Suchia

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Suchians
Temporal range: Early Triassic - Recent, 248–0  Ma
Paratypothorax andressorum.JPG
Paratypothorax andressorum , an aetosaur
Prestosuchus chiniquensis white background.jpg
Prestosuchus chiniquensis , an early loricatan
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Suchia
Krebs, 1974
Subgroups

Suchia is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing Aetosaurus ferratus , Rauisuchus tiradentes , Prestosuchus chiniquensis , and Crocodylus niloticus (the living Nile crocodile) by Nesbitt (2011). [1] 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 (which are usually considered suchians) and aetosaurs (which are suchians by definition of the group). [2] Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place. [1] [3]

There is some controversy over which traits, if any, can be used to distinguish suchians from non-suchian archosaurs. Anatomical features which evolve at the base of a group, and can thus be used to characterize the group, are known as synapomorphies. The two most comprehensive recent studies on archosaur relationships, Nesbitt (2011) [1] and Ezcurra (2016), [3] each came to different conclusions on what synapomorphies characterize Suchia. Nevertheless, they did agree on one trait that likely qualified as a synapomorphy of Suchia: a calcaneal tuber which was significantly wider than it was tall.

Description

As Suchia is a clade, it is defined by relationships rather than shared characteristics between its members. However, several traits are found in most suchians but not in ornithosuchids, phytosaurs, or other archosaurs and archosaur relatives, so they can be used to help determine whether a reptile is a suchian or not. These traits are spread out throughout the body, but different paleontologists disagree on whether they were true synapomorphies (derived distinguishing features).

While Benton & Clark (1988) found numerous characteristics for the clade later renamed Suchia, Paul Sereno's 1991 study on archosaur relationships argued that many of them were also present in ornithosuchids, phytosaurs, or avemetatarsalians, so they could not be considered traits which diagnose Suchia (under his definition of the group). Instead, Sereno listed a single synapomorphy for Suchia. The postorbital-squamosal bar, which separates the upper and lower temporal fenestrae at the rear part of the skull, is short. This has the effect of making the lower temporal fenestra triangular in shape. However, Sereno also noted that this feature was not present in Prestosuchus, which he excluded from the group. [4] Later studies found support for a placement of Prestosuchus deep within Suchia, so Sereno's synapomorphy is likely invalid. [1]

Nesbitt (2011)'s synapomorphies

The left calcaneum (heel bone) of Batrachotomus, showing several of the calcaneal features Nesbitt (2011) considered to be synapomorphic for Suchia Batrachotomus calcaneum labelled.png
The left calcaneum (heel bone) of Batrachotomus , showing several of the calcaneal features Nesbitt (2011) considered to be synapomorphic for Suchia

Sterling Nesbitt's major 2011 analysis of early archosaurs found several more stable syapomorphies of Suchia. The jugal (cheek bone, below the eye) has a low, rounded longitudinal ridge in suchians, contrasting with other archosaurs which typically have no ridge, or a sharp ridge instead. As is the case for many early archosaurs, the structure of the calcaneum (outer heel bone) is important for diagnosing Suchia. This bone connects to both the fibula (outer shin bone) and astragalus (inner ankle bone) by means of two distinct surfaces, the convex fibular condyle and the concave astragalar facet. In other archosaurs, these surfaces are continuous, rather than separate. Moreover, the fibular condyle is hemicylindrical (half-cylinder shaped) in contrast to the more spherical structure of ornithosuchids, avemetatarsalians, and archosaur relatives such as Euparkeria and proterochampsians. A hemicylindrical fibular condyle is also present in phytosaurs, likely convergently. Lastly, the rear edge of the calcaneum has a backwards-extending tube of bone known as a calcaneal tuber. In early archosauriforms, the calcaneal tuber was taller than wide, while later archosaurs and archosaur relatives had a calcaneal tuber which was circular in cross-section. Suchians take this one step further, as early members of the group had calcaneal tubers which were much wider than tall. Although bipedal poposauroids and most loricatans ( Luperosuchus and crownward) had circular calcaneal tubers, these features were likely reversions, since the earliest poposauroids ( Qianosuchus , for example) and loricatans ( Batrachotomus ) retained wide calcaneal tubers. [1]

Some suchians have a shallow longitudinal groove on the rear edge of the radius (inner forearm bone), although the distribution of this characteristic is poorly understood. Likewise, some early suchians have a triangular medial condyle of the femur (thigh bone), but this characteristic is also known in some archosaur relatives so it is unclear whether the presence of the trait in suchians was a synapomorphy or a plesiomorphy ("primitive" trait retained from an earlier ancestor). In addition, most loricatans reverted to the state present in other archosaurs: a rounded medial condyle. [1] Many of Nesbitt (2011)'s suchian synapomorphies are not present in Nundasuchus , an unusual crurotarsan sometimes considered a suchian close to Ticinosuchus and Paracrocodylomorpha. [5]

Ezcurra (2016)'s synapomorphies

The skull of Gracilisuchus. Note the rounded ridge on the jugal bone beneath the eye (one of Nesbitt [2011]'s synapomorphies) visible in A, and the posterodorsal process of the maxilla (one of Ezcurra [2016]'s synapomorphies) visible in E (pdpm) Gracilisuchus.tif
The skull of Gracilisuchus . Note the rounded ridge on the jugal bone beneath the eye (one of Nesbitt [2011]'s synapomorphies) visible in A, and the posterodorsal process of the maxilla (one of Ezcurra [2016]'s synapomorphies) visible in E (pdpm)

Martin Ezcurra's 2016 analysis of archosauromorphs provided an alternative diagnosis for Suchia. Ezcurra found that Koilamasuchus was a member of the group, but also argued that Suchia excluded Nundasuchus. As a result, his diagnosis for Suchia depended primarily on several traits visible in Koilamasuchus, which was known from significantly less fossil material than most other Suchia. These traits are the presence of large, well-rimmed pits on either side of the dorsal (back) vertebrae, a humerus (upper arm bone) which has a symmetrical proximal portion when seen from the front, and a preacetabular process (front blade) of the ilium (upper hip bone) which has a moderate length, longer than it is high, but not longer than the pubic peduncle. He also noted that in some suchians, the tip of the maxillary bone's posterior process (rear branch), near the rear lower tip of the antorbital fenestra, is actually taller than the middle portion of the rear branch. [3] This trait, formally known as the posterodorsal process of the maxilla, has been considered a synapomorphy of gracilisuchids and is also present in aetosaurs and Qianosuchus. [6] Despite the broad distribution of this characteristic, its absence in some suchians makes it ambiguous whether it qualifies as a synapomorphy of Suchia, or alternatively evolved in several independent lineages within the group. Ezcurra also agreed with one of Nesbitt (2011)'s synapomorphies of the calcaneum, namely the calcaneal tuber being wider than tall. [3]

Classification

Suchia was originally formalized by German paleontologist Bernard Krebs in 1974. His intention was to link the Triassic pseudosuchians (which at that time were considered to have left no descendants) with the true crocodilians that would evolve later in the Mesozoic. Suchia was named as an order united by the possession of a crurotarsal ankle, with Pseudosuchia and Crocodilia as independent suborders within it. [7] However, newer studies starting in the 1980s have significantly altered the traditional classification scheme of crocodilian ancestors. Rather than pseudosuchians and crocodilians being sister groups as Krebs believed, the crocodilians are now considered to be a subset of Pseudosuchia. [8] This necessitated Suchia to acquire a new definition, as Kreb's original classification scheme was invalidated.

Sereno (1991) assigned the name Suchia to a clade containing most pseudosuchians, which had previously been found by Benton & Clark (1988). Under this new definition, Suchia included Gracilisuchus stipanicicorum , Aetosauria, "Rauisuchia", Poposauridae, Crocodylomorpha, and all descendants of their common ancestor. [4] Nesbitt (2011) altered Sereno's definition slightly, as he defined Suchia as the least inclusive clade containing Aetosaurus ferratus , Rauisuchus tiradentes , Prestosuchus chiniquensis , and Crocodylus niloticus (the living Nile crocodile). Suchia, therefore, is a node-based taxon including crocodylomorphs and several more basal Triassic pseudosuchian groups. Such groups include Gracilisuchidae, Revueltosaurus, Aetosauria, Ticinosuchus, Poposauroidea, Loricata, [1] [9] and possibly Erpetosuchidae, [2] Koilamasuchus, [3] Nundasuchus, [5] and/or Euscolosuchus . [10]

Phylogeny

Below is a phylogenetic cladogram by Butler et al. in 2011 showing the cladistics of Archosauriformes, focusing mostly on Pseudosuchia: [9] Clade names follow Nesbitt 2011. [1]

Crurotarsi

Related Research Articles

<span class="mw-page-title-main">Archosaur</span> Group of diapsids broadly classified as reptiles

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.

<span class="mw-page-title-main">Crurotarsi</span> Clade of reptiles

Crurotarsi is a clade of archosauriform reptiles that includes crocodilians and stem-crocodilians and possibly bird-line archosaurs too if the extinct, crocodile-like phytosaurs are more distantly related to crocodiles than traditionally thought. Prior to 2011, the group had invariably included only archosaurs closer to crocodilians than to birds and other dinosaurs. An equivalent term for the crocodilian side of the archosaur family tree is Pseudosuchia. This traditional definition of Crurotarsi assumed that phytosaurs were crown-group archosaurs and more closely related to crocodilians than to birds. However, a 2011 study argued that the phytosaur lineage evolved prior to the split between birds and crocodilians. This would mean that phytosaurs were not true archosaurs, and therefore could not be considered representatives of croc-line archosaurs.

<span class="mw-page-title-main">Rauisuchia</span> Informal group of Triassic archosaurs with pillar-erect posture

"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

<span class="mw-page-title-main">Aetosaur</span> Extinct order of heavily armoured reptiles

Aetosaurs are heavily armored reptiles belonging to the extinct order Aetosauria. They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs more closely related to crocodilians than to birds and other dinosaurs. All known aetosaurs are restricted to the Late Triassic, and in some strata from this time they are among the most abundant fossil vertebrates. They have small heads, upturned snouts, erect limbs, and a body ornamented with four rows of plate-like osteoderms. Aetosaur fossil remains are known from Europe, North and South America, parts of Africa, and India. Since their armoured plates are often preserved and are abundant in certain localities, aetosaurs serve as important Late Triassic tetrapod index fossils. Many aetosaurs had wide geographic ranges, but their stratigraphic ranges were relatively short. Therefore, the presence of particular aetosaurs can accurately date a site in which they are found.

<i>Gracilisuchus</i> Genus of fossil reptiles

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.

<span class="mw-page-title-main">Pseudosuchia</span> Clade of reptiles

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.

<span class="mw-page-title-main">Prestosuchidae</span> Extinct family of reptiles

Prestosuchidae is a polyphyletic grouping of carnivorous archosaurs that lived during the Triassic. They were large active terrestrial apex predators, ranging from around 2.5 to 7 metres in length. They succeeded the Erythrosuchidae as the largest archosaurs of their time. While resembling erythrosuchids in size and some features of the skull and skeleton, they were more advanced in their erect posture and crocodile-like ankle, indicating more efficient gait. "Prestosuchids" flourished throughout the whole of the middle, and the early part of the late Triassic, and fossils are so far known from Europe, India, Africa (Tanzania), Argentina, and Paleorrota in Brazil. However, for a long time experts disagree regarding the phylogenetic relationships of the group, what genera should be included, and whether indeed the "Prestosuchidae" constitute a distinct family.

<i>Prestosuchus</i> Extinct genus of reptiles

Prestosuchus is an extinct genus of pseudosuchian in the group Loricata, which also includes Saurosuchus and Postosuchus. It has historically been referred to as a "rauisuchian", and was the defining member of the family Prestosuchidae, though the validity of both of these groups is questionable: Rauisuchia is now considered paraphyletic and Prestosuchidae is polyphyletic in its widest form.

<span class="mw-page-title-main">Ornithosuchidae</span> Extinct family of reptiles

Ornithosuchidae is an extinct family of pseudosuchian archosaurs from the Triassic period. Ornithosuchids were quadrupedal and facultatively bipedal, meaning that they had the ability to walk on two legs for short periods of time. They had distinctive, downturned snouts, unique, "crocodile-reversed" ankle bones, and several other features that distinguish them from other archosaurs. Ornithosuchids were geographically widespread during the Carnian and Norian stages of the Late Triassic with members known from Argentina, Brazil, and the United Kingdom. Four genera, comprising Ornithosuchus, Venaticosuchus, Dynamosuchus, and Riojasuchus are presently known. The family was first erected by German paleontologist Friedrich von Huene in 1908.

<span class="mw-page-title-main">Crurotarsal</span>

A crurotarsal joint is one that’s situated between the bones of crus, i.e. shin and the proximal tarsal bones, i.e. astragalus and calcaneum.

<i>Turfanosuchus</i> Extinct genus of reptiles

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.

<i>Luperosuchus</i> Extinct genus of reptiles

Luperosuchus is an extinct genus of loricatan pseudosuchian reptile which contains only a single species, Luperosuchus fractus. It is known from the Chañares Formation of Argentina, within strata belonging to the latest Ladinian stage of the late Middle Triassic, or the earliest Carnian of the Late Triassic. Luperosuchus was one of the largest carnivores of the Chañares Formation, although its remains are fragmentary and primarily represented by a skull with similarities to Prestosuchus and Saurosuchus.

<span class="mw-page-title-main">Loricata</span> Extinct clade of reptiles

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.

<span class="mw-page-title-main">Poposauroidea</span> Extinct clade of reptiles

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.

<i>Decuriasuchus</i> Extinct genus of reptiles

Decuriasuchus is an extinct genus of loricatan from the Middle Triassic period. It is a carnivorous archosaur that lived in what is now southern Brazil, in Paleorrota. It was first named by Marco Aurélio G. França, Jorge Ferigolo and Max C. Langer in 2011 and the type species is Decuriasuchus quartacolonia. The generic name means "unit of ten crocodiles" in Latin and Greek in reference to the ten known specimens and the animal's possible group behavior. The specific name refers to the Quarta Colonia region where the fossils were collected.

<span class="mw-page-title-main">Paracrocodylomorpha</span> Clade of reptiles

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.

<span class="mw-page-title-main">Erpetosuchidae</span> Extinct family of reptiles

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.

Schultzsuchus is an extinct genus of pseudosuchian from the Triassic Santa Maria Formation of Brazil. Initially described as a species of Prestosuchus, a 2024 study found the taxon to differ in multiple aspects of its anatomy, enough to warrant it being placed in a distinct genus. Unlike Prestosuchus, which is a more derived member of Loricata, Schultzsuchus appears to have been a basal member of the clade Poposauroidea.

References

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