Luperosuchus

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Luperosuchus
Temporal range: Late Ladinian or Early Carnian,
~236.7–235.5  Ma
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Luperosuchus cranium diagram.png
Skull diagram
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Family: Prestosuchidae
Genus: Luperosuchus
Romer, 1971
Type species
Luperosuchus fractus
Romer, 1971

Luperosuchus (meaning "vexing" or "difficult crocodile") is an extinct genus of loricatan pseudosuchian reptile (historically known as a "rauisuchian") 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 . [1]

Description

Life restoration Luperosuchus1DB.jpg
Life restoration

Luperosuchus is known only from a single incomplete skull, with an associated atlas intercentrum representing the only known postcranial material. [1] However, it is likely that it was a large quadruped, similar to other basal loricatans. Isolated osteoderms from the same region were initially attributed this genus based on their size and similarity to those of other "rauisuchian" osteoderms, [2] however they were later found to be belong to the contemporary erpetosuchid Tarjadia . [3] The presence of osteoderms in Luperosuchus is therefore left ambiguous. The length has been estimated to be 4.1–4.4 metres (13–14 ft) based on rauisuchids and 3.7 metres (12 ft) based on Prestosuchus .

Skull

The skull belongs to a large animal, with the preserved portions measuring 54.5 centimetres (21.5 in) and an estimated complete length of approximately 60 centimetres (2.0 ft), resembling those of other basal loricatans. The material largely consists of the skull roof, including the nasals, maxilla and part of the premaxilla. The snout is narrow and pointed, with a tall maxilla. Only a small, rounded front portion of the antorbital fenestra is preserved, though it was likely triangular based on the height of the skull. The antorbital fossa, the basin surrounding the antorbital fenestra, was deep. The premaxilla has a characteristically long posterodorsal process that articulates with the nasals, excluding the maxilla from the external naris, which tapers posteriorly. A narrow slit was identified between the premaxilla and maxilla, as in other loricatans, however this feature is likely to be from postmortem distortion rather than anatomical. [1] An unusual feature found in Luperosuchus is the tall, mediolaterally compressed crest that sits on the front of the snout. This structure is formed by the dorsally arching nasals, and has been described as a 'roman-nose' following Romer's initial description. A similar structure is found in other loricatans, particularly Prestosuchus and Batrachotomus , although it is most prevalent in Luperosuchus. [2] [1]

The orbit is tall and has smooth boundaries, without the "keyhole" shape of many other loricatans. The upper part of the infratemporal fenestra was also tall and narrow, although it may have been much wider near the lower rear corner of the skull, which was not preserved. The prefrontal bone had a pronounced finger-like process which stretched forwards above the lacrimal bone, although it did not stretch as far as the front edge of the frontals which were adjacent to it. A palpebral bone was situated above the orbit, though it was small and fused to the frontal and postfrontal. The postorbital bone had two distinguishing features: a large rounded knob on its upper portion and an elongated lower branch. [1]

Discovery and naming

The holotype material of Luperosuchus, specimen PULR 04, was collected by Ruth Romer, wife of palaeontologist Alfred S. Romer, on January 17, 1965. [4] This specimen is stored at the La Plata Museum in La Plata, Argentina. [1] It was found near the remains of a large dicynodont at a locality about 5 km northeast of where the Chañares River emerges onto the Plano de Talampaya. The site is believed to represent the top of the lower member of the Chañares Formation, based on similarities in preservation and the composition of the surrounding matrix to other fossils known from this time interval. This would be positioned below the strata recently dated by Marsicano et al., [5] providing a minimum age of 236.1 +/- 0.6 Ma for the Luperosuchus locality. Ezcurra et al. (2017) assigned the Luperosuchus specimen to the base of the formation, within the relatively fossil-poor Tarjadia Assemblage Zone below the much more prolific Massetognathus - Chanaresuchus Assemblage Zone. [6]

Romer published his description of Luperosuchus in 1971, as part of a series of articles describing the numerous "thecodonts" he had collected from the Chañares Formation. The binomial name, Luperosuchus fractus, refers to the fragmentary and "perplexing" nature of the fossil, and is derived from the Greek lyperos ("vexing", "difficult", "troublesome") and souchos ("crocodile"), with the specific name is from the Latin fractus, meaning "broken". [2]

Romer had collected additional fragments of bone from the holotype locality, but was unsure if they belonged to Luperosuchus or the dicynodont collected with it. After rediscovering and examining these fragments, Nesbitt & Desojo determined that many of the identifiable pieces were consistent with loricatan archosaurs, and so referred them to the holotype of Luperosuchus. These additional pieces include portions of the maxilla, quadrate, fragments of the braincase and the atlas intercentrum, the only postcranial material for Luperosuchus. [1]

A second specimen, PULR 057, was reported in 2009 by Desojo & Arcucci and was referred to this genus. This specimen however was a third smaller than the type specimen, and differed from it in a number of aspects, including a less developed 'roman-nose', but these were interpreted as ontogenetic variation as the material appeared to exhibit juvenile features (including loose sutures and less developed ornamentation of the bones). [7] The referral of this material to Luperosuchus was questioned by Nesbitt & Desojo in 2017, who suggested that the differences are taxonomic, rather than ontogenetic. The holotype PULR 04 remains the only known material of Luperosuchus, with the taxonomic identity of the former referred specimen PULR 057 currently unknown. [1]

Classification

Romer identified Luperosuchus as a member of the family Rauisuchidae in 1971, however at the time of his description the relationships of Rauisuchidae to other archosaurs were poorly understood and a detailed examination of its relationships was not performed. Luperosuchus was largely ignored in later phylogenetic analyses of Triassic pseudosuchians, likely due to the poor preservation of its remains and lack of identifying characteristics, as well as an unclear understanding of the interrelationships of close relatives. [1] As such, it was typically only ever provisionally assigned to "Rauisuchia" based on general similarities, without further justification.

Luperosuchus was incorporated into a phylogenetic analysis for the first time by Nesbitt & Desojo in 2017, where it was recovered in a clade with two other South American loricatans, Saurosuchus and Prestosuchus , at the base of Loricata. [1] They found two equally plausible hypotheses for the least inclusive position of Luperosuchus; one as a sister taxon to both Saurosuchus and Prestosuchus, and another as a sister taxon to Saurosuchus, the former of which is reproduced here:

Archosauria  

A study on new Chañares vertebrates (including new specimens of Tarjadia) published in 2017 incorporated Luperosuchus and several other suchians into an archosauromorph dataset used by Ezcurra (2016). This study found Luperosuchus as the sister taxon to Decuriasuchus based on a prominent vertical peg on the supraoccipital bone of the braincase. The Luperosuchus + Decuriasuchus clade was found outside of a more restrictive Paracrocodylomorpha, crownward of Ticinosuchus and stemward of Prestosuchus. [6]

Nesbitt & Desojo (2017) was one of several phylogenetic analyses combined during the course of Da-Silva et al. (2019)'s study of a new Prestosuchus specimen. This study did not fully support the Luperosuchus + Saurosuchus + Prestosuchus clade, although it did note that the clade was only barely less optimal than their most parsimonious tree (MPT). Their MPT found these three taxa in a series leading up to more crownward loricatans, with Prestosuchus occupying the bottom (stemward) rung, Saurosuchus in the middle, and Luperosuchus at the top (crownward) rung just below Batrachotomus . [8]

Palaeobiology

Histology and growth

The skull of Luperosuchus has very few visible sutures, partly due to the material's poor preservation but also from the addition of consolidates to the surface of the specimen that obscure finer anatomical details. [1] Romer struggled to identify the suture pattern, but identified fused sutures which he interpreted as a possible indicator that the specimen was mature. [2] Later examination by Nesbitt & Desojo confirmed that many of the sutures in the skull of Luperosuchus were obliterated, with the bones in the skull fully fused, in addition to well developed sculpting of the bones. [1]

In 2008, Ricqlès et al. reported that they performed a histological study on a limb bone belonging to Luperosuchus, identified as specimen MCZ 4077. [9] However, as Luperosuchus is only definitively known from the holotype material, the referral of this bone to Luperosuchus is dubious. [1] MCZ 4077 was later referred to Tarjadia ruthae . [6]

Palaeoecology

Luperosuchus is one of the largest animals in the Chañares Formation, and indeed was one of the few large carnivores known from the formation. In the Tarjadia Assemblage Zone of the early part of the formation, Luperosuchus shared its environment with large herbivorous dicynodonts, small burrowing cynodonts, herbivorous rhynchosaurs, and the large predatory erpetosuchid Tarjadia. Other indeterminate taxa include the small loricatan represented by PULR 057, as well as specimens of very large paracrocodylomorphs, possibly representing the largest predatory archosaurs in the ecosystem. The early part of the Chañares Formation corresponds to a braided river system with periodic flooding events. The environment later shifts to encompass deposits of pyroclastic and volcaniclastic materials, such as tuff and ash, indicating a complex history of regional volcanism in the environment. Volcanic mass mortality events may have contributed to the abundance of well-preserved skeletal material within the Massetognathus - Chanaresuchus Assemblage Zone near the stratigraphic middle of the Chañares Formation. [6] [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>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.

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

Lewisuchus is a genus of archosaur that lived during the Late Triassic. As a silesaurid dinosauriform, it was a member of the group of reptiles most commonly considered to be the closest relatives of dinosaurs. Lewisuchus was about 1 metre (3.3 ft) long. Fossils have been found in the Chañares Formation of Argentina. It exhibited osteoderms along its back.

<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>Erythrosuchus</i> Extinct genus of reptiles

Erythrosuchus is an extinct genus of archosauriform reptiles from the Triassic of South Africa. Remains have been found from the Cynognathus Assemblage Zone of the Beaufort Group in the Karoo of South Africa.

<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>Doswellia</i> Extinct genus of reptiles

Doswellia is an extinct genus of archosauriform from the Late Triassic of North America. It is the most notable member of the family Doswelliidae, related to the proterochampsids. Doswellia was a low and heavily built carnivore which lived during the Carnian stage of the Late Triassic. It possesses many unusual features including a wide, flattened head with narrow jaws and a box-like rib cage surrounded by many rows of bony plates. The type species Doswellia kaltenbachi was named in 1980 from fossils found within the Vinita member of the Doswell Formation in Virginia. The formation, which is found in the Taylorsville Basin, is part of the larger Newark Supergroup. Doswellia is named after Doswell, the town from which much of the taxon's remains have been found. A second species, D. sixmilensis, was described in 2012 from the Bluewater Creek Formation of the Chinle Group in New Mexico; however, this species was subsequently transferred to a separate doswelliid genus, Rugarhynchos. Bonafide Doswellia kaltenbachi fossils are also known from the Chinle Formation of Arizona.

<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.

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.

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

Chanaresuchus is an extinct genus of proterochampsid archosauriform. It was of modest size for a proterochampsian, being on average just over a meter in length. The type species is Chanaresuchus bonapartei was named in 1971. Its fossils were found in from the early Carnian-age Chañares Formation in La Rioja Province, Argentina. Chanaresuchus appears to be one of the most common archosauriforms from the Chañares Formation due to the abundance of specimens referred to the genus. Much of the material has been found by the La Plata-Harvard expedition of 1964-65. Chanaresuchus is the most well-described proterochampsid in the subfamily Rhadinosuchinae.

The Chañares Formation is a Carnian-age geologic formation of the Ischigualasto-Villa Unión Basin, located in La Rioja Province, Argentina. It is characterized by drab-colored fine-grained volcaniclastic claystones, siltstones, and sandstones which were deposited in a fluvial to lacustrine environment. The formation is most prominently exposed within Talampaya National Park, a UNESCO World Heritage Site within La Rioja Province.

<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>Tarjadia</i> Extinct genus of reptiles

Tarjadia is an extinct genus of erpetosuchid pseudosuchian, distantly related to modern crocodilians. It is known from a single species, T. ruthae, first described in 1998 from the Middle Triassic Chañares Formation in Argentina. Partial remains have been found from deposits that are Anisian-Ladinian in age. Long known mostly from osteoderms, vertebrae, and fragments of the skull, specimens described in 2017 provided much more anatomical details and showed that it was a fairly large predator. Tarjadia predates known species of aetosaurs and phytosaurs, two Late Triassic groups of crurotarsans with heavy plating, making it one of the first heavily armored archosaurs. Prior to 2017, most studies placed it outside Archosauria as a member of Doswelliidae, a family of heavily armored and crocodile-like archosauriforms. The 2017 specimens instead show that it belonged to the Erpetosuchidae.

<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.

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

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.

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

Mambawakale is a genus of large sized basal paracrocodylomorph, possibly a poposauroid, from the Manda Beds of Tanzania. It was informally named Pallisteria before being officially published under its current name almost 60 years after its discovery. It contains a single species, Mambawakale ruhuhu.

References

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