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''Raptorex'' is a
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
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 ...
. Its fossil remains consist of a single juvenile specimen probably uncovered in
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
, or possibly northeastern China. 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 specime ...
is ''R. kriegsteini'', described in 2009 by Sereno and colleagues. The genus name is derived from
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
', "robber", and ', "king". The specific name honours Roman Kriegstein, a survivor of the
Holocaust The Holocaust, also known as the Shoah, was the genocide of European Jews during World War II. Between 1941 and 1945, Nazi Germany and its collaborators systematically murdered some six million Jews across German-occupied Europe; ...
, whose son Henry Kriegstein donated the specimen to the
University of Chicago The University of Chicago (UChicago, Chicago, U of C, or UChi) is a private university, private research university in Chicago, Illinois. Its main campus is located in Chicago's Hyde Park, Chicago, Hyde Park neighborhood. The University of Chic ...
for scientific study. While initially considered to have come from the
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its exq ...
of China, dated to approximately 125 million years ago during the early
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period, later studies showed that such an early date for the fossil are unlikely, and given its extremely close similarity to juvenile tyrannosaurids of the late Cretaceous, it probably came from the Nemegt or similar formation. Because the specimen is a juvenile, and the changes undergone by tyrannosaurids during growth are not yet well understood, many researchers now consider it to be a ''nomen dubium'', because it cannot be confidently paired with an adult skeleton (though it is extremely similar to juvenile ''
Tarbosaurus bataar ''Tarbosaurus'' ( ; meaning "alarming lizard") is a genus of tyrannosaurid dinosaur that flourished in Asia about 70 million years ago, at the end of the Late Cretaceous Period (geology), Period, considered to contain a single known species, ' ...
'' skeletons of the same size and age).


Description

The only known ''Raptorex'' specimen shows the same basic proportions as juvenile tyrannosaurids: a comparatively large and solidly-constructed skull, long legs with adaptations for running, and tiny, two-fingered forelimbs. This is in contrast with more basal tyrannosauroids such as ''
Dilong ''Dilong'' (; lit. "earth dragon") is a Chinese dragon name that is also used to mean "earthworm" in traditional Chinese medicine and '' Geosaurus'' in zoological nomenclature. Dragon In Chinese mythology, ''dilong'' "earth dragon" is one of ...
'', which retained features characteristic of more basal
coelurosaurs 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, tyran ...
such as a small head and long, three-fingered forelimbs. The specimen is a very small juvenile, estimated at long and about . 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 sever ...
(LH PV18) measured about and died in its third year.


History


Discovery

According to Peter Larson, who attempted to re-trace the origins of the specimen, the holotype fossil of ''Raptorex'' (currently designated LH PV18 and housed in the collections of the Long Hao Institute of Geology and Paleontology in Hohhot, Inner Mongolia, China) was purchased from a Mongolian fossil dealer by an American businessman in Tokyo, Japan, and subsequently taken to the United States, where it was again put up for sale at the Tucson, Arizona Gem, Mineral and Fossil Show. There it was sold by Hollis Butts to Dr. Henry Kriegstein, an ophthalmologist and fossil collector. Up until this point, the specimen had been identified as a juvenile specimen of ''
Tarbosaurus ''Tarbosaurus'' ( ; meaning "alarming lizard") is a genus of tyrannosaurid dinosaur that flourished in Asia about 70 million years ago, at the end of the Late Cretaceous Period, considered to contain a single known species, ''Tarbosaurus ba ...
'', which had been collected from Mongolia. Kriegstein notified American paleontologist
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
of the specimen, who proposed it represented a subadult of a new species from the Yixian Formation of China. Sereno arranged to publish a description of the specimen and to have it sent to China, from where he assumed it had been smuggled.


Initial interpretation

In Sereno's description of the specimen, he and his co-authors interpreted it as a near-adult aged about six years. Sereno initially stated to the press that numerous
index fossil Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...
s present in the rock slab surrounding the specimen showed it belonged to Early Cretaceous. In an interview with the ''Chicago Tribune'', Sereno said “From sediments, fossil fish bones, turtles, clam shells and other fauna we recovered from the rock matrix alongside the ''Raptorex'' fossil, we could generally pinpoint where it had been dug up in an area along the border with Inner Mongolia."Mullen W (2009) Fossil identified as mini-T. rex. Chicago Tribune, Sept 18th 2009. Available: http://articles.chicagotribune.com/2009-09-18/news/0909170802_1_fossil-bones-tyrannosaurus-rex-chicago-paleontologist-paul-sereno. Accessed 2011 March 8. However, only a single fish vertebra and a crushed, unidentifiable mollusk shell were actually present alongside the ''Raptorex'' specimen. Sereno and colleagues identified the fish vertebra as being similar to the genus ''
Lycoptera ''Lycoptera'' is an extinct genus of fish that lived from the late Jurassic to Cretaceous periods in present-day China, North Korea, Mongolia and Siberia. It is known from abundant fossils representing sixteen species, which serve as important ind ...
'', a key index fossil of the early Cretaceous Yixian formation. However, they did not describe the bone or note any features in common with known ''Lycoptera'' specimens. Sereno's interpretation of the specimen as an early, primitive, non-tyrannosaurid
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 ...
would have major implications for the evolution of tyrannosaurs. Rather than evolving their distinct anatomy of a large head, long legs and tiny, two-fingered arms only after large body size, a small, early Cretaceous species with similar proportions to adults of true tyrannosaurids would indicate that the characteristic tyrannosaur shape appeared prior to the advent of giant size in the group. This would contradict previous evidence, as all previously known primitive tyrannosaurs had small skulls and long, three-fingered arms, including contemporary species and those that lived later in the Cretaceous than Sereno believed ''Raptorex'' to have lived.


Controversy and re-interpretation

In October 2010 an online ''Nature News'' report (not peer-reviewed) challenged the provenance and classification of ''Raptorex'' as a basal tyrannosauroid.
Peter Larson Peter Lars Larson (born 1952) is an American fossil expert and president of the Black Hills Institute of Geological Research. He led the team that excavated " Sue", one of the largest and most complete specimen of ''Tyrannosaurus rex'' found to ...
, president of the Black Hills Institute of Geological Research, Inc., a private fossil excavation and supply company, inspected the fossil and told ''Nature'' that he concluded it was a juvenile ''
Tarbosaurus ''Tarbosaurus'' ( ; meaning "alarming lizard") is a genus of tyrannosaurid dinosaur that flourished in Asia about 70 million years ago, at the end of the Late Cretaceous Period, considered to contain a single known species, ''Tarbosaurus ba ...
''. Because the specimen was donated by a collector without detailed provenance information, Larson doubted the assigned age, which was based only on the ''Lycoptera'' vertebra and mollusk shell found alongside the dinosaur fossil. Larson speculated that the fossil could have come from beds in Mongolia that yield fossils of ''Tarbosaurus'', dating from 70 million years ago. He has suggested "a more detailed analysis of the fossil matrix—including dating of any pollen associated with the fossil." Sereno is quoted in the report as standing by his conclusions, noting that definite evidence or a publication had not been produced refuting them. In June 2011, a more detailed re-study was published in the peer-reviewed journal '' PLoS ONE'' by Denver Fowler, Peter Larson and others reanalyzing the published data, which they stated to be equivocal regarding stratigraphic position, and that ontogenetic interpretation in the description had overestimated the maturity of the specimen. While Sereno had claimed that the vertebrae of ''Raptorex'' were nearly fused, and that the bone histology of the specimen indicated it was a subadult about six years old, Fowler and colleagues argued that Sereno and his team had misinterpreted the growth stage data, and found that the specimen was actually a young juvenile only about three years old. They also found fault with the Sereno team's interpretation of the specimen's age. Fowler and colleagues showed that the fish bone which Sereno had identified as ''Lycoptera'' without comment is actually very different in shape and much larger in size than any known specimen of ''Lycoptera'' and cannot even be assigned to the same order as that genus. Rather, it probably belonged to an ellimmichthyiform fish, which span the entire Cretaceous period, rendering the bone useless for dating. In light of this, they noted that there is no reason to believe the fossil dates to the early Cretaceous, and that given its extreme similarity to juvenile tyrannosaurids, a late Cretaceous age is far more likely. Based on this analysis, Fowler and colleagues concluded that ''Raptorex'' was much more likely to represent a juvenile tyrannosaurid similar to ''Tarbosaurus'', though its exact identity cannot be known without more information about growth patterns in tyrannosaurids, and further efforts to discover its age. Consequently, Sereno's hypothesis that the derived features of tyrannosaurids evolved in the Early Cretaceous cannot be supported by current evidence. Newbrey ''et al.'' (2013) identified the fish vertebra centrum found with the holotype specimen of ''Raptorex kriegsteini'' as belonging to a hiodontid, probably representing the same taxon as hiodontid fossils described by the authors from the Late Cretaceous
Nemegt Formation The Nemegt Formation (also known as Nemegtskaya Svita) is a geological formation in the Gobi Desert of Mongolia, dating to the Late Cretaceous. The formation consists of river channel sediments and contains fossils of fish, turtles, crocodilians, ...
of
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
. According to Newbrey ''et al.'', centra like those of the Nemegt hiodontid are not known from outside the Nemegt Formation; according to the authors, this implies that the vertebra found with the holotype specimen of ''Raptorex kriegsteini'' – and by extension the holotype of ''R. kriegsteini'' itself – most likely come from the Nemegt Formation and are Late Cretaceous in age. Kim ''et al.'' (2022) compared a fish centrum found with 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 sever ...
of ''Raptorex'' with '' Harenaichthys lui'' from the Nemegt Formation and Chinese '' Xixiaichthys tongxinensis'', and interpreted their findings as supporting the conclusion that the holotype of ''R. kriegsteini'' comes from the Nemegt Formation.


Classification

''Raptorex'' has been interpreted as a tyrannosaurid, tyrannosauroid and a synonym of ''Tarbosaurus''. It is currently placed as a basal tyrannosauroid more derived than ''
Bagaraatan ''Bagaraatan'' (/'ba-ɣa-raa-tan/ meaning 'small' ''baɣa'' + 'carnivorous animal, beast of prey' ''araatan'' in Mongolian) is a genus of theropod dinosaur from the Late Cretaceous period. Its fossils were found in the Nemegt Formation of Mongoli ...
'' but more basal than ''
Dryptosaurus ''Dryptosaurus'' ( ) is a genus of tyrannosauroid that lived approximately 67 million years ago (mya) during the latter part of the Cretaceous period, New Jersey. ''Dryptosaurus'' was a large, bipedal, ground-dwelling carnivore, that grow up to ...
''. Below is a cladogram placing ''Raptorex'' as a basal tyrannosauroid. It was published by Loewen ''et al.'' in 2013 on PLoS ONE.


See also

*
Timeline of tyrannosaur research This timeline of tyrannosaur research is a chronological listing of events in the history of paleontology focused on the tyrannosaurs, a group of predatory theropod dinosaurs that began as small, long-armed bird-like creatures with elaborate cr ...


References


External links


Science Daily: Tiny Tyrannosaur: T. Rex Body Plan Debuted In Raptorex, But 100th The Size
{{Taxonbar, from=Q131717 Late Cretaceous dinosaurs of Asia Nemegt fauna Tyrannosaurids Fossil taxa described in 2009 Nomina dubia Taxa named by Paul Sereno Taxa named by Stephen L. Brusatte Controversial dinosaur taxa