Siats Temporal range: Cenomanian, | |
---|---|
Skeletal reconstruction of known elements | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Tetanurae |
Clade: | Orionides |
Clade: | Avetheropoda |
Genus: | † Siats Zanno & Makovicky, 2013 |
Type species | |
†Siats meekerorum Zanno & Makovicky, 2013 |
Siats (/see-ats/) is an extinct genus of large theropod dinosaur known from the Late Cretaceous Cedar Mountain Formation of Utah, United States. It contains a single species, Siats meekerorum. [1] [2] It was initially classified as a megaraptoran, a clade of large theropods with very controversial relationships. Siats may be a neovenatorid allosauroid, [1] a coelurosaur of uncertain phylogenetic position, [3] or a tyrannosauroid . [4]
It was discovered in 2012 and named by Lindsay E. Zanno and Peter J. Makovicky in 2013 and the type species is Siats meekerorum. The generic name is derived from the name of Siats, a man-eating monster in the Ute mythology. [1] The specific name meekerorum honors the late geologist John Caldwell Meeker who bequeathed a fund for the support of paleontological research, his widow Withrow Meeker and their daughter Lis Meeker, one of the volunteers in the research project. [1]
Siats is known from the holotype FMNH PR 2716, a partial postcranial skeleton housed at the Field Museum of Natural History, Chicago. FMNH PR 2716 consists of five dorsal and eight caudal vertebrae, a chevron, partial right ilium, ischium and fibula, a partial left tibia, and several right and left pedal phalanges. [1] FMNH PR 2716 was discovered by Lindsay Zanno, as a part of a 2008 expedition of the Field Museum led by Peter Makovicky. [5] The bones were first seen protruding out of a hillside, prompting the excavation. It was collected between 2008 and 2010 from the Mussentuchit Member of the Cedar Mountain Formation, in Emery County of Utah, dating to the early Cenomanian stage of the Late Cretaceous, approximately 94.5 million years ago. [6]
The holotype specimen, FMNH PR 2716, consists of material from a single individual that is considered skeletally immature on the basis of the incomplete fusion of neural arches to the centra of the dorsal vertebrae. Siats is characterized by seven diagnostic, including four autapomorphic (i.e. unique), traits. Its autapomorphies include the subtriangular cross section of the distal caudal vertebrae, elongated centrodiapophyseal laminae lacking noticeable infradiapophyseal fossae on the proximal caudals, a transversely concaved acetabular rim of iliac pubic peduncle, and the presence of a notch on the end of the truncated lateral brevis shelf. Other notable traits include the broad neural spines on the dorsal vertebrae. [1]
Siats represents one of the largest known theropods from its place and time. Zanno and Mackovicky (2013), using a femur circumference regression, estimated its mass at roughly 3.9 metric tons (4.3 short tons). The authors also wrote that the holotype specimen was already comparable in size to Saurophaganax and Acrocanthosaurus despite the visible neurocentral sutures showing absence of ossification, suggesting a skeletally immature individual. [1] In 2019, its body length was estimated up to 11.7 m (38 ft). [7]
If true, the discovery of Siats also reveals that allosauroids did not yield dominance in North America to tyrannosauroids until the late Cretaceous. [8] However, There is substantial disagreement on the classification of this animal as well as megaraptorans, which are usually found to be either neovenatorids or tyrannosauroids, often with both interpretations appearing in the same paper. [9] The placement of Siats within Megaraptora itself has also been a source of controversy, with recent analyses placing it as a non-megaraptoran theropod irrespective of the placement of Megaraptora. [10]
Siats was initially classified as a megaraptoran neovenatorid closely related to Chilantaisaurus upon its discovery in 2013, based on the presence of pronounced centrodiapophyseal laminae bracketed by deep infradiapophyseal fossa on the caudal neural arches, similar to that of the megaraptoran Aerosteon . In a 2014 description of a juvenile Megaraptor specimen, the referral of Siats to Megaraptora was contested, and megaraptorans were found to more likely be tyrannosauroids rather than neovenatorids such as Siats. The paper noted that, although sharing various features with Neovenator , Siats was very different from megaraptorans in the structure of its dorsal vertebrae, ilium, and fibula. [10] A subsequent analysis conducted by Coria and Currie (2016), which even placed megaraptorans as neovenatorids, still placed Siats and Chilantaisaurus as neovenatorids outside of Megaraptora. [11] However, Bell et al. (2016) recovered Siats as a member of Coelurosauria of uncertain phylogenetic placement within this group; the analysis indicated that Siats might either be a relative of ornithomimosaurs, a theropod more closely related to maniraptorans than tyrannosauroids, a basal megaraptoran, or a tyrannosauroid more closely related to tyrannosaurids than Xiongguanlong . [3] Naish and Cau (2022) recovered the taxon as the basalmost Megaraptoran. [12]
The cladogram presented below (created prior to these more recent studies) follows Zanno & Makovicky (2013). [1]
Allosauroidea | |
The cladogram below follows Naish and Cau (2022), who found Siats within Megaraptora as part of Tyrannosauroidea: [12]
Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods.
Proceratosaurus is a genus of carnivorous theropod dinosaur from the Middle Jurassic (Bathonian) of England. It contains a single species. P. bradleyi, known from a mostly complete skull and lower jaws. Proceratosaurus was a small dinosaur, estimated to measure around 3 m (9.8 ft) in length. Its name refers to how it was originally thought to be an ancestor of Ceratosaurus, due to the partially preserved portion of the crest of Proceratosaurus superficially resembling the small crest of Ceratosaurus. Now, however, it is considered a coelurosaur, specifically a member of the family Proceratosauridae, and amongst the earliest known members of the clade Tyrannosauroidea.
Eotyrannus is a genus of tyrannosauroid theropod dinosaur hailing from the Early Cretaceous Wessex Formation beds, included in Wealden Group, located in the southwest coast of the Isle of Wight, United Kingdom. The remains (MIWG1997.550), consisting of assorted skull, axial skeleton and appendicular skeleton elements, from a juvenile or subadult, found in a plant debris clay bed, were described by Hutt et al. in early 2001. The etymology of the generic name refers to the animal's classification as an early tyrannosaur or "tyrant lizard", while the specific name honors the discoverer of the fossil.
Chilantaisaurus is a genus of large theropod dinosaur, possibly a neovenatorid or a primitive coelurosaur, from the Late Cretaceous Ulansuhai Formation of China. The type species, C. tashuikouensis, was described by Hu in 1964.
Megaraptor is a genus of large theropod dinosaur that lived in the ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina, South America. Initially thought to have been a giant dromaeosaur-like coelurosaur, it was classified as a neovenatorid allosauroid in previous phylogenies, but more recent phylogeny and discoveries of related megaraptoran genera has placed it as either a basal tyrannosauroid or a basal coelurosaur with some studies still considering it a neovenatorid.
Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period. By the end of the Cretaceous Period, tyrannosauroids were the dominant large predators in the Northern Hemisphere, culminating in the gigantic Tyrannosaurus. Fossils of tyrannosauroids have been recovered on what are now the continents of North America, Europe and Asia, with fragmentary remains possibly attributable to tyrannosaurs also known from South America and Australia.
Aniksosaurus is a genus of avetheropod dinosaur from what is now Chubut Province, Argentina. It lived during the Cenomanian to Turonian of the Cretaceous period, between 96-91 million years ago. The type species, Aniksosaurus darwini, was formally described from the Bajo Barreal Formation of the Golfo San Jorge Basin by Rubén Dario Martínez and Fernando Emilio Novas in 2006; the name was first coined in 1995 and reported in the literature in 1997. The specific epithet honors Charles Darwin who visited Patagonia in 1832/1833 during the Voyage of the Beagle.
Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. Definitive fossils attributed to the subfamily have only been found in the Late Cretaceous deposits of East Asia, with Kansaignathus as the basalmost member, while numerous taxa from North America previously assigned to the velociraptorines are reclassified within Saurornitholestinae, Dromaeosaurinae or Eudromaeosauria outside these clades. Several teeth that may belong to indeterminate velociraptorines have also been discovered in Germany, dating to the Kimmeridgian stage of the Late Jurassic.
Orkoraptor is a genus of medium-sized megaraptoran theropod dinosaur from the late Cretaceous Period of Argentina. It is known from incomplete fossil remains including parts of the skull, teeth, tail vertebrae, and a partial tibia. The specialized teeth resemble those of some maniraptoriform theropods, namely the deinonychosaurs and compsognathids. This and other anatomical features led the authors who described it to suggest that it was a maniraptoran coelurosaur. However, subsequent studies found it to be a megaraptoran. Found in the Cerro Fortaleza Formation of Southern Patagonia, it is one of the southernmost carnivorous dinosaurs known from South America.
Aerosteon is a genus of megaraptoran dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, which is from the late Campanian. The type and only known species is A. riocoloradensis. Its specific name indicates that its remains were found 1 km north of the Río Colorado, in Mendoza Province, Argentina.
Peloroplites is a monospecific genus of nodosaurid dinosaur from Utah that lived during the Late Cretaceous in what is now the Mussentuchit Member of the Cedar Mountain Formation. The type and only species, Peloroplites cedrimontanus, is known from a partial skull and postcranial skeleton. It was named in 2008 by Kenneth Carpenter and colleagues. Peloroplites was 6 metres long and weighed 2 tonnes, making it one of the largest known nodosaurids, and came from a time when ankylosaurids and nodosaurids were attaining large sizes.
Bahariasauridae is a potential family of averostran theropods that might include a handful of African and South American genera, such as Aoniraptor, Bahariasaurus, Deltadromeus, and Gualicho. The placement of these theropods is controversial, with some studies placing them as basal ceratosaurs possibly related to Noasauridae, others classifying them as megaraptorans, basal neovenatorids, or basal coelurosaurs. There is also a possibility the group might not be monophyletic, as a monograph on the vertebrate diversity in the Kem Kem Beds published in 2020 found Bahariasaurus to be nomen dubium. In the same paper Deltadromeus is classified as an noasaurid, a result also recovered by some previous studies. A 2024 phylogenetic analysis found Aoniraptor, Bahariasaurus, Deltadromeus, and Gualicho to form a monophyletic clade as the sister taxon to Elaphrosaurus near the base of Ceratosauria.
Xiongguanlong is an extinct genus of tyrannosauroid theropod from the Early Cretaceous period of what is now China. The type and only species is X. baimoensis. The generic name comes from Jiayuguan City and the Mandarin word "long" which means dragon. The specific epithet, "baimoensis" is a latinization of the Mandarin word for "white ghost" in reference to one of the geological features of the type locality.
Australovenator is a genus of megaraptoran theropod dinosaur from Cenomanian -age Winton Formation of Australia. It is known from partial cranial and postcranial remains which were described in 2009 by Scott Hocknull and colleagues, although additional descriptions and analyses continue to be published. It is the most complete predatory dinosaur discovered in Australia. It has been suggested that Australovenator is a sister taxon to Fukuiraptor, although some phylogenetic analyses find it to be a more derived member of the Megaraptora, possibly being part of the main Megaraptoridae family itself.
Megaraptora is a clade of carnivorous theropod dinosaurs with controversial relationships to other tetanuran theropods. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods.
Paludititan is a genus of titanosaurian sauropod dinosaur which lived in the area of present Romania during the Late Cretaceous. It existed in the island ecosystem known as Hațeg Island.
Zuolong is an extinct genus of tetanuran theropod from the Late Jurassic period of China. The type and only species is Z. salleei. The generic name of Zuolong is in honor of General Zuo Zōngtáng with the Chinese word "long" which means dragon. The specific epithet "salleei" is in honor of Hilmar Sallee, who funded the expedition which led to the specimen's discovery.
Gualicho is a genus of theropod dinosaur. The type species is Gualicho shinyae. It lived in what is now northern Patagonia, on what was then a South American island continent split off from the supercontinent Gondwana. The fossils were found in the Huincul Formation, dating to the late Cenomanian-early Turonian age of the upper Cretaceous Period, around 91 million years ago.
Phuwiangvenator is an extinct genus of megaraptoran theropod that lived during the Early Cretaceous period in what is now Thailand. It contains only the type species, P. yaemniyomi. The generic name of Phuwiangvenator comes from the Phu Wiang mountains, where the holotype was discovered, and the Latin word "venator" meaning hunter. The specific name, "yaemniyomi", is in honor of Sudham Yaemniyom, who was a historical paleontologist from Thailand and the first person to discover fossils there.
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