Proterosuchids | |
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Restored skull of Archosaurus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Archosauriformes |
Family: | † Proterosuchidae Huene, 1914 |
Genera | |
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Synonyms | |
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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. [1] The name comes from Greek πρότερο- ("first") and σοῦχος ("crocodile").
They were slender, medium-sized (about 1.5 m (4 ft 11 in) long, largest specimens reached 3.5–4 m (11–13 ft) [2] ), long-snouted and superficially crocodile-like animals, although they lacked the armoured scutes of true crocodiles, and their skeletal features are much more primitive. The limbs are short and indicate a sprawling posture, like contemporary lizards but unlike most later archosaurs.
Their most characteristic feature is a distinct down-turning of the premaxilla (the front of the upper jaw, which overhangs the lower jaw).
The terminal Permian catastrophe, which killed off 95% of all types of life, cleared the world of all large therapsids and allowed the proterosuchids to become the top predators. Within the space of five million years the proterosuchids had evolved to fill a wide variety of terrestrial and semi-aquatic niches. [3] The proterosuchids represent perhaps the earliest adaptive radiation of the archosaurs. They gave rise to the Erythrosuchidae some time in the Early Triassic.[ citation needed ]
A 2023 study recovered a majority of proterosuchids outside of Proterosuchus and Sarmatosuchus as belonging to a distinct subfamily, the Chasmatosuchinae, defined as "the most inclusive clade containing Chasmatosuchus rossicus , but not Proterosuchus fergusi , " Chasmatosaurus " yuani, Proterosuchus alexanderi , Proterosuchus goweri , Erythrosuchus africanus , or Alligator mississippiensis (the American alligator)". Chasmatosuchinae contains many Triassic proterosuchids in addition to the only Permian proterosuchid, Archosaurus ; this would suggest that an initial taxonomic diversification of the Proterosuchidae (including an initial taxonomic divergence between Proterosuchus and chasmatosuchines) had already occurred in the Permian. [1]
Recent studies consider Proterosuchidae to be at least a partially paraphyletic grouping, meaning that it does not form a true clade with a single common ancestor and proterosuchids as its only descendants. Instead, they are a chain of successively basal archosauriforms. [4] [5] Proterosuchidae sensu stricto is defined as "all taxa more closely related to Proterosuchus fergusi than to Erythrosuchus africanus , Crocodylus niloticus (the Nile crocodile), or Passer domesticus (the house sparrow). [5] 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 are 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 Proterosuchidae containing only Archosaurus and the species of Proterosuchus , however some species (e.g. Chasmatosuchus spp, Vonhuenia friedrichi ) are too fragmentary to resolve whether they also fall into Proterosuchidae. Tasmaniosaurus , Fugusuchus , Sarmatosuchus , Cuyosuchus and the "Long Reef proterosuchid" (SAM P41754) on the other hand were recovered confidently outside of Proterosuchidae. [5]
| previously assigned to Proterosuchidae |
The following phylogeny for the Proterosuchidae sensu stricto was recovered by Ezcurra et al (2023): [1]
Proterosuchidae |
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Genus [5] | Status | Age | Location | Description | Images |
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nomen dubium | Early Triassic | India | |||
Valid | Late Permian, Changhsingian | Russia | |||
nomen dubium | Early Triassic, Induan | Russia | |||
Valid | Early Triassic | Russia | |||
Gamosaurus | Valid [1] | Early Triassic | Russia | ||
Jaikosuchus | Valid [1] | Early Triassic | Russia | ||
Koilamasuchus | Disputed | Middle - Late Triassic | Argentina | ||
Valid | Early Triassic | South Africa | |||
Samsarasuchus | Valid | Early Triassic | India | ||
Sarmatosuchus | Disputed | Early Triassic | Russia | ||
Tsylmosuchus | Valid [1] | Early Triassic | Russia | ||
Valid [1] | Early Triassic, Induan | Russia | |||
Archosauria or archosaurs is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistic sense of the term includes all living and extinct relatives of birds and crocodilians such as non-avian dinosaurs, pterosaurs, phytosaurs, aetosaurs and rauisuchians as well as many Mesozoic marine reptiles. 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.
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.
Ankistrodon is an extinct genus of archosauriform known from the Early Triassic Panchet Formation of India. First thought to be a theropod dinosaur, it was later determined to be a proterosuchid. The type species is A. indicus, described by prolific British zoologist Thomas Henry Huxley in 1865. One authority in the 1970s classified Ankistrodon as a senior synonym of Proterosuchus. Ezcurra (2023) found Ankistrodon to be a nomen dubium, as the teeth are indistinguishable from those of Proterosuchus. A second Indian proterosuchid from the same formation, Samsarasuchus, was also described in the same study, making it the only known valid proterosuchid from India.
Chasmatosuchus was an archosauriform reptile from the early Triassic period of European Russia. One of the earliest described archosauriforms, it was over 2 m long and is thought to have behaved like a modern crocodile. Its mouth had two distinct features: the top of its jaw hooked downwards to aid in holding prey, and the upper palate was lined with a row of teeth—a primitive feature lost in later archosaurs.
Erythrosuchidae are a family of large basal archosauriform carnivores that lived from the later Early Triassic (Olenekian) to the early Middle Triassic (Anisian).
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.
Kalisuchus was a genus of basal archosauriform known from remains unearthed from the Arcadia Formation of the Early Triassic of the Crater, Southwest of Rolleston, south central Queensland, Australia. It was named after Kali, the Hindu goddess of destruction, a reference to the very fragmentary nature of its remains. The type species of Kalisuchus is K. rewanensis, which refers to the Rewan Group. The Arcadia formation is dated to the Induan age at the very beginning of the Triassic, making Kalisuchus one of the oldest archosauromorphs known in Australia.
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.
Tsylmosuchus is an extinct genus of proterosuchid archosauriform reptile known from Western Russia. Fossils referred to Tsylmosuchus occurred over a wide area in sediments corresponding to the Induan and Olenekian stages of the Early Triassic. Most of these fossils are fragmentary neck vertebrae which were originally reported as sharing similarities with crocodile-line archosaurs (pseudosuchians) such as Mandasuchus. As a result, Tsylmosuchus was first described as part of the family Rauisuchidae, making it supposedly one of the oldest known archosaurs. However, its fragmentary remains do not show any of the distinguishing features of rauisuchids or even pseudosuchians in general, so Tsylmosuchus has more recently been interpreted as an indeterminate archosauriform. Although three species of Tsylmosuchus have been named, they lack diagnostic traits and are probably not distinct from each other. In 2023, Tsylmosuchus was reinterpreted as a proterosuchid, specifically a member of the subfamily Chasmatosuchinae.
Vjushkovisaurus is an extinct genus of Middle Triassic archosauriform. It is known from the Anisian-aged Donguz Gorizont in Sol-Iletsk, Orenburg Oblast, Russia. The genus was named in 1982, with the type species being V. berdjanensis. Material has been collected in the Berdyanka II locality from a fossil assemblage called the Eryosuchus Fauna along the Berdyanka River, specifically in a sand-carbonate concretion in the upper part of the main river channel. Vjushkovisaurus is known only from the holotype PIN 2865/62, a partial postcranial skeleton which consists of 12 presacral vertebrae, left humerus, ribs, a fragment of the coracoid and a fragment of the fibula.
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.
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.
Vonhuenia is an extinct genus of proterosuchid, a basal archosauriform from the Early Triassic of Russia. Fossils have been found in the Vokhminskaya Formation, along the Vetluga River that are Induan in age, making Vonhuenia one of the earliest archosauriforms.
Gamosaurus is an extinct genus of proterosuchid archosauriform. It contains a single species, Gamosaurus lozovskii, named by Vitalii Georgievich Ochev in 1979.
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.
Aenigmastropheus is an extinct genus of early archosauromorph reptiles known from the middle Late Permian Usili Formation of Songea District, southern Tanzania. It contains a single species, Aenigmastropheus parringtoni, known solely from UMZC T836, a partial postcranial skeleton of a mature individual. It was collected in 1933, and first described in 1956, as a "problematic reptile" due to its unique morphology. Therefore, a binomial name was erected for this specimen in 2014. Aenigmastropheus was probably fully terrestrial.
Samsarasuchus is an extinct genus of archosauriform reptile from the Early Triassic of India. This genus has one known species, Samsarasuchus pamelae. Samsarasuchus lived a few million years after the Permian-Triassic extinction, the largest known mass extinction event. It was a member of the Proterosuchidae, a group of successful crocodile-like reptiles that survived the extinction event and were among the earliest successful archosauromorphs.
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