Benggwigwishingasuchus Temporal range: Middle Triassic | |
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Holotype of Benggwigwishingasuchus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | † Poposauroidea |
Genus: | † Benggwigwishingasuchus Smith et al., 2024 |
Species: | †B. eremicarminis |
Binomial name | |
†Benggwigwishingasuchus eremicarminis Smith et al., 2024 | |
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.
The fossil remains of Benggwigwishingasuchus were discovered by Elaine Kramer and Monica Shaffer in the outcrops of the Fossil Hill Member of the Favret Formation, located in Pershing County, Nevada. [1] [2] The holotype specimen LACM-DI 158616 consists of a partially articulated skeleton which includes parts of the skull as well as most of the spinal column, limb girdles and limbs. [1]
The genus name of Benggwigwishingasuchus combines the Shoshone term "Benggwi Gwishinga", meaning "to catch fish", with the Greek "suchus", a commonly used suffix among fossil pseudosuchians derived from the Egyptian deity Sobek. The species name meanwhile means "desert song", derived from the Latin "erema" and "carminis", and was chosen to honor the discoverers of the fossil, who are noted for their love of opera. [1]
The skull of Benggwigwishingasuchus is poorly understood, with the holotype specimen only preserving the prefrontal bones, the parietal and the frontal bone. These elements do however showcase some features considered diagnostic for the genus, like the prominent supratemporal fossa. The prefrontals are also well developed, especially the ventral ramus, a peg of bone that is directed downwards and frames the front of the eye sockets. Usually, the lacrimal would also contribute to this area, but in Benggwigwishingasuchus this part of the prefrontal is so well developed that it is likely that it entirely excludes the lacrimals from forming part of the orbits, similar to the condition seen in Litorosuchus . Another diagnostic feature of the prefrontal is a crescent-shaped (semilunate) eminence that is situated down the middle of the ventral ramus. [1]
Compared to the skull, the axial skeleton is understood much better, in large part due to the completeness of the cervical (neck) and dorsal (torso) vertebrae. Benggwigwishingasuchus has been noted for having both a large number of neck vertebrae and a low number of dorsal vertebrae, both ranging from about 10 to 11 individual bones, the ambiguity stemming from transitional vertebrae that could belong to either series. The neural spine of the axis, the second of the cervical vertebrae, is weakly convex and the middle vertebrae of the neck possess centra that are slightly elongated relative to those of the torso. This elongation is a feature shared not only by other poposauroids but also by members of the Avemetatarsalia (dinosaurs, pterosaurs and relatives). The dorsal vertebrae, though lacking pneumatic foramina, are laterally depressed and possess prominent laminae. These house deep infrazygapophseal and infraprezygapophyseal fossae, which is yet another feature common among poposauroids and avemetatarsalians. The sacral vertebrae are unfused and partially obscured, making it uncertain how many there are. At least two were identified, with the possibility of a third being present. The vertebrae that form the base of the tail are unique among archosaurs since they preserve spatulate ribs that appear asymmetrical when seen from above. Some of the neural spines of the mid-caudal vertebrae possess accessory laminar processes, a feature shared with paracrocodylomorphs and Ticinosuchus . The chevrons have split upper ends, as seen in Qianosuchus , Poposaurus and Gracilisuchus . [1]
As with many other paracrocodylomorphs, the ilium possesses a second crest above the supracetabular crest, though, in the case of Benggwigwishingasuchus, it has been heavily weathered following preservation. The humerus has been noted for its expanded ends, both distal and proximal, and its relative shortness compared to the length of the femur. The femur bears two tubers on its inner (medial) side, one smaller and directed towards the front (a feature seen in the majority of archosaurs) and a second, larger tuber that faces towards the back. The proximal end of the femur, the region closest to the hip, bears a deep groove as in other paracrocodylomorphs and the head itself is oriented at a 45° angle relative to the opposing distal end. The fibula has a weakly sinusoidal shaft and ends in a symmetrical and weakly rounded distal end, a feature that Benggwigwishingasuchus shares with other poposauroids. The metatarsals, the bones that make up the middle of the foot, show some atypical anatomy among archosaurs, with the fourth metatarsal being longer than both the third and second. The fifth metatarsal has a hooked proximal end and its underside features a robust tubercle similar to that of Nundasuchus . The proximal-most phalanx of the fifth toe is robust and roughly as long as it is wide. [1]
Unlike derived poposauroids like Poposaurus, Benggwigwishingasuchus still retains its osteoderm armour, which is arranged in a simple paramedial double-row as in many pseudosuchians. The osteoderm pairs do not correlate 1:1 with the underlying vertebrae and instead, each vertebra is covered by multiple pairs, similar to Ticinosuchus, Qianosuchus and gracilisuchids. The individual osteoderms are exact mirrors of each other and are nearly square up until the sacrum. The edges of the osteoderms in Benggwigwishingasuchus are concave due to prominent indentations that give them an hourglass shape. The edges of the osteoderms are further marked by a series of irregular spikes and crenulations, which bear resemblance to those of some phytosaurs and Litorosuchus. [1]
Benggwigwishingasuchus was a medium-sized paracrocodylomorph, reaching a total body length of approximately 1.5 m (4 ft 11 in). [1]
The phylogenetic analysis published in the type description of Benggwigwishingasuchus recovered it as a member of the Poposauroidea, a group of highly diverse paracrocodylomorphs that form the sister clade to Loricata. Nathan Smith and colleagues recovered poposauroids in a polytomy with loricatans as well as the genera Stagonosuchus and Mambawakale , though both of them have been recovered as poposauroids themselves in other phylogenies. Within the clade Benggwigwishingasuchus is recovered as one of the most basal taxa, more derived than Mandasuchus but more basal than Qianosuchus. [1]
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The histology of Benggwigwishingasuchus resembles that of crocodylomorphs more so than that of poposauroids, which indicates that it grew rather slowly compared to more derived members of its clade. Evidence further suggests that Benggwigwishingasuchus was ectothermic and that its growth was highly dependant on environmental factors. Given the presence of multiple resting lines in the bone, this would suggest a seasonal environment. The inferred ectothermic metabolism is further supported by the histology, showing that the 8 year old individual was relatively small but still growing. [2]
Benggwigwishingasuchus is known from the Anisian Fossil Hill Member of the Favret Formation, which is thought to have been deposited around 10 km (6.2 mi) off the Panthalassic coast of Pangea. The formation is chiefly known for preserving the remains of enormous ichthyosaurs, [1] [2] yet lacks shallow-water inhabitants or coastal animals, except for Benggwigwishingasuchus, which Smith and colleagues hypothesize to have been a coastal animal. In this case, Benggwigwishingasuchus may be a panthalassic equivalent to some pseudosuchians found along the Tethys coast, namely Qianosuchus from the Guanling Formation of China and Ticinosuchus from Switzerland. [1]
Despite the circumstances of its fossilisation, there is little evidence to suggest that Benggwigwishingasuchus was a marine animal. [1] [2] The bone density of the femoral midshaft suggests that it was not a diver and the overall ratio between the length of the femur and the humerus suggests a terrestrial lifestyle. This is supported by the osteohistology, which revealed no signs of the animal possessing any adaptation to secondarily aquatic habits. [2] However, the posture of the fossil with a pronounced arched back, similar to what is seen in Qianosuchus, would suggest that it was not transported over any long distances following the death of the individual, meaning that it could have died near the coast or that it was swept out to sea and died there while foraging. [1] [2] Following the work of Motani and Vermeij, Smith and colleagues suggest that Benggwigwishingasuchus was only in the early stages of adapting to marine life, specifically by foraging directly in the sea while not consuming fresh water directly. [1] [2]
Such a lifestyle may have been more widespread across Triassic paracrocodylomorphs, as Benggwigwishingasuchus, Qianosuchus and Ticinosuchus do not form a distinct clade and thus likely acquired their adaptations independently from one another. This lifestyle may also be shared by some even more distantly related archosaurs such as Diandongosuchus , Litorosuchus and Heteropelta . Smith and colleagues reason that archosaurs may have been more predisposed to exploiting such niches, but also argue that these instances highlight the conclusion reached by Motani and Vermeij, which was that although many animals show early adaptations for and signs of transitioning to marine habitats, few fully transition to a marine lifestyle. [1]
Lagosuchus is an extinct genus of avemetatarsalian archosaur from the Late Triassic of Argentina. The type species of Lagosuchus, Lagosuchus talampayensis, is based on a small partial skeleton recovered from the early Carnian-age Chañares Formation. The holotype skeleton of L. talampayensis is fairly fragmentary, but it does possess some traits suggesting that Lagosuchus was a probable dinosauriform, closely related to dinosaurs.
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.
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".
Qianosuchus is an extinct genus of aquatic poposauroid archosaur from the middle Triassic (Anisian) Guanling Formation of Pan County, China. It is represented by two nearly complete skeletons and a crushed skull preserved in the limestone. Qianosuchus was at least 3 metres long, and had several skeletal adaptations which indicate a semi-marine lifestyle, similar to modern-day saltwater crocodiles. These adaptations have not been seen in any other archosaur from the Triassic.
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.
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.
Mandasuchus is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956, a formal description was not published until 2018.
Koilamasuchus is an extinct genus of indeterminate archosauriform from the Triassic of Argentina. It is based on an external mold of a partial postcranial skeleton from the Quebrada de los Fósiles Formation. Due to its incomplete nature, the relationships of this reptile are difficult to establish. Originally described as a non-archosaur archosauriform, later studies tentatively considered it a doswelliid or a suchian archosaur.
Suchia is a clade of archosaurs containing the majority of pseudosuchians. It was defined as the least inclusive clade containing Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, and Crocodylus niloticus by Nesbitt (2011). Generally the only pseudosuchian group which is omitted from Suchia is the family Ornithosuchidae, although at least one analysis classifies ornithosuchids as close relatives of erpetosuchids and aetosaurs. Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place.
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.
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.
Archeopelta is an extinct genus of carnivorous archosaur from the late Middle or early Late Triassic period. It was a 2 m (6 ft) long predator which lived in what is now southern Brazil. Its exact phylogenetic placement within Archosauriformes is uncertain; it was originally classified as a doswelliid, but subsequently it was argued to be an erpetosuchid archosaur.
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.
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.
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.
Litorosuchus is a genus of armored, semiaquatic archosauriform reptile from the Middle Triassic of China, closely related to the morphologically similar Vancleavea. It contains one species, L. somnii.
Aphanosauria is an extinct group of reptiles distantly related to dinosaurs. They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchia, includes modern crocodilians. Aphanosaurs possessed features from both groups, indicating that they are the oldest and most primitive known clade of avemetatarsalians, at least in terms of their position on the archosaur family tree. Other avemetatarsalians include the flying pterosaurs, small bipedal lagerpetids, herbivorous silesaurids, and the incredibly diverse dinosaurs, which survive to the present day in the form of birds. Aphanosauria is formally defined as the most inclusive clade containing Teleocrater rhadinus and Yarasuchus deccanensis but not Passer domesticus or Crocodylus niloticus. This group was first recognized during the description of Teleocrater. Although only known by a few genera, Aphanosaurs had a widespread distribution across Pangaea in the Middle Triassic. They were fairly slow quadrupedal long-necked carnivores, a biology more similar to basal archosaurs than to advanced avemetatarsalians such as pterosaurs, lagerpetids, and early dinosaurs. In addition, they seemingly possess 'crocodile-normal' ankles, showing that 'advanced mesotarsal' ankles were not basal to the whole clade of Avemetatarsalia. Nevertheless, they possessed elevated growth rates compared to their contemporaries, indicating that they grew quickly, more like birds than other modern reptiles. Despite superficially resembling lizards, the closest modern relatives of aphanosaurs are birds.
Scolotosuchus is an extinct genus of rauisuchid from the Early Triassic of Russia. The only known species, Scolotosuchus basileus, has been discovered at the Donskaya Luka Locality. It differs from all other rauisuchids in the morphology of its spine, which allowed it to both resist great stress and gave it a great range of mobility. Additionally, it may have lacked osteoderms, with its spine instead being supported in a similar manner as in dinosaurs. At 3 m (9.8 ft) it was likely the apex predator of its ecosystem.
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.