Arganasuchus

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Arganasuchus
Temporal range: Carnian
Arganasuchus dutuiti hemimandible left.jpg
Left mandible, specimen ALM 2
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Loricata
Genus: Arganasuchus
Jalil & Peyer, 2007
Type species
Arganasuchus dutuiti
Jalil & Peyer, 2007

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. [1] [2]

Discovery

Fossils of Arganasuchus were first reported by Jean-Michel Dutuit in 1979, who referred a maxilla, dentary, femur, and fibula to Ticinosuchus. These fossils were found in the lower part of unit T5 (the Irohalene Member) of the Timezgadiouine Formation. This geological formation, which is found in the Argana Basin of Morocco, may have been deposited in the Carnian stage of the Late Triassic period. [1]

Arganasuchus dutuiti was named and described in 2007 by Nour-Eddine Jalil and Karin Peyer. Its name translates to "Dutuit's Argana crocodile" in honor of its discoverer and place of discovery. The genus was based on the holotype AZA 904, a partial pubis bone found at a fossil site near the village of Azarifen. Other Arganasuchus fossils from Azarifen include ribs and a chevron (AZA 903-1-3, and 407-1-5), teeth (AZA 408-1-3), a femur (AZA 900), a fibula (AZA 901), part of a cervical vertebra (AZA 902), part of a neural arch (AZA 905), and part of a tibia (AZA 906). Skull bones have been found at a different locale, the Alili n’yifis site near the village of Alma. These skull bones include a maxilla (ALM 1), and several lower jaws (ALM 2, 3, 5, 6, and 7). One of the jaws, ALM 2, connected to a small angled bone tentatively identified as a quadratojugal. The referral of the skull fossils to Arganasuchus is uncertain but likely considering their "rauisuchian" identity and similar size and occurrence. [1]

Description

Skull

The maxilla has a straight lower edge with at least 12 large, knife-like teeth set in deep sockets and covered by interdental plates. A longitudinal ridge runs along the inner portion of the maxilla and is continuous with a low facet for the palatine bone. This continuity is seemingly unique to Arganasuchus. The maxilla as a whole is Y-shaped like Batrachotomus and Fasolasuchus . It has a long and tapering posterior process (rear branch) and a shorter ascending process (upper branch) separated by a triangular antorbital fenestra which forms a 40 degrees angle. The maxilla also has an unusually long anterior process (forward branch), forming the "stem" of the Y shape. It is uncertain whether the front edge of the maxilla formed part of the naris (like Batrachotomus) or contacted the premaxilla (like other loricatans). The possible quadratojugal fossil is thick and sharply angled, apparently contacting a long jugal but not the quadrate, unlike its relatives. The front of the lower jaw is narrow and taper towards the chin, resembling that of Saurosuchus in most respects. Although the rear of the lower jaw is unknown, enough of the front half was preserved to indicate that at least 14 teeth were present. Arganasuchus has an alternating tooth replacement scheme, like many other archosaurs. [1]

Postcrania

The single known vertebra is a spool-shaped cervical (neck) centrum with a low keel running along its underside. A partial neural arch is also preserved, though the low inclination of the postzygapophyses indicates that it may belong to a dorsal (back) vertebra. The rib is similar to those of Saurosuchus (thick, two-headed, with a longitudinal ridge) while the chevron is similar to those of Ticinosuchus (thickened at its tip). The pubis is among the most unusual and unique bones in Arganasuchus. The acetabulum (hip socket) expands down the shaft of the pubis and is delineated from the front by a thick ridge which projects out as a tuberosity at its lower extent. The size and orientation of the acetabulum on the pubis is an autapomorphy (unique defining feature) of Arganasuchus. The femur is large and robust, with a prominent knob-like fourth trochanter overlooking a smaller knob which may be an area of diseased bone. The tibia is thick but incomplete while the fibula is thinner and has a pronounced iliofibularis muscle scar almost halfway down its shaft. [1]

Classification

Arganasuchus is universally considered a "rauisuchian", a grouping of large carnivorous pseudosuchians (crocodile-line archosaurs) from the Triassic. However, "Rauisuchia" is currently considered to be a paraphyletic grade of archosaurs incrementally closer to crocodilians, rather than a clade (a natural grouping defined by shared relations). The most recent cladistic interpretations of archosaur classification are mainly inspired by an analysis by Nesbitt (2011). He split "rauisuchians" (relabeled as Paracrocodylomorpha) into two branches: the unusual poposauroids and a more diverse branch called Loricata, which includes crocodylomorphs and ancestral forms like Postosuchus and Saurosuchus . [3] [2] Although most frequently compared with loricatans, Arganasuchus has yet to be properly incorporated into this newer understanding of crocodilian ancestry, and as a result its classification relative to other "rauisuchians" is poorly understood.

The original description by Jalil & Peyer (2007) made many comparisons between the anatomy of Arganasuchus and other loricatans. The shape of the maxilla was similar to Batrachotomus and Fasolasuchus , the vertebrae were similar to Postosuchus, the femur was comparable to Prestosuchus , and the pubis had a few similarities with both Batrachotomus and Postosuchus. The question of Arganasuchus's relations was left unresolved, partly because its describers were aware of the uncertainty within rauisuchian taxonomy as a whole. [1]

Some phylogenetic analyses have attempted to deal with Arganasuchus, although there is no consensus on its precise affinities. Brusatte et al. (2010) found weak support for a small clade of basal "rauisuchoids" (loricatans) containing Arganasuchus, Fasolasuchus, Stagonosuchus , and Ticinosuchus. [4] França et al. (2011) found that Arganasuchus was an unstable "wildcard" taxon in their analysis, though a connection to Decuriasuchus , Prestosuchus, and Batrachotomus was one possibility. [5]

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>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">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>Saurosuchus</i> Paracrocodylomorph reptile genus from Late Triassic period

Saurosuchus is an extinct genus of large loricatan pseudosuchian archosaurs that lived in South America during the Late Triassic period. It was a heavy, ground-dwelling, quadrupedal carnivore, likely being the apex predator in the Ischigualasto Formation.

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

Dromicosuchus is an extinct genus of sphenosuchian, a type of basal crocodylomorph, the clade that comprises the crocodilians and their closest kin. It was found in Upper Triassic rocks of North Carolina, United States, and is known from a nearly complete skull and partial skeleton. This specimen is unusual in that it was found beneath the skeleton of a larger rauisuchian and has apparent bite damage, suggesting that it was attacked by the larger carnivore before both died and were buried together.

<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.

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.

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

Fasolasuchus is an extinct genus of loricatan. Fossils have been found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina that date back to the Norian stage of the Late Triassic, making it one of the last "rauisuchians" to have existed before the order became extinct at the end of the Triassic.

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

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

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

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.

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">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">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.

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.

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

  1. 1 2 3 4 5 6 Jalil, Nour-Eddine; Peyer, Karin (2007). "A new rauisuchian (Archosauria, Suchia) from the Upper Triassic of the Argana Basin, Morocco". Palaeontology. 50 (2): 417–430. doi:10.1111/j.1475-4983.2007.00640.x. S2CID   140681396.
  2. 1 2 Nesbitt, Sterling J.; Brusatte, Stephen L.; Desojo, Julia B.; Liparini, Alexandre; França, Marco A. G. De; Weinbaum, Jonathan C.; Gower, David J. (2013-01-01). "Rauisuchia" (PDF). Geological Society, London, Special Publications. 379 (1): 241–274. Bibcode:2013GSLSP.379..241N. doi:10.1144/SP379.1. ISSN   0305-8719. S2CID   219193351.
  3. Nesbitt, S.J. (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.
  4. Brusatte, Stephen L.; Benton, Michael J.; Desojo, Julia B.; Langer, Max C. (2010-03-15). "The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida)". Journal of Systematic Palaeontology. 8 (1): 3–47. doi:10.1080/14772010903537732. hdl: 20.500.11820/24322ff3-e80e-45f2-8d53-d35fd104195c . ISSN   1477-2019. S2CID   59148006.
  5. França, Marco Aurélio G.; Ferigolo, Jorge; Langer, Max C. (2011-03-29). "Associated skeletons of a new middle Triassic "Rauisuchia" from Brazil". Naturwissenschaften. 98 (5): 389–395. Bibcode:2011NW.....98..389F. doi:10.1007/s00114-011-0782-3. ISSN   1432-1904. PMID   21445632. S2CID   4811617.