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''Tarbosaurus'' ( ; meaning "alarming lizard") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of large
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen 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 Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived in
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
epoch, about 70
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
(
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age Age or AGE may refer to: Time and its effects * Age, the amount of time someone has been alive or something has existed ** East Asian age reckoning, an Asian system of marking age starting at 1 * Ageing or aging, the process of becoming older ...
). It contains the single
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
: ''Tarbosaurus bataar'', which is known from the
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 is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
, with more fragmentary remains found further afield in the
Subashi Formation The Subashi Formation () is a Late Cretaceous (Campanian to Maastrichtian)Xi et al., 2018 formation from the Xinjiang Autonomous Region of western China.China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. ''Tarbosaurus'' is represented by dozens of
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
specimens, including several complete skulls and skeletons. These remains have allowed studies focusing on its
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
, skull mechanics, and brain structure. Further fossil remains have been reported from other
geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
s of Asia, however, these remains are fragmentary and cannot be confidently assigned to ''Tarbosaurus'' or the type species. Like most known tyrannosaurids, ''Tarbosaurus'' was a large
biped Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ' ...
al
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
, with the type specimen measuring approximately long, tall at the hips, and weighing up to . It had a unique locking mechanism in its jaw, equipped with about sixty large teeth, and the smallest arms relative to body size of all tyrannosaurids, renowned for their disproportionately tiny, two-fingered hands. Although many
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
have been named, modern paleontologists recognize only one species, ''T. bataar''. Some experts see this species as an Asian representative of the
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
n genus ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
'', which would make the genus ''Tarbosaurus'' redundant. ''Tarbosaurus'' and ''Tyrannosaurus'', if not
synonymous A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
, are considered to be very closely related genera. ''
Alioramus ''Alioramus'' (; meaning 'different branch') is a genus of tyrannosaurid theropod dinosaurs from the Late Cretaceous period of Asia. It currently contains two species. The type species, ''A. remotus'' is known from a partial skull and three foo ...
'', also from Mongolia, has previously been thought by some authorities to be the closest relative of ''Tarbosaurus'', though this has since been disproven with the discovery of ''
Qianzhousaurus ''Qianzhousaurus'' (meaning "Qianzhou lizard") is a genus of tyrannosaurid dinosaur that lived in Asia during the Maastrichtian age of the Late Cretaceous period. There is currently only one species named, the type species ''Qianzhousaurus sinens ...
'' and the description of the tyrannosaurine tribe
Alioramini Alioramini is a clade of long-snouted tyrannosaurine tyrannosaurid dinosaurs from the Late Cretaceous epoch. It includes the genera '' Alioramus'' and '' Qianzhousaurus''. Although tyrannosaurids are known from a variety of places around the glo ...
. ''Tarbosaurus'' lived in a humid
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
dominated by deserts, forests and plains, and criss-crossed by river channels. In this environment, it was an
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
preying on other large dinosaurs, like
ankylosaurids Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
, such as ''
Tarchia ''Tarchia'' (meaning "brainy one") is a genus of herbivorous ankylosauridae, ankylosaurid dinosaur from the late Cretaceous of Mongolia. Discovery and naming In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. ...
'' and ''
Saichania ''Saichania'' (Mongolian meaning "beautiful one") is a genus of herbivorous ankylosaurid dinosaur from the Late Cretaceous period of Mongolia. The first fossils of ''Saichania'' were found in the early 1970s in Mongolia. In 1977 the type speci ...
'',
hadrosaurids Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
, such as ''
Saurolophus ''Saurolophus'' (; meaning "lizard crest") is a genus of large hadrosaurid dinosaur from the Late Cretaceous period of Asia and North America, that lived in what is now the Horseshoe Canyon and Nemegt formations about 70 million to 66 million ...
'' and ''
Barsboldia ''Barsboldia'' (meaning "of Rinchen Barsbold, Barsbold", a well-known Mongolian people, Mongolian paleontologist) is a genus of large hadrosaurid dinosaur from the Maastrichtian, early Maastrichtian Nemegt Formation of Ömnögovi Province, Ömno ...
'', and
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
, such as ''
Nemegtosaurus ''Nemegtosaurus'' (meaning 'Reptile from the Nemegt') was a sauropod dinosaur from the Late Cretaceous Period of what is now Mongolia. Measuring an estimated long and weighing , it was named after the Nemegt Basin in the Gobi Desert, where the ...
'' and ''
Opisthocoelicaudia ''Opisthocoelicaudia'' is a genus of sauropod dinosaur of the Late Cretaceous Period discovered in the Gobi Desert of Mongolia. The type species is ''Opisthocoelicaudia skarzynskii''. A well-preserved skeleton lacking only the head and neck wa ...
''.


Discovery and naming

In 1946, a joint
Soviet The Union of Soviet Socialist Republics. (USSR), commonly known as the Soviet Union, was a List of former transcontinental countries#Since 1700, transcontinental country that spanned much of Eurasia from 1922 until Dissolution of the Soviet ...
-
Mongolian Mongolian may refer to: * Something of, from, or related to Mongolia, a country in Asia * Mongolian people, or Mongols * Bogd Khanate of Mongolia, the government of Mongolia, 1911–1919 and 1921–1924 * Mongolian language * Mongolian alphabet * ...
expedition to the
Gobi Desert The Gobi Desert (, , ; ) is a large, cold desert and grassland region in North China and southern Mongolia. It is the sixth-largest desert in the world. The name of the desert comes from the Mongolian word ''gobi'', used to refer to all of th ...
in the
Ömnögovi Province Ömnögovi ( ''Ömnögovǐ'', ''South Gobi'') is an aimag (province) of Mongolia, located in the south of the country, in the Gobi Desert. Ömnögovi is Mongolia's largest aimag. The capital is Dalanzadgad. The province is rich in mineral dep ...
turned up a large theropod skull and some
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e from the
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 ...
. In 1955,
Evgeny Maleev Evgeny Aleksandrovich Maleev (, ; 25 February 1915 – 12 April 1966) was a Soviet and Russian paleontologist who did most of his research on reptiles and Asian fossils, such as the naming of the ankylosaur '' Talarurus'' and theropods '' Tar ...
, a Soviet
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
, made this specimen the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(
PIN A pin is a device, typically pointed, used for fastening objects or fabrics together. Pins can have the following sorts of body: *a shaft of a rigid inflexible material meant to be inserted in a slot, groove, or hole (as with pivots, hinges, an ...
551–1) of a new species, which he called ''Tyrannosaurus bataar''. The specific name is a misspelling of the Mongolian word баатар/''baatar'', meaning "hero". In the same year, Maleev also described and named three new theropod skulls, each associated with skeletal remains discovered by the same expedition in 1948 and 1949. The first of these, PIN 551–2, was named ''Tarbosaurus efremovi'', a new generic name composed of the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words ''τάρβος'' ('), meaning "terror", "alarm", "awe", or "reverence", and ('), meaning "lizard", and the species was named after
Ivan Yefremov Ivan Antonovich (Antipovich) Yefremov, sometimes Efremov (; 23 April 1908 – 5 October 1972) was a Soviet paleontologist, science-fiction author and social thinker. He founded taphonomy, the study of fossilization patterns. Biography He ...
, a Russian paleontologist and
science fiction Science fiction (often shortened to sci-fi or abbreviated SF) is a genre of speculative fiction that deals with imaginative and futuristic concepts. These concepts may include information technology and robotics, biological manipulations, space ...
author. The other two, PIN 553-1 and PIN 552–2, were also named as new species and assigned to the North American genus ''
Gorgosaurus ''Gorgosaurus'' ( ; ) is a genus of tyrannosaurid theropod dinosaur that lived in western North America during the Late Cretaceous Period (Campanian), between about 76.5 and 75 million years ago. Fossil remains have been found in the Ca ...
'' as ''G. lancinator'' and ''G. novojilovi'', respectively. All three of these latter specimens are smaller than the first. A 1965 paper by A. K. Rozhdestvensky recognized all of Maleev's specimens as different growth stages of the same species, which he believed to be distinct from the North American ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
''. He created a new combination, ''Tarbosaurus bataar'', to include all the specimens described in 1955 and newer material. Later authors, including Maleev himself, agreed with Rozhdestvensky's analysis, although some used the name ''Tarbosaurus efremovi'' instead of ''T. bataar''. American paleontologist Ken Carpenter re-examined the material in 1992. He concluded that it belonged to the genus ''Tyrannosaurus'', as originally published by Maleev, and lumped all the specimens into the species ''Tyrannosaurus bataar'' (except the remains that Maleev had named ''Gorgosaurus novojilovi''). Carpenter thought this specimen represented a separate, smaller genus of tyrannosaurid, which he called ''Maleevosaurus novojilovi''. George Olshevsky created the new generic name ''Jenghizkhan'' (after
Genghis Khan Genghis Khan (born Temüjin; August 1227), also known as Chinggis Khan, was the founder and first khan (title), khan of the Mongol Empire. After spending most of his life uniting the Mongols, Mongol tribes, he launched Mongol invasions and ...
) for ''Tyrannosaurus bataar'' in 1995, while also recognizing ''Tarbosaurus efremovi'' and ''Maleevosaurus novojilovi'', for a total of three distinct, contemporaneous genera from the Nemegt Formation. A 1999 study subsequently reclassified ''Maleevosaurus'' as a juvenile ''Tarbosaurus''. All research published since 1999 recognizes only a single species, which is either called ''Tarbosaurus bataar'' or ''Tyrannosaurus bataar''. After the original Soviet-Mongolian expeditions in the 1940s,
Polish Polish may refer to: * Anything from or related to Poland, a country in Europe * Polish language * Polish people, people from Poland or of Polish descent * Polish chicken * Polish brothers (Mark Polish and Michael Polish, born 1970), American twin ...
-Mongolian joint expeditions to the Gobi Desert began in 1963 and continued until 1971, recovering many new fossils, including new specimens of ''Tarbosaurus'' from the Nemegt Formation. Expeditions involving
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
ese and Mongolian scientists between 1993 and 1998, as well as private expeditions hosted by
Canadian Canadians () are people identified with the country of Canada. This connection may be residential, legal, historical or cultural. For most Canadians, many (or all) of these connections exist and are collectively the source of their being ''C ...
paleontologist
Phil Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
around the turn of the 21st century, discovered and collected even more ''Tarbosaurus'' material. More than 30 specimens are known, including more than 15 skulls and several complete postcranial skeletons.


Synonyms

Chinese Chinese may refer to: * Something related to China * Chinese people, people identified with China, through nationality, citizenship, and/or ethnicity **Han Chinese, East Asian ethnic group native to China. **'' Zhonghua minzu'', the supra-ethnic ...
paleontologists discovered a partial skull and skeleton of a small theropod ( IVPP V4878) in the
Xinjiang Xinjiang,; , SASM/GNC romanization, SASM/GNC: Chinese postal romanization, previously romanized as Sinkiang, officially the Xinjiang Uygur Autonomous Region (XUAR), is an Autonomous regions of China, autonomous region of the China, People' ...
Autonomous Region of China in the mid-1960s. In 1977,
Dong Zhiming Dong Zhiming (Chinese language, Chinese: 董枝明, Pinyin: ''Dǒng Zhimíng''; January 1937 – 20 October 2024) was a Chinese vertebrate paleontologist formerly employed at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) ...
described this specimen, which was recovered from the
Subashi Formation The Subashi Formation () is a Late Cretaceous (Campanian to Maastrichtian)Xi et al., 2018 formation from the Xinjiang Autonomous Region of western China.Shanshan County Shanshan County ( zh, s=鄯善县) as the official romanized name, also transliterated from Uyghur as Piqan County (; zh, s=皮羌县, links=no), is a county within the Xinjiang Uyghur Autonomous Region and is under the administrative jurisdict ...
, as a new genus and species, ''Shanshanosaurus huoyanshanensis''.
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
recognized ''Shanshanosaurus'' as a tyrannosaurid in 1988, referring it to the now-defunct genus ''
Aublysodon ''Aublysodon'' (“backwards-flowing tooth") is a dubious genus of carnivorous dinosaurs known only from the Judith River Formation in Montana, which has been dated to the late Campanian age of the late Cretaceous period (about 75 million years ...
''. Dong and Currie later re-examined the specimen and deemed it to be a juvenile of a larger species of tyrannosaurine. These authors refrained from assigning it to any particular genus, but suggested ''Tarbosaurus'' as a possibility. ''Albertosaurus periculosus'', ''Tyrannosaurus luanchuanensis'', ''Tyrannosaurus turpanensis'', and '' Chingkankousaurus fragilis'' were all considered synonyms of ''Tarbosaurus'' in the second edition of the Dinosauria, but ''Chingkankousaurus'' has been assessed as dubious by Brusatte et al. (2013). Named in 1976 by
Sergei Kurzanov Sergei Mikhailovich Kurzanov (Сергей Михайлович Курзанов, born 1947) is a Russian (formerly USSR, Soviet) paleontologist at the Paleontological Institute of the Russian Academy of Sciences. He is known mainly for his work in ...
, ''
Alioramus ''Alioramus'' (; meaning 'different branch') is a genus of tyrannosaurid theropod dinosaurs from the Late Cretaceous period of Asia. It currently contains two species. The type species, ''A. remotus'' is known from a partial skull and three foo ...
'' is another genus of tyrannosaurid from slightly older sediments in Mongolia. Several analyses have concluded ''Alioramus'' was quite closely related to ''Tarbosaurus''. It was described as an adult, but its long, low skull is characteristic of a juvenile tyrannosaurid. This led Currie to speculate that ''Alioramus'' might represent a juvenile ''Tarbosaurus'', but he noted that the much higher tooth count and row of crests on top of the snout suggested otherwise.


Additional specimens

In 1979 Dong Zhiming described several dinosaur remains from the strata of South China, reporting teeth, a dorsal vertebra, and several fragmentary foot bones of a tyrannosaurid. He tentatively referred these to ''Tarbosaurus'' sp. given some similarities in tooth morphology as well as the general geographic proximity to the Nemegt Formation. In 1993 Tomasz Jerzykiewiczz examined the general outcrops from the
Bayan Mandahu Formation The Bayan Mandahu Formation (also known as Wulansuhai Formation or Wuliangsuhai Formation) is a geological unit of "redbeds" located near the village of Bayan Mandahu in Inner Mongolia, China, in the Gobi Desert. It dates from the late Cretaceous ...
of China. In this paper, he reported isolated tyrannosaurid premaxillary and maxillary teeth, which he assigned to ''Tarbosaurus'' without anatomical comparisons. Lev A. Nessov in 1995 reported a partial femur from the
Bostobe Formation The Bostobe Formation () is a geological formation in Qaraghandy & Qyzylorda, Kazakhastan whose strata date back to the Late Cretaceous (Santonian to early Campanian stages, approximately 85 Ma). The sandstones and claystones of the formation ...
of
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
and referred it as ''Tarbosaurus'' sp. without any diagnosis. In 2004, David B. Weishampel and team listed a dubious ''Tarbosaurus''? sp. as a component of the known dinosaur taxa of the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
-aged
Djadokhta Formation The Djadochta formation (sometimes transcribed and also known as Djadokhta, Djadokata, or Dzhadokhtskaya) is a highly fossiliferous geological formation in Central Asia, Gobi Desert, dating from the Late Cretaceous period, about 75 million ...
without further ado. Despite this referral, scattered, and very sparse tyrannosaurid remains are occasionally found in the strata of the Djadokhta Formation of Mongolia and none of these remains have ever been diagnostically assigned to ''Tarbosaurus'' or even ''T. bataar''. In addition, the extreme conditions of the Djadokhta Formation suggest that the remains of tyrannosaurids and other large-bodied dinosaurs from this unit represent passing by taxa foreign to the region. During a large fossil prospection led by the Hayashibara Museum of Natural Sciences-Mongolian Paleontological Center Joint Expedition, a juvenile tyrannosaurid was discovered in 2006 from the highly fossiliferous Bügiin Tsav locality where adult specimens of ''Tarbosaurus'' have been recovered. The specimen was found preserving a partial skeleton with a fairly complete skull. In 2011 this juvenile was formally described and referred to ''Tarbosaurus bataar'', catalogued as MPC-D 107/7 within the collections of the
Mongolian Paleontological Center Mongolian may refer to: * Something of, from, or related to Mongolia, a country in Asia * Mongolian people, or Mongols * Bogd Khanate of Mongolia, the government of Mongolia, 1911–1919 and 1921–1924 * Mongolian language * Mongolian alphabet * ...
.


Poached specimens

''Tarbosaurus'' fossils are only found around the Gobi Desert of Mongolia and China, both of which ban their export, though some specimens have been looted by private collectors. A $1 million smuggling deal was uncovered when suspicions were raised about a catalog put out by Heritage Auctions for an event in New York City on May 20, 2012. By Mongolian law, any specimen found in the
Gobi Desert The Gobi Desert (, , ; ) is a large, cold desert and grassland region in North China and southern Mongolia. It is the sixth-largest desert in the world. The name of the desert comes from the Mongolian word ''gobi'', used to refer to all of th ...
was to rest at an appropriate Mongolian institution and there was little reasonable doubt that the ''Tarbosaurus bataar'' advertised on the catalog was a stolen one. The president of Mongolia and many paleontologists raised objections to the sale, which led to a last-minute investigation that confirmed that it was a specimen that can only be found in the Gobi Desert, rightfully belonging to Mongolia. During the court case ( United States v. One ''Tyrannosaurus Bataar'' Skeleton), Eric Prokopi, the smuggler, pleaded guilty to illegal smuggling and the dinosaur was returned to Mongolia in 2013, where it is temporarily displayed on Sukhbaatar Square, the center of the city of Ulaanbaatar. Prokopi had sold the dinosaur with a partner and fellow commercial hunter in England named Christopher Moore. The case led to the repatriation of dozens of Mongolian dinosaurs, including several skeletons of ''Tarbosaurus bataar''.


Skin impressions and footprints

Skin impressions were recovered from a large skeleton at the Bugiin Tsav locality that was subsequently destroyed by poachers. These impressions show non-overlapping scales with an average diameter of and pertain to the thoracic region of the individual, although the exact position can no longer be assessed due to the destruction of the skeleton.
Phil Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
and colleagues (2003) described two footprints from the Nemegt locality that probably pertain to ''Tarbosaurus''. These tracks represent natural casts, which means that only the sandy infill of the tracks, not the tracks themselves, are preserved. The better-preserved track features skin impressions over large areas on and behind the toe impressions that are similar to those discovered in Bugiin Tsav. It also features vertical parallel slide marks that were left by scales when the foot was pushed into the ground. The track measures in length, thus representing a large individual. The second track, although even larger, was affected by
erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...
and does not show any detail. In 1997, Ken Carpenter reported a damaged ''Tarbosaurus'' skull with impressions of a dewlap or throat pouch beneath the lower jaws, based on a personal communication from Konstantin Mikhailov. Carpenter speculated that the pouch may have been used for display, possibly being brightly colored and inflatable like a frigatebird's. In a 2019 communication to Mickey Mortimer, Mikhailov confirmed that this specimen had not been collected because it was on a heavy stone slab. He revealed that it had been discovered by
Sergei Kurzanov Sergei Mikhailovich Kurzanov (Сергей Михайлович Курзанов, born 1947) is a Russian (formerly USSR, Soviet) paleontologist at the Paleontological Institute of the Russian Academy of Sciences. He is known mainly for his work in ...
and that it was Kurzanov himself who had originally interpreted the impressions as a throat structure. This specimen may be the same as one that was purportedly destroyed by poachers in 1992.


Description

Although slightly smaller than ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
'', ''Tarbosaurus'' was one of the largest tyrannosaurines, with the type specimen PIN 551–1 measuring approximately long, tall at the hips, and weighing up to . Other adult specimens were a bit smaller, as MPC-D 107/2, ZPAL.MgD-I/4 and PIN 552-1 likely weighed around to . The largest known ''Tarbosaurus'' skull is about long, which is larger than that of all other tyrannosaurids, aside from ''Tyrannosaurus''.


Skull

The skull was tall, like that of ''Tyrannosaurus'', but not as wide, especially towards the rear. The unexpanded rear of the skull meant that ''Tarbosaurus''‘s eyes did not face directly forwards, suggesting that it lacked the
binocular vision Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
of ''Tyrannosaurus''. Large
fenestra A fenestra (fenestration; : fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biology, biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomy, ...
e in the skull reduced its overall weight and served as attachment points for muscles. Between 58 and 64 teeth lined its jaws. This tooth count is slightly more than that of ''Tyrannosaurus'', but fewer than in smaller tyrannosaurids, like ''
Gorgosaurus ''Gorgosaurus'' ( ; ) is a genus of tyrannosaurid theropod dinosaur that lived in western North America during the Late Cretaceous Period (Campanian), between about 76.5 and 75 million years ago. Fossil remains have been found in the Ca ...
'' and ''Alioramus''. Most of its teeth were ovalur in
cross section Cross section may refer to: * Cross section (geometry) ** Cross-sectional views in architecture and engineering 3D *Cross section (geology) * Cross section (electronics) * Radar cross section, measure of detectability * Cross section (physics) **A ...
, although the teeth of the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
at the tip of the upper jaw had a ''D''-shaped cross section. However, this
heterodonty In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals where ...
is characteristic of the family. The longest teeth were in the
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
(upper jaw bone), with crowns up to long. In the
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
, a ridge on the outer surface of the
angular bone The angular is a large bone in the lower jaw (mandible) of amphibians and reptiles (birds included), which is connected to all other lower jaw bones: the dentary (which is the entire lower jaw in mammals), the splenial, the suprangular, and the ...
articulated with the rear of the
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
bone, creating a locking mechanism unique to ''Tarbosaurus'' and ''Alioramus''. Other tyrannosaurids lacked this ridge and had more flexibility in the lower jaw.


Postcranial skeleton

Tyrannosaurids varied little in overall body form and ''Tarbosaurus'' was no exception. The head was supported by an ''S''-shaped neck, while the rest of the
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
, including the tail, was held horizontally. ''Tarbosaurus'' had tiny arms, proportionably to body size the smallest of all members in the family. The hands had two clawed digits each, with an additional unclawed third metacarpal found in some specimens, similar to those of closely related genera. Holtz has suggested that ''Tarbosaurus'' also has a theropod reduction of fingers IV-I "developed further" than in other tyrannosaurids,Carpenter K, Tanke D.H. & Skrepnick M.W. (2001), ''Mesozoic Vertebrate Life'' (
Indiana University Press Indiana University Press, also known as IU Press, is an academic publisher founded in 1950 at Indiana University that specializes in the humanities and social sciences. Its headquarters are located in Bloomington, Indiana. IU Press publishes ...
, ), p. 71.
as the second metacarpal in the ''Tarbosaurus'' specimens he studied is less than twice the length of the first metacarpal. Other tyrannosaurids have a second metacarpal about twice the length of the first metacarpal. Also, the third metacarpal in ''Tarbosaurus'' is proportionally shorter than in other tyrannosaurids. In other tyrannosaurids, like ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of large tyrannosaurid theropod dinosaur that lived in northwestern North America during the early to middle Maastrichtian age of the Late Cretaceous period, about 71 million yea ...
'' and ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 77 and 74.4 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three named ...
'', the third metacarpal is often longer than the first metacarpal, while in the ''Tarbosaurus'' specimens studied by Holtz, the third metacarpal is shorter than the first. In contrast to the arms, the three-toed legs were long, thick, and muscular to support the body in a bipedal posture. The long, heavy tail served as a
counterweight A counterweight is a weight (object), weight that, by applying an opposite force, provides balance and stability of a machine, mechanical system. The purpose of a counterweight is to make lifting the load faster and more efficient, which saves e ...
to the head and torso, while also placing the
center of gravity In physics, the center of mass of a distribution of mass in space (sometimes referred to as the barycenter or balance point) is the unique point at any given time where the weighted relative position of the distributed mass sums to zero. For ...
directly over the hips.


Classification

''Tarbosaurus'' is classified as a
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
in the subfamily Tyrannosaurinae of the family
Tyrannosauridae Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
. Other members include ''Tyrannosaurus'' and the earlier ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 77 and 74.4 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three named ...
'', both from North America, and possibly the Mongolian genus ''Alioramus''. Animals in this subfamily are more closely related to ''Tyrannosaurus'' than to ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of large tyrannosaurid theropod dinosaur that lived in northwestern North America during the early to middle Maastrichtian age of the Late Cretaceous period, about 71 million yea ...
'' and are known for their robust build with proportionally larger skulls and longer
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
s than in the other subfamily, Albertosaurinae. ''Tarbosaurus bataar'' was originally described as a species of ''Tyrannosaurus'', an arrangement that has been supported by some more recent studies. Others prefer to keep the genera separate, while still recognizing them as
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
. A 2003
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis based on skull features instead identified ''Alioramus'' as the closest known relative of ''Tarbosaurus'', as the two genera share skull characteristics that are related to stress distribution and are not found in other tyrannosaurines. If proven, this relationship would argue against ''Tarbosaurus'' being a synonym of ''Tyrannosaurus'' and would suggest that separate tyrannosaurine lineages
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
in Asia and North America. The two known specimens of ''Alioramus'', which show juvenile characteristics, are not likely juvenile individuals of ''Tarbosaurus'' because of their much higher tooth count (76 to 78 teeth) and their unique row of bony bumps along the top of their snouts. The discovery of ''Lythronax argestes'', a much earlier tyrannosaurine, further reveals the close relationship between ''Tyrannosaurus'' and ''Tarbosaurus''. It was also discovered that ''Lythronax'' is a sister taxon to a clade consisting of Campanian genus ''Zhuchengtyrannus'', and the Maastrichtian genera ''Tyrannosaurus'' and ''Tarbosaurus''. Further studies of ''Lythronax'' also suggest that the Asian tyrannosauroids were part of one evolutionary radiation. Below is the cladogram of Tyrannosaurinae based on the phylogenetic analysis conducted by Voris and team in 2020.


Paleobiology


Ontogeny

Most specimens of ''Tarbosaurus'' represent adult or subadult individuals, while juveniles remain very rare. Nevertheless, the 2006 discovery of a juvenile individual (MPC-D 107/7) with a complete, long skull was reported and described in 2011, providing information on the life history of this dinosaur. This individual was probably 2 to 3 years old at the time of death. Compared to adult skulls, the juvenile skull was weakly constructed and the teeth were thin, indicating different food preferences in juveniles and adults that reduced competition between different age groups. Examination of the sclerotic rings in this juvenile ''Tarbosaurus'' suggests they may also have been crepuscular or nocturnal hunters. Whether or not the adult ''Tarbosaurus'' were also nocturnal is currently unknown due to the lack of fossil evidence to suggest so. Ontogenetic changes identified in the frontal bones suggests that the allometric growth of ''Tarbosaurus'' was similar to that of North American tyrannosaurids.


Senses

A ''Tarbosaurus'' skull found in 1948 by Soviet and Mongolian scientists (PIN 553–1, originally called ''Gorgosaurus lancinator'') included the skull cavity that held the brain. Making a plaster Casting, cast, called an ''endocranial cast, endocast'', of the inside of this cavity allowed Maleev to make preliminary observations about the shape of a ''Tarbosaurus'' brain. A newer polyurethane rubber cast allowed a more detailed study of ''Tarbosaurus'' brain structure and function. The endocranial structure of ''Tarbosaurus'' was similar to that of ''Tyrannosaurus'', differing only in the positions of some cranial nerve roots, including the trigeminal nerve, trigeminal and accessory nerves. Tyrannosaurid brains were more similar to those of crocodilians and other nonavian reptiles than they were to birds. The total brain volume for a long ''Tarbosaurus'' is estimated at only . The large size of the olfactory bulbs, as well as the terminal nerve, terminal and olfactory nerves, suggest that ''Tarbosaurus'' had a highly keen sense of smell, as was also the case with ''Tyrannosaurus''. The Olfactory bulb#Accessory olfactory bulb, vomeronasal bulb is large and differentiated from the olfactory bulb, which was initially suggested as being indicative of a well-developed Vomeronasal organ, Jacobsen's organ, which was used to detect pheromones. This may imply that ''Tarbosaurus'' had complex mating behavior. However, the identification of the vomeronasal bulb has been challenged by other researchers because they are not present in any living archosaurs. The auditory nerve was also large, suggesting good Hearing (sense), hearing, which may have been useful for auditory animal communication, communication and spatial awareness. The nerve had a well-developed Vestibular system, vestibular component as well, which implies a good sense of balance and coordination. In contrast, the nerves and brain structures associated with eyesight were smaller and undeveloped. The midbrain tectum, responsible for visual processing in reptiles, was very small in ''Tarbosaurus'', as were the optic nerve and the oculomotor nerve, which controls eye movement. Unlike ''Tyrannosaurus'', which had forward-facing eyes that provided accurate binocular vision, ''Tarbosaurus'' had a narrower skull more typical of other tyrannosaurids in which the eyes faced primarily sideways. All of this suggests that ''Tarbosaurus'' relied more on its senses of smell and hearing than on its eyesight. It has been suggested that the lack of binocular vision in Asian tyrannosaurs, like ''Tarbosaurus'', might have been correlated with a greater amount of scavenging resources provided by sauropod carcasses, which might have afforded them a less active predatory lifestyle when compared to the North American forms, meaning they would need less predatory adaptations. However, this is contradicted by numerous lines of evidence indicating ''Tarbosaurus'' was actively preying on hadrosaurs, titanosaur sauropods, and other large bodied herbivores in its ecosystem.


Skull mechanics

The skull of ''Tarbosaurus'' was completely described for the first time in 2003. Scientists noted key differences between ''Tarbosaurus'' and the North American tyrannosaurids. Many of these differences are related to the handling of stress by the skull bones during a bite. When the upper jaw bit down on an object, force was transmitted up through the maxilla, the primary tooth-bearing bone of the upper jaw, into surrounding skull bones. In North American tyrannosaurids, this force went from the maxilla into the fused nasal bones on top of the snout, which were firmly connected in the rear to the lacrimal bones by bony struts. These struts locked the two bones together, suggesting that force was then transmitted from the nasals to the lacrimals. ''Tarbosaurus'' lacked these bony struts and the connection between the nasals and lacrimals was weak. Instead, a backwards projection of the maxilla was massively developed in ''Tarbosaurus'' and fit inside a sheath formed from the lacrimal. This projection was a thin, bony plate in North American tyrannosaurids. The large backwards projection suggests that force was transmitted more directly from the maxilla to the lacrimal in ''Tarbosaurus''. The lacrimal was also more firmly anchored to the frontal bone, frontal and prefrontal bones in ''Tarbosaurus''. The well-developed connections between the maxilla, lacrimal, frontal, and prefrontal would have made its entire upper jaw much more rigid. Another major difference between ''Tarbosaurus'' and its North American relatives was its more rigid mandible. While many theropods, including North American tyrannosaurids, had some degree of flexibility between the bones in the rear of the mandible and the dentary in the front, ''Tarbosaurus'' had a locking mechanism formed from a ridge on the surface of the angular, which articulated with a square process on the rear of the dentary. Some scientists have hypothesis, hypothesized that the more rigid skull of ''Tarbosaurus'' was an adaptation to hunting the massive titanosaurid sauropods found in the Nemegt Formation, which did not exist in most of North America during the Late Cretaceous. The differences in skull mechanics also affect tyrannosaurid phylogeny. ''Tarbosaurus''-like articulations between the skull bones are also seen in ''Alioramus'' from Mongolia, suggesting that it is the closest relative of ''Tarbosaurus''. Similarities between ''Tarbosaurus'' and ''Tyrannosaurus'' might be related to their large size, independently developed through convergent evolution.


Bite force and feeding

In 2001, Bruce Rothschild and others published a study examining evidence for stress fractures and Avulsion fracture, tendon avulsions in Theropoda, theropod dinosaurs and the implications for their behavior. Since stress fractures are caused by repeated trauma rather than singular events, they are more likely to be caused by regular behavior than other types of injuries. None of the eighteen ''Tarbosaurus'' foot bones examined in the study were found to have a stress fracture, but one of the ten examined hand bones was found to have one. Stress fractures in the hands have special behavioral significance compared to those found in the feet, since stress fractures there can be obtained while running or during migration. Hand injuries, by contrast, are more likely to be obtained while in contact with struggling prey. The presence of stress fractures and tendon avulsions, in general, provide evidence for a "very active" predation-based diet instead of obligate scavenging.Rothschild, B., Tanke, D. H., and Ford, T. L., 2001, Theropod stress fractures and tendon avulsions as a clue to activity: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 331-336. As for its bite force, it was revealed in 2005 that ''Tarbosaurus'' had a bite force of around 8,000 to 10,000 pounds per square inch of force, meaning that it could crush bones just like its North American relative, ''Tyrannosaurus''. David W. E. Hone and Mahito Watabe in 2011 reported the left humerus of a nearly complete ''
Saurolophus ''Saurolophus'' (; meaning "lizard crest") is a genus of large hadrosaurid dinosaur from the Late Cretaceous period of Asia and North America, that lived in what is now the Horseshoe Canyon and Nemegt formations about 70 million to 66 million ...
'' skeleton (MPC-D 100/764) from the Bügiin Tsav locality of the Nemegt Formation, which was Paleopathology, heavily damaged from bite marks attributed to ''Tarbosaurus''. As suggested by the lack of damage to the rest of the skeleton (such as large wounds in skeletal remains indicative of predation), this tyrannosaurid was likely scavenging an already dead ''Saurolophus''. It is unlikely that a large-bodied predator, such as ''Tarbosaurus'', would have left sparse feeding traces on a single humerus when having an entire carcass to feed on. The humerus shows three distinctive feeding methods, interpreted as punctures, drag marks, and bite−and−drag marks. Hone and Watabe noted that bite marks were mostly located at the deltopectoral crest, suggesting that this ''Tarbosaurus'' was actively selecting which biting style to employ so it could scavenge the bone. In 2012, bite marks on two fragmentary Gastralium, gastralia of the holotype specimen of the large ornithomimosaur ''Deinocheirus, Deinocheirus mirificus'' were reported. The size and shape of the bite marks match the teeth of ''Tarbosaurus'', the largest known predator from the Nemegt Formation. Various types of feeding traces were identified. These include punctures, gouges, striae, fragmentary teeth, and combinations of the above marks. The bite marks probably represent feeding behavior instead of aggression between the species and the fact that bite marks were not found elsewhere on the body indicates the predator focused on internal organs. ''Tarbosaurus'' bite marks have also been identified on hadrosaur and sauropod fossils, but theropod bite marks on bones of other theropods are very rare in the fossil record. A 2020 study involving stable isotopes found that ''Tarbosaurus'' primarily hunted large dinosaurs in its environment, most notably titanosaurs and hadrosaurs.


Paleoenvironment


Nemegt Formation

The vast majority of known ''Tarbosaurus'' fossils were recovered from the
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 ...
in the
Gobi Desert The Gobi Desert (, , ; ) is a large, cold desert and grassland region in North China and southern Mongolia. It is the sixth-largest desert in the world. The name of the desert comes from the Mongolian word ''gobi'', used to refer to all of th ...
of southern Mongolia. This
geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
has never been radiometric dating, dated radiometrically, but the fauna present in the fossil record indicate it was probably deposited during the early
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
stage at the near end of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
, about 70 million years ago.Sulliban, R.M. (2006). "A taxonomic review of the Pachycephalosauridae (Dinosauria: Ornithischia)." Pp. 347-366 in Lucas, S.G. and Sullivan, R.M. (eds.), ''Late Cretaceous vertebrates from the Western Interior'.' New Mexico Museum of Natural History and Science Bulletin 3. ''Tarbosaurus'' is found chiefly in the Nemegt Formation, whose sediments preserve large river channels and soil deposits that indicate a far more humid climate than those suggested by the underlying Barun Goyot Formation, Barun Goyot and Djadochta Formations. However, Caliche (mineral), caliche deposits indicate at least periodic droughts. Sediment was deposited in the channels and floodplains of large rivers. The rock facies of this formation suggest the presence of mudflats and shallow lakes. Sediments also indicate that there existed a rich habitat, offering diverse food in abundant amounts that could sustain massive Cretaceous dinosaurs.Novacek, M. (1996). Dinosaurs of the Flaming Cliffs. Bantam Doubleday Dell Publishing Group Inc. New York, New York. Occasional mollusk fossils are found, as well as a variety of other aquatic animals, such as fish and turtles. Crocodilians included several species of ''Paralligator'', a genus with teeth adapted for crushing shells. Mammal fossils are exceedingly rare in the Nemegt Formation, but many birds have been found, including the enantiornithine ''Gurilynia'' and the Hesperornithes, hesperornithiform ''Judinornis'', as well as ''Teviornis'', an early representative of the still-existing Anseriformes. Scientists have described many dinosaurs from the Nemegt Formation, including the ankylosaurids ''
Tarchia ''Tarchia'' (meaning "brainy one") is a genus of herbivorous ankylosauridae, ankylosaurid dinosaur from the late Cretaceous of Mongolia. Discovery and naming In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. ...
'' and ''
Saichania ''Saichania'' (Mongolian meaning "beautiful one") is a genus of herbivorous ankylosaurid dinosaur from the Late Cretaceous period of Mongolia. The first fossils of ''Saichania'' were found in the early 1970s in Mongolia. In 1977 the type speci ...
'' and the pachycephalosaur ''Prenocephale''. By far the largest predator known from the formation, adult ''Tarbosaurus'' most likely preyed upon large hadrosaurs, such as ''
Saurolophus ''Saurolophus'' (; meaning "lizard crest") is a genus of large hadrosaurid dinosaur from the Late Cretaceous period of Asia and North America, that lived in what is now the Horseshoe Canyon and Nemegt formations about 70 million to 66 million ...
'' and ''
Barsboldia ''Barsboldia'' (meaning "of Rinchen Barsbold, Barsbold", a well-known Mongolian people, Mongolian paleontologist) is a genus of large hadrosaurid dinosaur from the Maastrichtian, early Maastrichtian Nemegt Formation of Ömnögovi Province, Ömno ...
'', or sauropods, such as ''
Nemegtosaurus ''Nemegtosaurus'' (meaning 'Reptile from the Nemegt') was a sauropod dinosaur from the Late Cretaceous Period of what is now Mongolia. Measuring an estimated long and weighing , it was named after the Nemegt Basin in the Gobi Desert, where the ...
'' and ''
Opisthocoelicaudia ''Opisthocoelicaudia'' is a genus of sauropod dinosaur of the Late Cretaceous Period discovered in the Gobi Desert of Mongolia. The type species is ''Opisthocoelicaudia skarzynskii''. A well-preserved skeleton lacking only the head and neck wa ...
''. Adults would have received little competition from small theropods, such as the small
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
''
Alioramus ''Alioramus'' (; meaning 'different branch') is a genus of tyrannosaurid theropod dinosaurs from the Late Cretaceous period of Asia. It currently contains two species. The type species, ''A. remotus'' is known from a partial skull and three foo ...
'', troodontids (''Borogovia'', ''Tochisaurus'', ''Zanabazar (dinosaur), Zanabazar''), oviraptorosaurs (''Elmisaurus'', ''Nemegtomaia'', ''Rinchenia'') or ''Bagaraatan'', sometimes considered a basal (phylogenetics), basal tyrannosauroid. Other theropods, like the gigantic ''Therizinosaurus'', might have been herbivore, herbivorous and ornithomimosaurs, such as ''Anserimimus'', ''Gallimimus'', and gigantic ''Deinocheirus'' might have been omnivores that only took small prey and were therefore no competition for ''Tarbosaurus''. However, as in other large tyrannosaurids, as well as modern Komodo dragons, juveniles and subadult ''Tarbosaurus'' would have filled niches between the massive adults and these smaller theropods.


Subashi Formation

The
Subashi Formation The Subashi Formation () is a Late Cretaceous (Campanian to Maastrichtian)Xi et al., 2018 formation from the Xinjiang Autonomous Region of western China.Skeletal image
of ''Shanshanosaurus'' a


Comparison between ''Tarbosaurus'' skulls from specimens of different age group and size
Readable material

at The Theropod Database.

by George Olshevsky (1995). * ''Scienceblogs:'
Juvenile, 5 years old ''Tarbosaurus'' specimen found in Mongolia 2006
{{Taxonbar, from=Q2491 Tyrannosauridae Dinosaur genera Maastrichtian dinosaurs Nemegt Formation Taxa named by Evgeny Maleev Fossil taxa described in 1955 Dinosaurs of Mongolia