Heptasuchus

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Heptasuchus
Temporal range: Middle - Late Triassic, 238–230  Ma
Heptasuchus skull B&W.jpg
Diagram of known skull elements
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Loricata
Genus: Heptasuchus
Dawley et al., 1979
Type species
Heptasuchus clarki
Dawley et al., 1979

Heptasuchus is an extinct genus of loricatan pseudosuchian known from the Middle or Late Triassic upper Chugwater Group of Wyoming, United States. [1] It contains a single species, Heptasuchus clarki, the first formally recognized "rauisuchian" or loricatan pseudosuchian from North America. [2]

Discovery and history

The Clark locality in Wyoming, where fossils of Heptasuchus have been discovered. Clark Locality.jpg
The Clark locality in Wyoming, where fossils of Heptasuchus have been discovered.

Collected in the summer of 1977 at the newly discovered Clark locality northwest of Casper Wyoming, the specimens assigned to H. clarki were described in a brief article [2] and two master's theses at Wayne State University, Detroit. Heptasuchus was first formally described and named by Robert M. Dawley, John M. Zawiskie and J. W. Cosgriff in 1979 and the type species is Heptasuchus clarki. The generic name is derived from epta (ἑπτά), "seven" in Ancient Greek, and suchus (συχος) which is the Latinised form of the Ancient Greek word for the crocodile god of ancient Egypt. The specific name honors George Clark of Casper for discovering the type locality of Heptasuchus. UW 11562, a partial skull and associated postcranial skeleton was designed as the holotype and UW 11563 through UW 11565, representing partial cranial and postcranial remains not associated with the holotype were designed as paratypes. [1] [2] Little research followed the initial report, although Heptasuchus has been listed in various publications summarizing fossils of Wyoming and the Triassic. Thought to be lost until 1997, most of the specimens now reside at the University of Wyoming, although some elements are still unaccounted for.

The type locality of H. clarki occurs within a succession of red beds and carbonates above the Crow Mountain Formation which in turn overlies the Alcova Limestone within the Chugwater Group. The Chugwater Group is exposed as an outcrop belt along the trend of the Casper Arch flanking the southeastern Big Horn Mountains. The age of the type locality is not precisely known, but since it is lithostratigraphically equivalent to the lower Popo Agie Formation, though possibly chronostratigraphically older, it is probably Carnian or late Ladinian. [1] Zawiskie et al. (2011) suggested that all of the pseudosuchian elements collected at the locality belong to Heptasuchus and the taxon is not a chimera as previously suggested by Wroblewski (1997). [3] About 50% of the osteology of the taxon is preserved from a minimum of four individuals with overlapping skeletal elements. [1]

Taphonomy

The remains of Heptasuchus accumulated at the base of a muddy sheet flood deposit that includes intraformational carbonate-rich conglomerate co-mingled with the skeletal elements, at what was once thought to be the base of the Popo Agie Formation in a silty, sandy matrix overlain by an oxidized mudstone. The fossil locality probably represent a drought assemblage since the specimens are weathered, disarticulated, and confined to a small area that later served as a locus for deposition of water and sediment. Thoroughly disarticulated and showing signs of subaerial weathering, the remains of Heptasuchus occur alongside those of small, unidentified archosaurs and phytosaurs (represented by teeth and possible skeletal elements or in the case of the small archosaur, a nearly complete series of vertebral centra). [3]

Description

Skeletal reconstruction of Heptasuchus holotype showing bones in white where specimens are figured or in collections, and those in grey that have been reported but not figured or available at the time of illustration Hepta Skel.jpg
Skeletal reconstruction of Heptasuchus holotype showing bones in white where specimens are figured or in collections, and those in grey that have been reported but not figured or available at the time of illustration

Heptasuchus was a medium-sized archosaur, with a skull about 65 cm (25.6 inches) long and an estimated total length of about 4.9-5.2 meters (16–17 feet) based on comparisons to Batrachotomus and 18–23 feet based on Dawley et al.'s restoration. It is unknown whether this taxon was an obligate quadruped or faculative biped and the only limb elements represented are a fragmentary humerus, an ulna, a tibia, calcanea, possible metapodials, and some phalanges that may be referable. No complete skull is known, but preserved elements indicate a taxon very similar to Batrachotomus from Germany with an arched nasal, large naris, and a postorbital bar that enters the orbit. Reconstructions of Heptasuchus depicting this taxon with unusually large premaxillary teeth are based on a single premaxillary tooth that has partially slipped out of its socket. The hip had a long, straight pubis which was curved in anterior view and had a pubic boot. The ulna was gracile and lacks a strong olecranon process seen in Postosuchus . [4] Vertebrae are represented by cervical centra, some neural spines from the dorsals or caudals, and caudal centra. Fragments of ribs, gastralia, and possible osteoderms (bony scutes) were collected as well, but have not been formally described or illustrated.

Zawiskie et al. (2011) diagnosed Heptasuchus based on two autapomorphies, anatomical features that differentiate it from all other known archosaurs: the presence of large, posteriorly directed flanges on the parabasisphenoid and a distinct, orbit-overhanging postfrontal. [1]

Classification

Life reconstruction Heptasuchus clarki Life Reconstruction Gabriel Ugueto.png
Life reconstruction

Heptasuchus was originally assigned to Rauisuchidae, based on similarities and traits shared with taxa like Prestosuchus , Postosuchus and Saurosuchus . [2] It was suggested that Heptasuchus may prove to be synonymous with Poposaurus since they were both thought to be from the Popo Agie Formation, [5] but this hypothesis has since been refuted. [6] Early cladistic studies divided "rauisuchian" taxa between the two families of Rauisuchia; the Rauisuchidae and the Prestosuchidae. [7] [8] [9] Subsequent phylogenetic analyses, that excluded Heptasuchus, found Rauisuchia and Prestosuchidae to be non-monophyletic, placing "prestosuchids" and some taxa previously assigned to Rauisuchidae outside the clade containing Rauisuchidae and Crocodylomorpha. [10] A phylogenetic analysis of archosaurs presented in an abstract found sister taxon relationship between Heptasuchus and Batrachotomus within Loricata, with a strong support of at least six shared discrete character states. [1]

Paleoecology

Heptasuchus may have shared its environment with other taxa similar to ones recovered from Popo Agie Formation outcrops along the Wind River Range. These include the phytosaurs Angistorhinus and Paleorhinus , the poposaurid Poposaurus , the dicynodont Eubrachiosaurus and trematosaurian amphibians of the family Metoposauridae. Indeterminate specimens of possible archosauriformes have also been collected from the site, but their identity remains unknown. [2] [3] Heptasuchus was a hypercarnivore and may have hunted or scavenged these animals and any others it came across.

Since several individuals are represented by the type and referred material in a single locality, it has been suggested that Heptasuchus may have lived in groups. Such aggregations are common among other loricatan genera (e.g. Batrachotomus , Decuriasuchus and Postosuchus ) [1] and predators in general (e.g. Allosaurus , dire wolf and modern alligators), [11] suggesting that these animals, being hypercarnivores, may have dominated dwindling water supplies during drought before later succumbing to the drought themselves. Similar aggregations of predators include the famous Allosaurus concentration at the Cleveland-Lloyd dinosaur quarry in the Morrison Formation of Utah and the asphalt pits at Rancho LaBrea from the Pleistocene of Los Angeles. [11] [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.

<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". 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. 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.

<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>Poposaurus</i> Extinct genus of Archosaur

Poposaurus is an extinct genus of pseudosuchian archosaur from the Late Triassic of the southwestern 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, and Texas. 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".

<i>Batrachotomus</i> Genus of reptiles

Batrachotomus is a genus of prehistoric archosaur. Fossils of this animal have been found in southern Germany and dated from the Ladinian stage of the Middle Triassic period, around 242 to 237 million years ago. Batrachotomus was described by palaeontologist David J. Gower 22 years after its discovery.

<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.

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.

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.

Procerosuchus is an extinct genus of loricatan archosaur. Fossils have been collected from the Late Triassic Santa Maria Formation in Geopark of Paleorrota, Rio Grande do Sul, Brazil, which is Carnian in age. The genus was first described by the German paleontologist Friedrich von Huene in 1942.

Stagonosuchus is an extinct genus of loricatan, or possibly a species of Prestosuchus. Fossils have been found from the Late Triassic Manda Formation in Tanzania that are Anisian in age.

<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.

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

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.

<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.

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

Vivaron is a genus of rauisuchid known from the Late Triassic Chinle Formation in New Mexico. It is the second rauisuchid known from the southwestern United States, and it highlights the wide biogeographic range similar rauisuchid taxa occupied during the Late Triassic across Pangaea, despite the varied faunal assemblages at different latitudes.

Etjosuchus is an extinct genus of "rauisuchian" (loricatan) archosaur from the Triassic of Namibia. It is known from a single species, Etjosuchus recurvidens, which is based on a partial skeleton from the Ladinian or Carnian-age Omingonde Formation.

References

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  3. 1 2 3 Wroblewski, A.F.-J. (1997). "Mixed assemblages and the birth of a chimera: an example from the Popo Agie Formation (Upper Triassic) Wyoming". Journal of Vertebrate Paleontology. 17 (Supp. 3): 86A. doi:10.1080/02724634.1997.10011028.
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  6. Sterling J. Nesbitt (2011). "The Early Evolution of Archosaurs: Relationships and the Origin of Major Clades". Bulletin of the American Museum of Natural History. 352: 1–292. doi: 10.1206/352.1 . hdl:2246/6112. S2CID   83493714.
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  9. Gower, D. J. (2002). "Archosaurian anatomy and palaeontology. Essays in memory of Alick D. Walker. Edited by D. B. Norman and D. J. Gower. Braincase evolution in suchian archosaurs (Reptilia: Diapsida): Evidence from the rauisuchian Batrachotomus kupferzellensis". Zoological Journal of the Linnean Society. 136: 49–76. doi: 10.1046/j.1096-3642.2002.00025.x .
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  11. 1 2 Gates, T. A. (2005). "The Late Jurassic Cleveland-Lloyd Dinosaur Quarry as a Drought-Induced Assemblage". PALAIOS. 20 (4): 363–375. Bibcode:2005Palai..20..363G. doi:10.2110/palo.2003.p03-22. PMC   4032541 . PMID   24806709.
  12. "Archived copy". Archived from the original on 2013-10-30. Retrieved 2013-09-18.{{cite web}}: CS1 maint: archived copy as title (link)