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''Siats'' (/see-ats/) is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of large
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
known from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain (Utah), Cedar Mountain in northern Emery Coun ...
of
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, United States. It contains a single species, ''Siats meekerorum''. It was initially classified as a
megaraptora Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n, a clade of large
theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
with very controversial relationships. ''Siats'' may be a
neovenatorid Neovenatoridae is a proposed clade of carcharodontosaurian dinosaurs uniting some primitive members of the group such as ''Neovenator'' with the Megaraptora, a group of theropods with controversial affinities. Other studies recover megaraptorans ...
allosauroid Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among th ...
, a
coelurosaur Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
of uncertain phylogenetic position, or a
tyrannosauroid Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent b ...
.


Discovery

''Siats'' was first described in
2012 File:2012 Events Collage V3.png, From left, clockwise: The passenger cruise ship Costa Concordia lies capsized after the Costa Concordia disaster; Damage to Casino Pier in Seaside Heights, New Jersey as a result of Hurricane Sandy; People gather ...
and named by
Lindsay E. Zanno Lindsay E. Zanno is an American vertebrate paleontology, paleontologist and a leading expert on Theropoda, theropod dinosaurs and Cretaceous paleoecosystems. She is the Head of Paleontology at the North Carolina Museum of Natural Sciences and an ...
and Peter J. Makovicky in
2013 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment fact ...
and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
is ''Siats meekerorum''. The generic name is derived from the name of ''Siats'', a man-eating
monster A monster is a type of fictional creature found in horror, fantasy, science fiction, folklore, mythology and religion. Monsters are very often depicted as dangerous and aggressive with a strange, grotesque appearance that causes terror and fe ...
in the Ute mythology. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
''meekerorum'' honors the late
geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althou ...
John Caldwell Meeker who bequeathed a fund for the support of
paleontological Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (geology), epoch (roughly 11,700 years before present). It includes t ...
research, his widow Withrow Meeker and their daughter Lis Meeker, one of the volunteers in the research project. ''Siats'' is known from the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
FMNH PR 2716, a partial
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
housed at the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educational ...
,
Chicago (''City in a Garden''); I Will , image_map = , map_caption = Interactive Map of Chicago , coordinates = , coordinates_footnotes = , subdivision_type = Country , subdivision_name ...
. FMNH PR 2716 consists of five
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...
and eight
caudal vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
, a
chevron Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * ''Eulithis testata'', a moth * Chevron (geology), a fold in rock lay ...
, partial right ilium,
ischium The ischium () form ...
and
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
, a partial left
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, and several right and left pedal
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
. FMNH PR 2716 was discovered by Lindsay Zanno, as a part of a 2008 expedition of the Field Museum led by Peter Makovicky. The bones were first seen protruding out of a hillside, prompting the excavation. It was collected between 2008 and 2010 from the
Mussentuchit Member The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain (Utah), Cedar Mountain in northern Emery Coun ...
of the
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain (Utah), Cedar Mountain in northern Emery Coun ...
, in
Emery County Emery County is a county in east-central Utah, United States. As of the 2010 United States Census, the population was 10,976. Its county seat is Castle Dale, and the largest city is Huntington. History Prehistory Occupation of the San Rafael ...
of Utah, dating to the early
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stage of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
, approximately 94.5
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
.


Description

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 In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
(i.e. unique), traits. Its autapomorphies include the subtriangular
cross section Cross section may refer to: * Cross section (geometry) ** Cross-sectional views in architecture & engineering 3D *Cross section (geology) * Cross section (electronics) * Radar cross section, measure of detectability * Cross section (physics) **Abs ...
of the
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
caudal vertebrae, elongated centrodiapophyseal laminae lacking noticeable infradiapophyseal fossae on the
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
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 The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
on the dorsal vertebrae. ''Siats'' represents one of the largest known theropods from North America. Zanno and Mackovicky (2013), using a femur circumference regression, estimated its mass at roughly . The authors also wrote that the holotype specimen was already comparable in size to ''
Saurophaganax ''Saurophaganax'' ("lord of lizard-eaters") is a genus of large allosaurid dinosaur from the Morrison Formation of Late Jurassic (latest Kimmeridgian age, about 151 million years ago) Oklahoma, United States.Turner, C.E. and Peterson, F., (1999) ...
'' and ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ' ...
'' despite the visible neurocentral sutures showing absence of ossification, suggesting a skeletally immature individual. In 2019, its maximum adult size was estimated up to in length and in body mass. If it is a neovenatorid, the discovery of ''Siats'' also reveals that
allosauroid Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among th ...
s did not yield dominance in North America to
tyrannosauroid Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent b ...
s until the late Cretaceous. However, There has been and still is substantial disagreement on the classification of this animal as well as with megaraptorans, which are usually found to be either neovenatorids or tyrannosauroids, often with both interpretations appearing in the same paper. The placement of ''Siats'' within Megaraptora itself has also been a source of controversy, with recent analyses placing it as a non-megaraptoran neovenatorid irrespective of the placement of Megaraptora.


Phylogeny and classification

''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 ''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. riocolorade ...
''. 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 ''Neovenator'' (nee-o-ven-a-tor meaning "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known from several skeletons found in the Early Cretaceous (Barremian~130-125 million years ago) Wessex Formation on the south coa ...
'', ''Siats'' was very different from megaraptorans in the structure of its dorsal vertebrae, ilium, and fibula. A subsequent analysis conducted by Coria and
Currie Currie ( gd, Currach, IPA: kʰuːᵲəx is a village and suburb on the outskirts of Edinburgh, Scotland, situated south west of the city centre. Formerly within the County of Midlothian, it now falls within the jurisdiction of the City of Edi ...
(2016), which even placed megaraptorans as neovenatorids, still placed ''Siats'' and ''Chilantaisaurus'' as neovenatorids outside of Megaraptora. However, Bell ''et al.'' (2016) recovered ''Siats'' as a member of
Coelurosauria Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
of uncertain phylogenetic placement within this group; the analysis indicated that ''Siats'' might either be a relative of
ornithomimosaurs Ornithomimosauria ("bird-mimic lizards") are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia (now Asia, Europe and North ...
, a theropod more closely related to
maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
ns than
tyrannosauroids Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent be ...
, a basal megaraptoran, or a tyrannosauroid more closely related to
tyrannosaurids Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family (biology), family of coelurosaurian Theropoda, theropod dinosaurs that comprises two subfamilies containing up to thirteen genus, genera, including the eponymous ''Tyrannos ...
than ''
Xiongguanlong ''Xiongguanlong'' ("Grand Pass dragon") is a genus of tyrannosauroid dinosaur that lived in the Early Cretaceous of what is now China. The type species is ''X. baimoensis'', described online in 2009 by a group of researchers from China and the Un ...
''. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
presented below (created prior to these more recent studies) follows Zanno & Makovicky (2013).


References

{{Taxonbar, from=Q15221367 Prehistoric tetanurans Late Cretaceous dinosaurs of North America Fossil taxa described in 2013 Taxa named by Lindsay Zanno Paleontology in Utah