Shartegosuchidae

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Shartegosuchidae
Temporal range: Late Jurassic - Early Cretaceous
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Crocodylomorpha
Clade: Solidocrania
Clade: Crocodyliformes
Clade: Mesoeucrocodylia
Clade: Shartegosuchoidea
Family: Shartegosuchidae
Efimov, 1988
Genera

Shartegosuchidae is an extinct family of Late Jurassic and Early Cretaceous crocodyliforms. The family is named after the Late Jurassic Shar Teeg Beds in southwestern Mongolia, from which most shartegosuchid remains have been found. Five genera are currently assigned to Shartegosuchidae: Shartegosuchus , Nominosuchus , Kyasuchus , Adzhosuchus , and Fruitachampsa . Shartegosuchus, Nominosuchus, and Adzhosuchus all come from Shar Teeg, while Kyasuchus is known from the Early Cretaceous of Russia. Fruitachampsa is known from the Late Jurassic Morrison Formation of the western United States. [1]

Contents

Description

Shartegosuchids are known only from cranial material, or parts of the skull and some partial skeletons. Shartegosuchids share several distinct characteristics in the skull. On the palate, the choanae (holes where the nasal cavity opens into the mouth) is placed within a deep depression of its midline. The palatal bones, located behind the choanae in what is known as the secondary palate, unite with each other at the midline of the palate. Shartegosuchids can also be diagnosed by the position of the teeth in their lower jaws, which are never found behind the mandibular fenestrae (holes found near the back of the jaw). The edges of the teeth are denticulated, or ridged. The shape and position of several bones of the skull, including the frontal, nasal, lacrimal, and quadrate, are also distinctive. Unlike most other archosaurs, shartegosuchids lack an antorbital fenestra, an opening in the skull in front of the eyes. [2]

Classification

Shartegosuchids are considered to be basal crocodyliforms, but their classification within the group is still debated. At various times since the construction of the family, shartegosuchids have been considered to be protosuchians, mesosuchians, and perhaps notosuchians or atoposaurids. [3]

Fruitachampsa , a crocodylomorph from the Morrison Formation in Colorado, is also referable to Shartegosuchidae. Before it was formally named in 2011, it was commonly known as the "Fruita form". The known material of Fruitachampsa consists of several nearly complete skeletons. [4] It was first classified in the suborder Mesosuchia, which has since been replaced by the clade Mesoeucrocodylia. However, Fruitachampsa can be linked to shartegosuchids on the basis of similarities in the structure of the front of the palate and the choana. [5]

In a 2004 phylogenetic study, Fruitachampsa, along with Gobiosuchus and Zosuchus , was found to be outside Mesoeucrocodylia, but still more derived than Protosuchidae, a family that is often thought to be closely related to shartegosuchids. [5]

A more recent study in 2006 found Shartegosuchidae to be the most basal clade of mesosuchians. A phylogenetic analysis found the family to be monophyletic, meaning that it forms a true clade with a common ancestor from which only shartegosuchids are derived. "Fruitachampsa" was found to be outside Shartegosuchidae, but was the sister taxon of the family. [2] This means that it is more closely related to Shartegosuchidae than any other crocodyliform.

Below is a cladogram based on the 2006 study, showing the relationships of members of the family: [2]

Mesoeucrocodylia  

Metasuchia

Hsisosuchus

Shartegosuchidae

Nominosuchus matutinus

Nominosuchus arcanus

Kyasuchus

Shartegosuchus

Adzhosuchus

Below is a cladogram showing the phylogenetic placement of Shartegosuchidae from Clark (2011): [1]

Crocodyliformes  

Protosuchidae

Gobiosuchus

Zaraasuchus

Zosuchus

Sichuanosuchus

Shantungosuchus

Shartegosuchidae

Nominosuchus

Fruitachampsa

Shartegosuchus

Adzhosuchus

Hsisosuchus

Mesoeucrocodylia

Paleobiology

The shartegosuchids from Mongolia and Russia comprise a group of crocodyliforms that was endemic to Central Asia during the Mesozoic. These crocodyliforms, along with other Asiatic tetrapods, make up a fauna that may be unique to the region due to geographic isolation that existed since the Late Triassic or Early Jurassic. This isolation first occurred when the supercontinent Pangaea began to break apart, forming the smaller northern supercontinent of Laurasia. [6] [7] The authors of a 2007 study of early crocodylomorphs did not include shartegosuchids in a phylogenetic analysis, indicating that they were so highly endemic to Asia that their relationship to other crocodylomorphs would be difficult to interpret. [6]

Related Research Articles

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

Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day.

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

Comahuesuchus is an extinct genus of notosuchian crocodylomorphs from the late Cretaceous of Argentina. It was described by palaeontologist José Bonaparte in 1991. The type species is C. brachybuccalis from the Santonian Bajo de la Carpa Formation. In 2023 another species, C. bonapartei from the Sierra Barrosa Formation and Portezuelo Formation, was assigned to the genus.

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

Sichuanosuchus is an extinct genus of crocodyliform from the Late Jurassic and possibly Early Cretaceous of China.

<span class="mw-page-title-main">Notosuchia</span> Extinct suborder of reptiles

Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group ; if Sebecosuchia is included within Notosuchia its existence is pushed into the Middle Miocene, about 11 million years ago. Fossils have been found from South America, Africa, Asia, and Europe. Notosuchia was a clade of terrestrial crocodilians that evolved a range of feeding behaviours, including herbivory (Chimaerasuchus), omnivory (Simosuchus), and terrestrial hypercarnivory (Baurusuchus). It included many members with highly derived traits unusual for crocodylomorphs, including mammal-like teeth, flexible bands of shield-like body armor similar to those of armadillos (Armadillosuchus), and possibly fleshy cheeks and pig-like snouts (Notosuchus). The suborder was first named in 1971 by Zulma Gasparini and has since undergone many phylogenetic revisions.

Neuquensuchus is an extinct genus of crocodyliform from the Santonian-age Upper Cretaceous Bajo de la Carpa Formation of Neuquén Province, Argentina. The known remains were discovered on the campus of Universidad Nacional del Comahue in the city of Neuquén. Neuquensuchus was named by Lucas E. Fiorelli and Jorge O. Calvo in a publication dated to 2007, but which became available in 2008. The type species is N. universitas, in recognition of its discovery locality. Unlike the great majority of crocodyliforms, its shin was longer than its thigh, suggesting it had some running ability.

Calsoyasuchus is a genus of crocodylomorph that lived in the Early Jurassic. Its fossilized remains were found in the Sinemurian-Pliensbachian-age Kayenta Formation on Navajo Nation land in Coconino County, Arizona, United States. Formally described as C. valliceps, it is known from a single incomplete skull which is unusually derived for such an early crocodile relative. This genus was described in 2002 by Ronald Tykoski and colleagues; the specific name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones. It has often been interpreted as the earliest diverging member of Goniopholididae, but other studies have recovered it in various other positions.

<span class="mw-page-title-main">Protosuchia</span> Extinct informal group of reptiles

Protosuchia is a group of extinct Mesozoic crocodyliforms. They were small in size and terrestrial. In phylogenetic terms, Protosuchia is considered an informal group because it is a grade of basal crocodyliforms, not a true clade.

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

Eutretauranosuchus is an extinct genus of goniopholidid crocodyliform. E. delfsi is the only known species within the genus.

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

Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to Crocodylus niloticus than to Notosuchus terrestris. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages in North America. It is often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.

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

Fruitachampsa is a genus of shartegosuchid crocodyliform from the Upper Jurassic Morrison Formation of Colorado. It is known from multiple specimens that show it to have been a relatively long-limbed terrestrial quadrupedal predator less than 1 metre (3.3 ft) long, with a short face and a prominent pair of canine-like teeth in the lower jaw. Before it was formally described in 2011, it was also known as the "Fruita form". Its type species is F. callisoni.

Nominosuchus is a genus of protosuchian-grade crocodyliform. It is known from several specimens discovered in ancient lake deposits of the Tithonian-age Upper Jurassic Tsagaantsav Formation, southwestern Mongolia. The type specimen is PIN 4174/4, a partial skull. Nominosuchus was not large; its skull length is estimated at 60 millimetres (2.4 in). It was similar to Shartegosuchus, and is assigned to the same family (Shartegosuchidae). Nominosuchus was described in 1996 by Mikhail Efimov, and the type species is N. matutinus.

Shartegosuchus is a genus of protosuchian-grade crocodyliform. It is known primarily from PIN 4174/2, the partial deformed skull and jaws of a juvenile. This specimen was discovered in ancient lake deposits of the Tithonian-age Upper Jurassic Tsagaantsav Formation, southwestern Mongolia. The estimated length of the holotype skull is 40 millimetres (1.6 in). This genus was similar to Nominosuchus, and both are assigned to the same family (Shartegosuchidae). Shartegosuchus was described in 1988 by Mikhail Efimov, and the type species is S. asperopalatum.

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

Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudosuchians to survive the K-Pg extinction event.

Kyasuchus is an extinct genus of shartegosuchid crocodyliform. Fossils have been found from the Ilek Formation outcropping in the Kemerovo Oblast of Russia, deposited during the Aptian and Albian stages of the Early Cretaceous. The localities from which specimens of this genus have been found have also yielded many other vertebrate remains such as those of palaeonisciform fishes, turtles, various lizards, troodontids, triconodonts, the ceratopsian Psittacosaurus, and the protosuchian-grade crocodylomorph Tagarosuchus.

The Ulan Malgait Formation is a Late Jurassic geologic formation in Mongolia. Dinosaur remains are among the fossils that have been recovered from the formation, although as of 2004 none have yet been referred to a specific genus.

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

Platyognathus is an extinct genus of protosuchian crocodyliform. Fossils are known from the Early Jurassic Lower Lufeng Formation in Yunnan, China and belong to the type and only species, P. hsui.

Shantungosuchus is an extinct genus of Early Cretaceous crocodyliform found in China. It includes three species: Shantungosuchus chuhsienensis and S. brachycephalus, which were both described by Yang Zhongjian – usually referred to as "Young" – in 1961 and 1982, and S. hangjinensis, which was described by Xiao-Chun Wu et al in 1994. S. chuhsienensis is the type for this genus.

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

Susisuchus is an extinct genus of neosuchian mesoeucrocodylian crocodyliform from the Early Cretaceous of Brazil. Fossils have been found from the Nova Olinda Member of the Aptian-age Crato Formation in the Araripe and Lima Campos Basins of northeastern Brazil. Named in 2003, Susisuchus is the sole member of the family Susisuchidae, and is closely related to the clade Eusuchia, which includes living crocodilians. The type species is S. anatoceps, known from a single partial articulated skeleton that preserves some soft tissue. A second species, S. jaguaribensis, was named in 2009 from fragmentary remains.

Tagarosuchus is an extinct genus of Early Cretaceous protosuchian-grade crocodyliform. Fossils of Tagarosuchus have been found from southern Siberia, including a nearly complete skull found near the village of Shestakovo in Kemerovo Oblast. Tagarosuchus was named in 1999, with the type species being T. kulemzini. Remains have been recovered from the Aptian-Albian Ilek Formation.

Almadasuchus is an extinct genus of crocodylomorph known from the early Late Jurassic Puesto Almada Member of the e Cañadón Calcáreo Formation of Patagonia, Argentina. It contains a single species, Almadasuchus figarii. It is known from the holotype MPEF-PV 3838, a well-preserved posterior region of the skull as well as other skull and postcranial remains. Almadasuchus was recovered from Puesto Almada, 30 m above the fish beds, dated as Oxfordian in age.

References

  1. 1 2 Clark, James M. (2011). "A new shartegosuchid crocodyliform from the Upper Jurassic Morrison Formation of western Colorado". Zoological Journal of the Linnean Society. 163 (s1): S152–S172. doi: 10.1111/j.1096-3642.2011.00719.x .
  2. 1 2 3 Fiorelli, L.E.; Juárez Valiera, R.D.; Salinas, G.C. (2006). "Relaciones filogenéticas de "Shartegosuchidae" Efimov (Crocodyliformes: Mesoeucrocodylia) del Jurásico y Cretácico de Asia Central" (PDF). Congreso Argentino de Paleontología y Biostratigrafía. Vol. 9. Córdoba: Resúmenes. p. 83. Archived from the original (PDF) on 2012-03-09.
  3. Storrs, G.W.; Efimov, M.B. (2003). "Mesozoic crocodyliforms of north-central Eurasia". In Benton, M.J.; Shishkin, M.A.; Unwin, D.M. (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge: Cambridge University Press. pp. 402–419. ISBN   9780521545822.
  4. Foster, J. (2007). "Chapter Five: Fins, Scales, and Wings". Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Bloomington, Indiana: Indiana University Press. pp. 153–154. ISBN   978-0-253-34870-8.
  5. 1 2 Pol, D.; Norell, M.A. (2004). "A new crocodyliform from Zos Canyon, Mongolia" (PDF). American Museum Novitates (3445): 1–36. doi:10.1206/0003-0082(2004)445<0001:ancfcm>2.0.co;2. S2CID   86287623.
  6. 1 2 Fiorelli, L.E.; Calvo, J.O. (2007). "The first "protosuchian" (Archosauria: Crocodyliformes) from the Cretaceous (Santonian) of Gondwana" (PDF). Arquivos do Museu Nacional, Rio de Janeiro. 65 (4): 417–459. ISSN   0365-4508. Archived from the original (PDF) on 2012-03-09.
  7. Russell, D.A. (1993). "The role of Central Asia in dinosaurian biogeography". Canadian Journal of Earth Sciences. 30 (10): 2002–2012. Bibcode:1993CaJES..30.2002R. doi:10.1139/e93-176.