Archosauriforms | |
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Row 1 (basal archosauriforms): Erythrosuchus africanus , Euparkeria capensis ; Row 2 (Pseudosuchia): Crocodylus mindorensis , Typothorax coccinarum ; Contents | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | Crocopoda |
Clade: | Archosauriformes Gauthier, 1986 |
Subgroups [1] | |
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Archosauriformes (Greek for 'ruling lizards', and Latin for 'form') 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. [3] Phil Senter (2005) defined it as the most exclusive clade containing Proterosuchus and Archosauria. [4] Gauthier as part of the Phylonyms (2020) defined the clade as the last common ancestor and all descendants of Gallus , Alligator , and Proterosuchus . [5] Archosauriforms are a branch of archosauromorphs which originated in the Late Permian (roughly 252 million years ago) and persist to the present day as the two surviving archosaur groups: crocodilians and birds.
Archosauriforms present several traits historically ascribed to the group Archosauria. These include serrated teeth set in deep sockets, a more active metabolism, and an antorbital fenestra (a hole in the skull in front of the eyes). Reptiles with these traits have also been termed "thecodonts" in older methods of classification. Thecodontia is a paraphyletic group, and its usage as a taxonomic category has been rejected under modern cladistic systems. The name Archosauriformes is intended as a monophyletic replacement compatible with modern taxonomy.
Early archosauriforms, informally termed "proterosuchians", were superficially crocodile-like animals with sprawling gaits, carnivorous habits, and long hooked snouts. Unlike the bulk of their therapsid contemporaries, archosauriforms survived the catastrophic end-Permian mass extinction. The Late Permian proterosuchid Archosaurus is similar in appearance to its Early Triassic relative, Proterosuchus. Within a few million years after the beginning of the Triassic, the archosauriformes had diversified past the "proterosuchian" grade. The next major archosauriform group was Erythrosuchidae, a family of apex predators with massive heads, the largest carnivorous reptiles up to that time.
In 2016, Martin Ezcurra provided the name Eucrocopoda for the clade including all archosauriforms more crownward (closer to archosaurs) than erythrosuchids. He defined the clade all taxa more closely related to Euparkeria capensis , Proterochampa barrionuevoi , Doswellia kaltenbachi , Parasuchus hislopi , Passer domesticus (the house sparrow), or Crocodylus niloticus (the Nile crocodile) than to Proterosuchus fergusi or Erythrosuchus africanus . The name translates to "true crocodile feet", in reference to the possession of a crocodilian-style crurotarsal ankle. [1] Eucrocopodans include the families Euparkeriidae (small, agile reptiles),Proterochampsidae (narrow-snouted predators endemic to South America), and Doswelliidae (heavily armored Laurasian reptiles similar to proterochampsids), as well as various other strange reptiles such as Vancleavea and Asperoris .
The most successful archosauriforms, and the only members to survive into the Jurassic, were the archosaurs. Archosauria includes crocodilians, birds, and all descendants of their common ancestor. Extinct archosaurs include aetosaurs, rauisuchids (both members of the crocodilian branch), pterosaurs, and non-avian dinosaurs (both members of the avian branch). [6]
Vascular density and osteocyte density, shape and area have been used to estimate the bone growth rate of archosaurs, leading to the conclusion that this rate had a tendency to grow in ornithodirans and decrease in pseudosuchians. [7] The same method also supports the existence of high resting metabolical rates similar to those of living endotherms (mammals and birds) in the Prolacerta -Archosauriformes clade that were retained by most subgroups, though decreased in Proterosuchus, Phytosauria and Crocodilia. [8] Erythrosuchids and Euparkeria are basal archosauriforms showing signs of high growth rates and elevated metabolism, with Erythrosuchus possessing a rate similar of the fastest-growing dinosaurs. Sexual maturity in those Triassic taxa was probably reached quickly, providing advantage in a habitat with unpredictable variation from heavy rainfall to drought and high mortality. Vancleavea and Euparkeria, which show slower growth rates compared to Erythrosuchus, lived after the climatic stabilization. Early crown archosaurs possessed increased growth rates, which were retained by ornithodirans. [9] Ornithosuchians and poposaurs are stem-crocodilians that show high growth rates similar to those of basal archosauriforms. [10]
Developmental, physiological, anatomical and palaeontological lines of evidence indicate that crocodilians evolved from endothermic ancestors. Living crocodilians are ambush predators adapted to a semi-aquatic lifestyle that benefits from ectothermy due to the lower oxygen intake that allows longer diving time. The mixing of oxygenated and deoxygenated blood in their circulatory system is apparently an innovation that benefits ectothermic life. Earlier archosaurs likely lacked those adaptations and instead had completely separated blood as birds and mammals do. [11] [12] A similar process occurred in phytosaurs, which were also semi-aquatic. [13]
The similarities between pterosaur, ornithischian and coelurosaurian integument suggest a common origin of thermal insulation (feathers) in ornithodirans at least 250 million years ago. [14] [15] Erythrosuchids living in high latitudes might have benefited from some sort of insulation. [13] If Longisquama was an archosauromorph, it could be associated with the origin of feathers. [16] [13]
Below is a cladogram from Nesbitt (2011): [17]
Archosauriformes | *Note: Phytosaurs were previously placed within Pseudosuchia, or crocodile-line archosaurs. |
Below is a cladogram from Sengupta et al. (2017), [18] based on an updated version of Ezcurra (2016) [1] 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. [1]
Crocopoda |
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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.
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.
Erythrosuchidae are a family of large basal archosauriform carnivores that lived from the later Early Triassic (Olenekian) to the early Middle Triassic (Anisian).
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").
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.
Dongusuchus is an extinct genus of archosaur. Fossils have been found from the Donguz Formation outcropping on the banks of the Donguz River in the Orenburg Oblast of Russia. They are associated with a fossil assemblage called the Eryosuchus Fauna, named after the capitosaurid Eryosuchus, the most common organism found from the assemblage. The locality dates back to the Anisian and early Ladinian stages of the Middle Triassic.
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.
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.
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.
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.
Prolacerta is a genus of archosauromorph from the lower Triassic of South Africa and Antarctica. The only known species is Prolacerta broomi. Prolacerta was a small and slender reptile, with a rather long neck, low skull, and serrated teeth. It would have resembled a modern monitor lizard at a quick glance, though this is an example of convergent evolution as opposed to close affinities.
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.
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.