Poposauroidea Temporal range: Early-Late Triassic, | |
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life restoration and scale diagram of Poposaurus gracilis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Suchia |
Clade: | Paracrocodylomorpha |
Clade: | † Poposauroidea Nopsca, 1923 |
Subgroups | |
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Synonyms | |
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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 (crocodile ancestors) 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. It was phylogenetically defined in 2011 by Sterling Nesbitt as Poposaurus gracilis and all taxa more closely related to it than to Postosuchus kirkpatricki , Crocodylus niloticus (the Nile crocodile), Ornithosuchus woodwardi , or Aetosaurus ferratus . [4]
Poposauroids went extinct at the end of the Triassic period along with other non-crocodylomorph pseudosuchians. They were among the most diverse and longest lasting members of non-crocodylomorph Pseudosuchia, with Xilousuchus (a ctenosauriscid) living near the very beginning of the Triassic and Effigia (a shuvosaurid) surviving up until near the end of the Triassic. Despite the high level of diversity and anatomical disparity within Poposauroidea, certain features of the clade can be determined, particularly in the structure of the snout and pelvis (hip). Many of these features are examples of convergent evolution with dinosaurs, with bipedal poposauroids such as Poposaurus and shuvosaurids having been mistaken for theropod dinosaurs in the past. [4]
Poposauroidea was a diverse group of pseudosuchians, containing genera with many different ecological adaptations. Some (Poposaurus and shuvosaurids) were short-armed bipeds, while others (ctenosauriscids and Lotosaurus) were robust quadrupeds with elongated neural spines, creating a 'sail' like that of certain "pelycosaurs" (like Dimetrodon ) and spinosaurids. Lotosaurus and shuvosaurids were toothless and presumably beaked herbivores while Qianosuchus, Poposaurus and ctenosauriscids were sharp-toothed predators. The ecological disparity of many members of this clade means that it is difficult to assess what the ancestral poposauroid would have looked like.
Poposauroids can be differentiated from other pseudosuchians by the structure of the tip of the snout, particularly the premaxillary bone which lies in front of the nares (nostril holes). This bone possesses two bony extensions ("processes") which wrap around the nares. The anterodorsal process, which wraps above the nares to contact the nasal bones on the top edge of the snout, is typically quite short in pseudosuchians. Poposauroids have elongated anterodorsal processes, longer than the main premaxillary body. The posterodorsal process, which wraps below the nares to contact the maxilla on the side of the snout, possesses the opposite state. It is much shorter in poposauroids (compared to other pseudosuchians), restricted to only a portion of the lower edge of the nares. This has the added effect of allowing the maxilla to form the rest of the hole's lower and rear edge, with the front edge of the maxilla becoming concave as well. Although these snout features are rare among pseudosuchians, they are much more common in certain avemetatarsalians (bird-line archosaurs) such as pterosaurs and saurischian dinosaurs. The rear branch of the maxilla tapers in most poposauroids, with the exception of Qianosuchus. This contrasts with loricatans, in which this branch is rectangular in shape. [4]
Poposauroids also possess several features which are unusual compared to archosaurs in general. For example, in most archosaurs each side of the braincase possesses a pit from where the internal carotid arteries may exit the brain. In early poposauroids, these pits migrated to the underside of the braincase, thereby resembling the primitive condition seen in archosaur relatives such as Euparkeria and proterochampsians. Nevertheless, this reversion is undone in shuvosaurids (and possibly earlier, although no braincase material is known in Poposaurus or Lotosaurus). In addition, most poposauroids possessed elongated necks, and all of them had long and thin cervical ribs. This second neck trait contrasts with the condition in other pseudosuchians, phytosaurs, and pterosaurs, which have short and stout cervical ribs. The neural spines of the dorsal (back) vertebrae are thin and plate-like, even in members of Poposauroidea without sails. This differs compared to the vertebrae of most other early pseudosuchians (as well as Euparkeria and phytosaurs), which have neural spines that expand outward to form a flat, rectangular surface when seen from above. [4]
Like other reptiles, the pelvis of poposauroids is formed by three plate-like bones: the ilium which lies above the acetabulum (hip socket), the pubis which is below and in front of the acetabulum, and the ischium which is below and behind the acetabulum. The ilium is a large, complex bone, with a forward-pointing (preacetabular) process, a rear-pointing (postacetabular) process, and a lower portion which forms the upper edge of the acetabulum. In most archosaurs, the lower portion of the ilium is wedge-shaped, forming the inner face of a "closed" acetabulum. In poposauroids the lower portion of the ilium is concave, creating a partially to completely "open" acetabulum formed by open space instead of bone. The only other archosaurs with open hip sockets are dinosaurs and (to a lesser extent) crocodylomorphs. The upper edge of the acetabulum is formed by a pronounced ridge on the ilium, known as a supraacetabular rim or crest. [4]
Although all poposauroids possessed open acetabula, most other specializations of the ilium did not evolve until the clade containing Poposaurus and the shuvosaurids. For example, the supraacetabular crest projects downward, rather than outward in this clade. This trait is rare in archosaurs, only evolving independently in a few early theropod dinosaurs such as Coelophysis and Dilophosaurus . Another feature is the presence of an additional diagonal crest which branches upward from the supraacetabular rim. Although such a crest evolved independently in a number of different archosaurs, this specific subset of poposauroids is unique in having the crest be inclined forward (rather than vertical) and confluent with the elongated preacetabular blade, which is another derived feature of the clade. [4]
The pubis and ischium were also specialized in poposauroids. In every other archosaur, the two bones contact each other on the lower edge of the acetabulum. In poposauroids other than Qianosuchus and Lotosaurus, the bones did not touch, leaving the acetabulum open from the sides and below. The width of the pubis is variable at different parts of its shaft. The portion near the acetabulum is thickened, but the tip of the bone (except in Qianosuchus) is very thin when seen head-on. In most other archosaurs, the pubis has a consistent width. Theropod dinosaurs and a few other archosaurs have a distal part of the pubis which is thinner than the proximal part. Shuvosaurids and Lotosaurus also possessed ischia (on either side of the body) which were fused to each other at the midline of the body. [4]
Although the ancestral archosaur only had two sacral (hip) vertebrae, many different archosaur groups acquired additional sacral vertebrae over the course of their evolution. Nesbitt (2011) argued that additional sacral vertebrae formed between these two "primordial" vertebrae. He gave the well-preserved sacrum of the poposauroid Arizonasaurus as evidence to this process. Poposauroids have three to four sacral vertebrae, with the last and third-to-last vertebrae articulating with the ilium in a way similar to the two primordial vertebrae of more primitive archosauriformes such as Euparkeria and phytosaurs. The second-to-last vertebra has a form unlike the vertebrae of these archosauriforms, and Nesbitt concluded that it was an "insertion", formed from the innermost sections of the two primordial vertebrae. Although this process is not unique to poposauroids, it is only known in a few other archosaur lineages, such as Batrachotomus , silesaurids, and dinosaurs. [4]
Basal poposauroids such as Arizonasaurus and Qianosuchus only had three sacral vertebra, with the second vertebra being the 'insertion'. More advanced poposauroids such as Poposaurus and shuvosaurids have four sacral vertebrae, the third recognizable as the insertion. This means that the first vertebra must have been another addition, seemingly the last dorsal vertebra which had been repurposed and transformed into a sacral vertebra. This incorporated dorsal vertebra called a dorsosacral. They were irregularly distributed among archosaurs, known in a few ornithosuchids and aetosaurs as well as a variety of dinosaurs (most commonly in ornithischians and theropods) [4] [5]
In almost all archosariforms, the sacral ribs of the first primordial sacral vertebra contact the ilium near the base of that bone, close to its contact with the pubis. Poposauroids had first primordial sacral ribs with additional forward branches, which lie on the inner edge of the ilium's preacetabular blade. In poposauroids more advanced than Qianosuchus, the sacral vertebrae fuse into a single bone, the sacrum. This fusion occurred incrementally, at different portions of the vertebra. For example, the zygapophyses fused together as early as the ctenosauriscids. The centra (main cylindrical portion) of the sacral vertebrae also may have fused as early as the ctenosauriscids. [4] The bases of the neural arches (the portion of the vertebrae above the spinal cord) were fused in some ctenosauriscids (Arizonasaurus) but not others ( Bromsgroveia ), and were also fused in all poposauroids more advanced than the ctenosauriscids. [6]
Unlike most pseudosuchians, poposauroids lack bony scutes known as osteoderms. The only exception to this is Qianosuchus, which possessed numerous tiny osteoderms, lying in a row extending down the neck and body. [7] In all poposauroids, the tip of the fibula (outer shin bone) is symmetrical and straight when seen from the side, rather than slanted as in other non-crocodylomorph pseudosuchians. Those more advanced than ctenosauriscids had flattened hooflike pedal unguals (toe claws). Some poposauroids had very short arms compared to the length of their legs, although disarticulation in Qianosuchus and a lack of limb material in ctenosauriscids means that it is unknown whether this trait was basal to the group as a whole. Missing data for ctenosauriscids also obscures when certain traits of the caudal vertebrae and ankle bones were gained or lost within Poposauroidea. [4]
Franz Nopcsa first used the term Poposauridae in 1923 to refer to poposauroids. At this time, the sole member of the group was Poposaurus , which was considered to be a theropod dinosaur. Over the following years, poposauroids were placed in various groups, including Saurischia, Theropoda, and Carnosauria. [8] This classification existed up until the 1970s, when better remains indicated that Poposaurus was a pseudosuchian rather than a dinosaur. Other genera such as Sillosuchus and Shuvosaurus were later erected. Like Poposaurus, Shuvosaurus was originally thought to be a theropod dinosaur. [9]
Sankar Chatterjee reclassified poposauroids as theropod dinosaurs with his description of the new genus Postosuchus in 1985. [10] Chatterjee even considered poposauroids to be the ancestors of tyrannosaurs. Postosuchus was widely considered to be a poposauroid for the next ten years and was included in many phylogenetic analyses of Triassic archosaurs. In 1995, Robert Long and Phillip A Murry noted that several specimens referred to Postosuchus were distinct from the holotype, and so they assigned those specimens to the new genera Lythrosuchus and Chatterjeea . [9]
In 2005, Sterling Nesbitt noted that "ctenosauriscids" such as Arizonasaurus, Bromsgroveia, and Lotosaurus shared many similarities with "poposaurids" such as Poposaurus, Sillosuchus, and "Chatterjeea" (now known as Shuvosaurus ). He proposed that they formed a clade (informally named "Group X") to the exception of other pseudosuchians. [6]
"Group X" was formally given the name "Poposauroidea" by Jonathan C. Weinbaum and Axel Hungerbühler in 2007. In their paper, Weinbaum and Hungerbühler described two new skeletons of Poposaurus and incorporated several new characters of the genus into a phylogenetic analysis. Poposauroidea was recovered as a monophyletic grouping, while other rauisuchians (namely Rauisuchidae and Prestosuchidae) were placed as basal forms of a new group called Paracrocodyliformes. [11]
Brusatte et al. (2010) conducted a phylogenetic study of archosaurs that resulted in a grouping referred to as Poposauroidea. Unlike many recent studies, they found Rauisuchia to be monophyletic, consisting of two major clades: Rauisuchoidea and Poposauroidea. The monophyly of Rauisuchia was not strongly supported in Brusatte et al.'s analysis. They noted that if their tree was enlarged by one step, Poposauroidea fell outside Rauisuchia to become the sister group of Ornithosuchidae, which is thought to closely related to, but outside, Rauisuchia. In their tree, Poposauroidea included genera usually classified as poposauroids as well as several other genera that were not previously placed in the group. One of these genera, Qianosuchus, is unique among pseudosuchians in its semiaquatic lifestyle. [12]
In his massive revision of archosaurs which included a large cladistic analysis, Sterling J. Nesbitt (2011) found Xilousuchus to be a poposauroid most closely related to Arizonasaurus . Nesbitt's analysis did not recover a monophyletic Rauisuchia or monophyletic Rauisuchoidea. Poposauroidea was found to be monophyletic, and more resolved than in previous analyses, with Qianosuchus as the most basal member of the group and Lotosaurus grouping with shuvosaurids instead of ctenosauriscids. The cladogram below follows Nesbitt (2011) with clade names based on previous studies. [4]
"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.
Effigia was an extinct genus of shuvosaurid known from the Late Triassic of New Mexico, south-western USA. With a bipedal stance, long neck, and a toothless beaked skull, Effigia and other shuvosaurids bore a resemblance to the ornithomimid dinosaurs of the Cretaceous Period. However, shuvosaurids were not dinosaurs, but were instead a specialized family of poposauroid pseudosuchians, meaning that their closest living relatives are crocodilians.
Arizonasaurus was a ctenosauriscid archosaur from the Middle Triassic. Arizonasaurus is found in the Middle Triassic Moenkopi Formation of northern Arizona. A fairly complete skeleton was found in 2002 by Sterling Nesbitt. The taxon has a large sailback formed by elongated neural spines of the vertebrae. The type species, Arizonasaurus babbitti, was named by Samuel Paul Welles in 1947.
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", in contrast to the "bird-like archosaurs" or Avemetatarsalia. Despite Pseudosuchia meaning "false crocodiles", the name is a misnomer as true crocodilians are now defined as a subset of the group.
Rauisuchidae is a group of large predatory Triassic archosaurs. Some disagreement exists over which genera should be included in the Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whether these should even be thought of as separate valid families. Rauisuchidae in the modern sense was defined by Sterling Nesbitt in 2011 as the most inclusive clade containing Rauisuchus tiradentes, but not Prestosuchus chiniquensis, Poposaurus gracilis, or Crocodylus niloticus. In this modern sense, rauisuchids are recovered as members of the clade Loricata, being the sister taxon of Crocodylomorpha, and being more derived than taxa such as Prestosuchus and Batrachotomus. Rauisuchids occurred throughout much of the Triassic, and may have first occurred in the Early Triassic if some archosaurian taxa such as Scythosuchus and Tsylmosuchus are considered to be within the family.
Poposaurus is an extinct genus of pseudosuchian archosaur from the Late Triassic of the southwestern and eastern 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, Texas, and Virginia. 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".
Lotosaurus is an extinct genus of sail-backed poposauroid known from Hunan Province of central China.
Ctenosauriscus is an extinct genus of sail-backed poposauroid archosaur from Early Triassic deposits of Lower Saxony in northern Germany. It gives its name to the family Ctenosauriscidae, which includes other sail-backed poposauroids such as Arizonasaurus. Fossils have been found in latest Olenekian deposits around 247.5-247.2 million years old, making it one of the first known archosaurs.
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.
Sillosuchus is a genus of shuvosaurid poposauroid archosaur that lived in South America during the Late Triassic period. Shuvosaurids were an unusual family of reptiles belonging to the group Poposauroidea; although their closest modern relatives are crocodilians, they were bipedal and lightly armored, with dinosaur-like hip and skull structures. Based on skull remains from members of the family such as Effigia, they were also toothless and likely beaked herbivores.
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.
Arganasuchus is an extinct genus of "rauisuchian" (loricatan) archosaur. It is known from a single species, Arganasuchus dutuiti. Fossils of this genus have been found in Upper Triassic rocks of the Argana Basin, Morocco. Though its remains were initially referred to Ticinosuchus when discovered during the 1970s, in 2007 it was identified as a distinct genus with unique features of the pubis and maxilla. Arganasuchus also had several anatomical details in common with Batrachotomus, Fasolasuchus, and Postosuchus, though its relations with other loricatans remains unresolved. Arganasuchus is considered a carnivore due to its large, knife-shaped teeth.
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.
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.
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. However, a subsequent study conducted by Stocker et al. indicated it to be the basalmost known phytosaur instead.
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.
Incertovenator is an extinct genus of archosauriform reptile, likely an archosaur, of uncertain affinities. Its unstable position is a result of possessing a number features found in both the bird-line avemetatarsalian archosaurs and the crocodylian-line pseudosuchians. The type and only known species is I. longicollum, which is known from single specimen discovered in the Late Triassic Ischigualasto Formation of Argentina. Incertovenator is known almost entirely by its vertebral column. This indicates that it had a relatively long neck, leading to its uncertain classification due to the convergent evolution of elongated neck vertebrae in both avemetatarsalian and pseudosuchian archosaurs.
Benggwigwishingasuchus is a genus of poposauroid pseudosuchian from the Anisian Favret Formation of Nevada. While the animal was found in marine deposits and possibly lived a coastal lifestyle, Benggwigwishingasuchus shows no clear adaptations towards marine life. The genus contains a single species: B. eremicarminis.