Poposaurus

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Poposaurus
Temporal range: Late Triassic (Carnian to Norian), 237–216  Ma
Poposaurus.jpg
Mounted skeleton, Yale Peabody Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Poposauroidea
Family: Poposauridae
Genus: Poposaurus
Mehl, 1915
Species
  • P. gracilisMehl, 1915 (type)
  • P. langstoni(Long and Murry, 1995 [originally Lythrosuchus langstoni])
Synonyms
  • LythrosuchusLong and Murry, 1995

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".

Contents

Like theropod dinosaurs, Poposaurus was an obligate biped, meaning that it walked on two legs rather than four. However, as a pseudosuchian, it is more closely related to living crocodilians than to dinosaurs. [1] Poposaurus is thought to have evolved this form of locomotion independently, possibly from early archosaurs' ability to high walk.

History

An ilium (hip bone) assigned to Paleorhinus bransoni by J. H. Lees and later identified as Poposaurus gracilis. Paleorhinus bransoni ilium Lees.jpg
An ilium (hip bone) assigned to Paleorhinus bransoni by J. H. Lees and later identified as Poposaurus gracilis.
Illustration of the holotype. Poposaurus gracilis Mehl.jpg
Illustration of the holotype.

The first remains of Poposaurus were found in 1904 near Lander, Wyoming. In 1907, paleontologist J. H. Lees described this fossil, an ilium (part of the hip) from the Popo Agie Formation, and identified it as that of the phytosaur Paleorhinus bransoni . [2] In 1915, paleontologist M. G. Mehl named Poposaurus based on more complete material from the Popo Agie Formation, including vertebrae, hips, and limb bones. He cited the holotype as [Walker Museum] 602, but in fact the holotype is UR 357. Mehl concluded that the ilium described by Lees, UR 358, also belonged to Poposaurus. He did not classify Poposaurus as a phytosaur because the shape of its ilium was different and it had more sacral vertebrae fused to the hip. Mehl made comparisons between Poposaurus and the earlier named Dolichobrachium , also from the Triassic of Wyoming. Dolichobrachium was only known from some teeth, a humerus, and part of the pectoral girdle, so Mehl suggested that the Poposaurus and Dolichobrachium material could belong to the same animal. Mehl noted similarities between Poposaurus and theropod dinosaurs, including its hollow leg bones and deep hip socket, but did not consider it a dinosaur because each sacral vertebra supported only one rib (theropods usually have multiple ribs projecting from each sacral vertebra). [3]

In the following years, Poposaurus was assigned to many different groups of reptiles. Hungarian paleontologist Franz Nopcsa classified it as an ornithischian dinosaur in 1921, identifying similarities with iguanodonts and camptosaurs. In 1928, Nopcsa placed it in a new family called Poposauridae and a new suborder called Poposauroidea. To Nopsca, Poposauroidea was one of three suborders that made up the order Ornithopoda. Over the following years, many paleontologists supported this classification. For example, German paleontologist Oskar Kuhn classified Poposaurus in its own suborder of ornithischians, which he called Poposauria. In 1930, American paleontologist Oliver Perry Hay placed Poposaurus in Anchisauridae, a family of sauropodomorph dinosaurs. German paleontologist Friedrich von Huene considered it a very early stegosaur in 1950. [4]

In 1961, American paleontologist Edwin Harris Colbert gave an extensive description of the known material of Poposaurus and classified it as a theropod dinosaur. Colbert thought that Poposaurus could not have been a more primitive archosaur because it had hollow leg bones and complex vertebrae. He placed it in the Carnosauria, but because its ilium was distinct from all other archosaurs, Colbert placed Poposaurus in its own family, Poposauridae. In the same paper, Colbert described an ilium from the Dockum Group of Howard County, Texas, which he assigned to P. gracilis. [4]

In his 1977 study of Late Triassic saurischians, Peter Galton reclassified Poposaurus as a thecodont pseudosuchian. In 1915, Mehl described a "distal femur" in the holotype specimen of Poposaurus, but Galton interpreted this to be the fused end of the hip's pubis bones. Galton noted similarities between the hips of Poposaurus, Arizonasaurus , Bromsgroveia , Postosuchus , and Teratosaurus , and grouped them all in Poposauridae. Like paleontologists before him, Galton distinguished Poposaurus based on the unique shape of its ilium. [5]

In 1995, paleontologists Robert Long and Phillip Murry described new fossils of Poposaurus from the Placerias quarry in the Chinle Formation of Arizona. Among the new material were parts of the lower limb, including the tibia and calcaneum. They removed Postosuchus from Poposauridae, claiming that the material used in this assignment was a chimera, or a collection of bones belonging to different animals. The pubis of Postosuchus was in fact a pubis of Poposaurus, leading to the mistaken classification. Long and Murry separated poposaurids like Poposaurus, Bromsgroveia, and the newly named Lythrosuchus from rauisuchians like Postosuchus, which they held in the family Rauisuchidae. [6]

Fossils of Poposaurus gracilis have been found from the Chinle Formation in Grand Staircase-Escalante National Monument, Utah. Chinle Badlands.jpg
Fossils of Poposaurus gracilis have been found from the Chinle Formation in Grand Staircase–Escalante National Monument, Utah.

The known material of Poposaurus was again described in 2007, along with two new specimens from the Tecovas Formation of Texas and the Petrified Forest of Arizona. Long and Murry's Lythrosuchus langstoni was reclassified as a new species of Poposaurus, P. langstoni. P. langstoni differs from P. gracilis in that it is larger, it does not have a ridge of bone behind the hip socket, and does not have a pit on the ischium that fits into the ilium. [7] In 2011, a nearly complete specimen of P. gracilis known as YPM VP 057100, and informally named "the Yale specimen", was found in the Chinle Formation of Grand Staircase–Escalante National Monument, Utah. It includes the forelimbs, hind limbs, hips, ribs, dorsal vertebrae, and much of the tail. [1] Another specimen of Poposaurus from Arizona, PEFO 34865, includes not only postcranial remains but also cranial remains, confirming that Poposaurus was a hyper-carnivorous predator. [8]

In 2022, the partial remains (fragmentary thoracic vertebrae and part a right humerus) of an immature P. gracilis were described from the early Carnian-aged Doswell Formation of Virginia, marking the first occurrence of this genus from eastern North America. [9] This material was found at the same locality as Doswellia and was first mentioned in the 1980 paper describing it, where it was tentatively referred to as indeterminate rauisuchian remains. [10]

Description

Size of P. gracilis relative to a human. Poposaurus gracilis scale.svg
Size of P. gracilis relative to a human.

With the tail comprising about half the body length, Poposaurus was about 4–5 metres (13–16 ft) long and weighed 90 to 100 kilograms (200 to 220 lb) as an adult. [1] [11] The body of Poposaurus is laterally compressed, with a long and narrow hip structure. The pubis and ischium are elongated. The end of the pubis forms a distinct hook that is unique to Poposaurus and a few other early pseudosuchians. Poposaurus has five sacral vertebrae connecting the spine to the hip, three more than most early archosaurs. The hind legs are about twice as long as the arms and placed close together. Five digits are present on the foot, but the fifth is reduced to a small splint of bone next to the metatarsals. The calcaneum bone extends far from the ankle to form a distinct heel.


Classification

Poposaurus is a member of the family Poposauridae, part of the larger pseudosuchian group Poposauroidea. It is closely related to other Triassic pseudosuchians like ctenosauriscids and shuvosaurids. Like Poposaurus, shuvosaurids were bipedal. When the specimen YPM VP 057100 was described by Gauthier et al. in 2011, Poposaurus was included in a phylogenetic analysis. Poposaurus was placed within Poposauroidea as the sister taxon to the large-bodied herbivorous Lotosaurus and the shuvosaurids. This means that Poposaurus is more closely related to Lotosaurus and shuvosaurids than it is to any other pseudosuchian. The analysis found ctenosauriscids and the aquatic Qianosuchus to be successively more basal poposauroids. Below is the cladogram from Gauthier et al. (2011): [1]

Life restoration of P. gracilis. Poposaurus gracilis.jpg
Life restoration of P. gracilis.
Archosauria  

Paleobiology

Locomotion

The hypothesized leg musculature of P. gracilis, with leg bones (top), superficial muscles (middle), and deep muscles (bottom). Poposaurus gracilis leg muscles.svg
The hypothesized leg musculature of P. gracilis, with leg bones (top), superficial muscles (middle), and deep muscles (bottom).

When M. G. Mehl first named Poposaurus in 1915, he described it as "a well-muscled creature light in weight, possibly bipedal in gait occasionally, and most assuredly swift in movement." [3] Mehl based this description on its long limb bones and deep hip socket, two features which link it with bipedal dinosaurs. Since the 1970s however, Poposaurus has been considered a pseudosuchian archosaur more closely related to crocodilians than dinosaurs. Most of its close relatives (such as the large-bodied rauisuchids and ctenosauriscids) were obligate quadrupeds that could not walk on two legs. Although the entire skeleton was unknown, Poposaurus was expected to be similar in appearance to its relatives. In 2011, the nearly complete subadult specimen YPM VP 057100 was described. The specimen confirmed Mehl's description, revealing that Poposaurus was indeed bipedal. The skeleton preserves both the fore and hind limbs, showing that Poposaurus had much shorter arms than legs. [1]

Although Poposaurus and early dinosaurs were both bipedal, the method of locomotion evolved independently in each group. The independent origins are shown through several differences in the skeletons of Poposaurus and dinosaurs. Unlike dinosaurs, Poposaurus has the characteristic crurotarsal ankle of pseudosuchians, usually associated with quadrupedal locomotion. Poposaurus also has a "pillar erect" stance in which the acetabulum or hip socket faces downward and is positioned directly over the head of the femur. In contrast, dinosaurs have "buttress erect" hip structures in which the acetabulum faces laterally and the head of the femur is angled to fit into it. [12]

Although they evolved bipedal locomotion independently, Poposaurus and dinosaurs inherited a propensity for erect hind-limb driven movement from an early archosaur ancestor. The posture of this ancestral archosaur can be inferred from a method called extant phylogenetic bracketing. [13] Archosauria is a crown group represented today by birds and crocodilians, meaning that the first archosaur was the last common ancestor of all birds and crocodilians. All birds have a fixed erect stance, and crocodilians have the ability to high walk with their limbs erect. If an erect stance is considered homologous in birds and crocodilians (most likely), phylogenetic bracketing implies that they inherited this trait from their common ancestor and that this ancestor also had an erect stance. With this reasoning, the first archosaurs are thought to have had the ability to high walk. Poposaurus and dinosaurs achieved a bipedal posture as their legs increased in size, their hips strengthened, and their spines adapted for dorsoventral flexion. Other adaptations that may have facilitated bipedal locomotion include the development of a chambered heart and lungs with unidirectional airflow (both of which are assumed present in Poposaurus through phylogenetic bracketing). [1] [13]

The leg musculature of Poposaurus was hypothesized in a 2011 study that examined muscle scars on the bones and made inferences based on phylogenetic bracketing. 26 muscles, three ligaments, and two connective tissue structures were described. While the hypothesized muscles of Poposaurus share many aspects with those of birds, they are more similar to those of crocodilians. Poposaurus is thought to have had a puboischiotibialis muscle, but this muscle is absent in birds and probably non-avian dinosaurs as well. The extensor digitorum brevis was probably present on the foot of Poposaurus, but not in birds. The puboischiofemoralis externus muscle of Poposaurus is also similar to those of living crocodilians. Other aspects of the muscles of Poposaurus differ from those of crocodilians. For example, the puboischiofemoralis internus muscle originates on the spine in crocodilians and on the hip in Poposaurus. The hip origin for this muscle is considered to be the original condition for archosaurs, since it is also seen in birds and non-avian dinosaurs. Poposaurus is thought to have had adductor muscles that were even larger than dinosaurs, as their insertion site runs along the entire length of the femur. [12]

Breathing

The 2011 study of the leg musculature of Poposaurus also suggested a distinctive form of respiration that involved abdominal muscles. The study hypothesized that Poposaurus had an ischiotruncus muscle running from the ischium at the back of the hip, across the pubis, and into the gastralia bones of the abdomen. In a form of respiration called cuirassal breathing, the ischiotruncus would contract and compress the trunk, pumping the lungs. [12]

Related Research Articles

<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.

<i>Postosuchus</i> Genus of reptiles

Postosuchus, meaning "Crocodile from Post", is an extinct genus of rauisuchid reptiles comprising two species, P. kirkpatricki and P. alisonae, that lived in what is now North America during the Late Triassic. Postosuchus is a member of the clade Pseudosuchia, the lineage of archosaurs that includes modern crocodilians. Its name refers to Post Quarry, a place in Texas where many fossils of the type species, P. kirkpatricki, were found. It was one of the apex predators of its area during the Triassic, larger than the small dinosaur predators of its time. It was a hunter which probably preyed on large bulky herbivores like dicynodonts and many other creatures smaller than itself.

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

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.

<i>Effigia</i> Extinct genus of Archosaurs

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.

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

Marasuchus is a genus of basal dinosauriform archosaur which is possibly synonymous with Lagosuchus. Both genera lived during the Late Triassic in what is now La Rioja Province, Argentina. Marasuchus contains a single species, Marasuchus lilloensis.

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

Poposauridae is a family of large carnivorous archosaurs which lived alongside dinosaurs during the Late Triassic. They were around 2.5 to 5 metres long. Poposaurids are known from fossil remains from North and South America. While originally believed to be theropod dinosaurs, cladistic analysis has shown them to be more closely related to crocodiles.

<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", 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.

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

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.

<i>Terrestrisuchus</i> Genus of terrestrial crocodylomorph

Terrestrisuchus is an extinct genus of very small early crocodylomorph that was about 76 centimetres (30 in) long. Fossils have been found in Wales and Southern England and date from near the very end of the Late Triassic during the Rhaetian, and it is known by type and only known species T. gracilis. Terrestrisuchus was a long-legged, active predator that lived entirely on land, unlike modern crocodilians. It inhabited a chain of tropical, low-lying islands that made up southern Britain, along with similarly small-sized dinosaurs and abundant rhynchocephalians. Numerous fossils of Terrestrisuchus are known from fissures in limestone karst which made up the islands it lived on, which formed caverns and sinkholes that preserved the remains of Terrestrisuchus and other island-living reptiles.

<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.

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

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.

The Popo Agie Formation is a Triassic geologic formation that crops out in western Wyoming, western Colorado, and Utah. It was deposited during the Late Triassic in fluvial (river) and lacustrine (lake) environments that existed across much of what is now the American southwest. Fragmentary fossils of prehistoric reptiles and amphibians, including pseudosuchian reptiles and temnospondyl amphibians, have been discovered in the Popo Agie Formation. Dinosaur remains are also among the fossils that have been recovered from the formation, although none have yet been referred to a specific genus.

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

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.

<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. 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, Ornithosuchus woodwardi, or Aetosaurus ferratus.

<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.

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

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.

<i>Incertovenator</i> Extinct genus of probable archosaur

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.

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