on its vertebrae. Unlike other quadrupedal Triassic reptiles, the limbs and shoulders of Yarasuchus were slender, and more like those of ornithodirans.
Fenhosuchus is an extinct genus of archosauriform. The holotype, IVPP V 2697, and referred materials have been found in the Hsishihwa locality at Wuhsiang, China, from the Upper Ermaying Formation. The locality dates back to the Anisian stage of the Middle Triassic. The genus was named after the Fen River in Shanxi Province from which specimens were found. It may prove to be a chimera being composed of material from several different animals. Some material were believed to represent a rauisuchid. The calcaneum of Fenhosuchus seems to belong to an erythrosuchid or other basal archosauriform. Much of the material of the tarsal bones seem to be similar to those of the genus Shansisuchus. According to Nesbitt (2009) the assessment of Gower (2000) was correct, the holotype is a mix of Shansisuchus remains and a possible fragment from a paracrocodylomorph or a dinosauriform. Thus, Fenhosuchus cannot be considered a rauisuchian.
Koilamasuchus is an extinct genus of indeterminate archosauriform from the Triassic of Argentina. It is based on an external mold of a partial postcranial skeleton from the Quebrada de los Fósiles Formation. Due to its incomplete nature, the relationships of this reptile are difficult to establish. Originally described as a non-archosaur archosauriform, later studies tentatively considered it a doswelliid or a suchian archosaur.
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 and all forms more closely related to it than Euparkeria, Erythrosuchus, 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.
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.
Eorasaurus is an extinct genus of archosauromorph reptile known from the middle Late Permian of Tatarstan, European Russia. It contains a single species, Eorasaurus olsoni. When originally described by Sennikov (1997), Eorasaurus was identified as an early archosauromorph and assigned to the family Protorosauridae, Ezcurra et al. (2014) and Ezcurra (2016) later reclassified Eorasaurus and placed it within the group Archosauriformes. Eorasaurus is based solely on scant fossil material from the neck region, and is thus considered an unstable taxon in phylogenetic analyses. If Eorasaurus is an archosauriform, it would be the oldest known member of the group and would pre-date the previous record holder.
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.
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.
{{cite book}}: CS1 maint: multiple names: authors list (link)ISBN 0-231-13522-X
