Alioramus Altai
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''Alioramus'' (; meaning 'different branch') 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
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 ...
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 ...
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 ...
s from 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 ...
period of
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 ...
. It currently contains two species. The
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 ...
, ''A. remotus'' is known from a partial
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
and three foot bones recovered from the
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 ...
n
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 ...
, which was deposited 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 ...
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 ...
. These remains were named and described by
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 ...
paleontologist
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 ...
in 1976. A second species, ''A. altai'', known from a much more complete skeleton also from the Nemegt Formation, was named and described by Stephen L. Brusatte and colleagues in 2009. Its relationships to other tyrannosaurid genera were at first unclear, with some evidence supporting a hypothesis that ''Alioramus'' was closely related to the contemporary species ''
Tarbosaurus bataar ''Tarbosaurus'' ( ; meaning "alarming lizard") is a genus of large tyrannosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 70 million years ago (Maastrichtian age). It contains the single type species: ''Tarbosaurus b ...
''. However, 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 ...
'' indicates that it belongs to a distinct branch of tyrannosaurs, namely the 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 ...
. ''Alioramus'' were
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 like all known theropods, and their sharp teeth indicate that they were
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s. Known specimens were smaller than other tyrannosaurids like ''Tarbosaurus bataar'' and ''
Tyrannosaurus rex ''Tyrannosaurus'' () is a genus of large theropoda, 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 live ...
'', but their adult size is difficult to estimate since both ''Alioramus'' species are known only from juvenile or sub-adult remains. The genus ''Alioramus'' is characterized by a row of five bony crests along the top of the snout, a greater number of teeth than any other genus of tyrannosaurid, and a lower skull than most other tyrannosaurids.


History of discovery

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 ...
3141/1) of ''Alioramus'' is a partial skull associated with three
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
s. 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 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 early 1970s found these remains at a locality known as Nogon-Tsav in the Mongolian province of
Bayankhongor Bayankhongor () is the capital of the Bayankhongor Province (aimag) in Mongolia. The administration of the Bayankhongor Sum (district) is also located in the same place. The city is at an elevation of 1859 m above sea level, and has a populatio ...
,
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 ...
. ''Alioramus'' was named and described by
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
n paleontologist Sergei Kurzanov in 1976. Its crests and low skull profile looked so different from other tyrannosaurids that Kurzanov believed his find was far removed from other members of the family. Accordingly, he gave it the generic name ''Alioramus'', derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''alius'' ('other') and ''ramus'' ('branch'), and the specific name ''A. remotus'', which means 'removed' in Latin. A second species, ''A. altai'', was discovered back in 2009 at the Tsagan Khushu locality also from the Nemegt Formation. However, several faunal differences may suggest that the respective locations of ''A. remotus'' and ''A. altai'' differ in age. The holotype IGM 100/1844 is a partial skeleton that includes a very complete skull—more so than ''A. remotus''—with partial
vertebrae 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 spinal ...
,
pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...
and hindlimbs. The name for this species, ''altai'', is in reference to the
Altai Mountains The Altai Mountains (), also spelled Altay Mountains, are a mountain range in Central Asia, Central and East Asia, where Russia, China, Mongolia, and Kazakhstan converge, and where the rivers Irtysh and Ob River, Ob have their headwaters. The ...
.


Description

''Alioramus remotus'' was estimated at in length when originally described by Sergei Kurzanov in 1976. In 1988
Paul Paul may refer to: People * Paul (given name), a given name, including a list of people * Paul (surname), a list of people * Paul the Apostle, an apostle who wrote many of the books of the New Testament * Ray Hildebrand, half of the singing duo ...
gave a similar length of and a weight of . In 2016 Molina-Pérez and Larramendi estimated ''A. remotus'' at and , and ''A. altai'' at and . Kurzanov, however, did not correct for lengthening of the skull by deformation during
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 ...
ization, which may indicate a shorter overall body length for this individual. If this specimen is a juvenile, then adult ''Alioramus'' would have reached greater lengths, but no confirmed adult specimens are known.


Skull

The skull of ''A. remotus'' was approximately long. In general, it is long and low, a shape typical of more basal
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 supercontinen ...
s and juveniles of larger tyrannosaurids. 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 ...
ry bones at the tip of the snout in ''Alioramus remotus'' have not been found, but are taller than wide in all tyrannosauroids for which they are known. The
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
s are fused and ornamented with a row of five irregular bony crests that protrude upwards from the midline, where the nasal bones are sutured together. These crests all measure more than tall. At the back of the skull there is a protrusion, called the ''nuchal crest'', arising from the fused
parietal bone The parietal bones ( ) are two bones in the skull which, when joined at a fibrous joint known as a cranial suture, form the sides and roof of the neurocranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four bord ...
s, a feature shared with all tyrannosaurids. In ''Alioramus'', the nuchal crest is greatly thickened, similarly to ''Tarbosaurus'' and ''Tyrannosaurus''. Like the rest of the skull, 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 ...
of ''Alioramus'' was long and slender, another possible juvenile characteristic. As in ''Tarbosaurus'', 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 ...
of the lower jaw articulated with the rear of the
dentary bone 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 ...
, locking the two bones together and removing much of the flexibility seen in other tyrannosaurids. Other tyrannosaurids had four premaxillary teeth, ''D''-shaped 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 ...
, on each side. Including 16 or 17 in each
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 ...
, and 18 in each
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 ...
, ''Alioramus'' had 76 or 78 teeth, more than any other tyrannosaurid. The braincase of ''A. altai'' was intermediate between the basal theropod and
avialan Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative defi ...
conditions.


Postcranial skeleton

The rest of the skeleton of ''Alioramus remotus'' is completely unknown except for three
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
s (bones of the upper foot), but the discovery of ''A. altai'', which is known from substantially more complete remains, has shed light on the anatomy of the genus.


Classification

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 ...
s have long
classified Classified may refer to: General *Classified information, material that a government body deems to be sensitive *Classified advertising or "classifieds" Music *Classified (rapper) (born 1977), Canadian rapper * The Classified, a 1980s American ro ...
''Alioramus'' within the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Tyrannosauroidea, but because its remains were for many years poorly known, a more precise classification had remained elusive until the discovery of ''A. altai''. A
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 published in 2003 found ''Alioramus'' could be further classified into the family Tyrannosauridae and the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Tyrannosaurinae, alongside ''Tyrannosaurus'', ''Tarbosaurus'' 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 ...
''. A 2004 study supported this result but suggested it was equally probable that ''Alioramus'' belonged outside the family Tyrannosauridae entirely, with its supposed juvenile characters actually reflecting a more basal position within Tyrannosauroidea. Another study omitted ''Alioramus'' altogether due to the only specimen's fragmentary nature. The description of ''A. altai'' in 2009 confirmed the placement of the genus within the Tyrannosaurinae. ''Tarbosaurus'' and ''Alioramus'' shared several skull features, including a locking mechanism in the lower jaw between the dentary and angular bones, and both lacked the prong of the nasal bones which connected to the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
s in all other tyrannosaurids except adult ''Daspletosaurus''. The two genera may be closely related, representing an Asian branch of the Tyrannosauridae. Some specimens of ''Tarbosaurus'' have a row of bumps on the nasal bones like those of ''Alioramus'', although much lower. The long and low shape of the only known ''Alioramus remotus'' skull indicated that it was immature when it died and might even have been a juvenile ''Tarbosaurus'', which lived in the same time and place. The more prominent nasal crests and much higher tooth count of ''Alioramus'', however, suggested it was a separate taxon, even if it is known only from juvenile remains, confirmed by the discovery of ''A. altai''. Specimens identified as immature ''Tarbosaurus'' have the same tooth count as adults. The description of ''Qianzhousaurus'' in 2014 erected a new branch of the tyrannosaur family named Alioramini; consisting of the long-snouted ''Q. sinensis'' and the two known species of ''Alioramus''. This clade had an uncertain placement relative to other members of the tyrannosaur branch in the initial analysis that discovered it. The primary phylogenetic analysis found Alioramini to be closer to ''Tyrannosaurus'' than to ''Albertosaurus'', and therefore a member of the group Tyrannosaurinae. However, a second analysis in the same paper found it to be located outside of the clade including Albertosaurinae and Tyrannosaurinae, and therefore the sister group of Tyrannosauridae. Below is the first analysis found by the authors:


Paleobiology


Feeding

Brusatte and colleagues in 2009 indicated that ''Alioramus'' lacks many of the robust and brute skull traits (such as a deep maxilla, robust lower jaws, or peg-like teeth) that are necessary to employ a "puncture-pull" feeding characteristic of large tyrannosaurids. They suggested that ''Alioramus'' may have exploited a different feeding style, such as focusing small-sized prey. This would also suggest that both ''Alioramus'' and ''Tarbosaurus''—whose remains have also been collected at the Tsagan Khushu locality, making them
sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
—used different feeding strategies, avoiding competition. Foster with team in 2022 hypothesized that due to their slim and gracile build, Alioramin genera may have been hunters of small, particularly fast and nimble prey, which would have allowed alioramins to avoid competition with larger tyrannosaurs that specialized in killing larger animals. The long and delicate snouts of alioramins like ''Alioramus'' and ''Qianzhousaurus'' may have also prevented them from killing the same prey species that juvenile and adult tyrannosaurids of tyrannosaurids like ''Tarbosaurus'' hunted, though these larger tyrannosaurs themselves may have hunted alioramins as prey on occasion. Alioramins may also have had a different feeding strategy than other tyrannosaurids, as their jaws seem to have been weaker than those of the larger genera, and even juveniles of larger species have proportionately higher bite forces than alioramins of equivalent size. Furthermore, Alioramins seemingly remained confined to Asia, suggesting some factor prevented them from colonizing the better-sampled fossil deposits from North America. Why this may be remains a mystery until more evidence is discovered. Examinations of the skulls of various genera of tyrannosauroids suggest that ''Alioramus'' experienced lower stresses to its skull when feeding. Additionally, the same study suggests it and its relative ''Qianzhousaurus'' did not use the "puncture-and-pull" feeding method used by larger genera such as ''Tarbosaurus'' or ''Tyrannosaurus.''


Ontogeny

Histological analyses performed on the holotype of ''A. altai'' (IGM 100/1844) by Brusatte and colleagues in 2009 determined that this individual had an internal bone structure corresponding to a nine year-old and actively growing ''Tyrannosaurus''. The team however, noted that in terms of body size this individual is closer to a seven/eight year-old ''Albertosaurus'' or ''Gorgosaurus'', and a five/six year-old ''Daspletosaurus'' or ''Tyrannosaurus'', which may suggest that ''Alioramus'' attained a comparably smaller adult size. Lastly, Brusatte and team argued against the skull shape and cranial ornamentation of ''Alioramus'' being juvenile traits, given that: IGM 100/1844 is smaller and slender than comparably aged ''Tyrannosaurus'' and has a longer snout than any known juvenile of large tyrannosaurids (''Albertosaurus'' or ''Tarbosaurus''); and several well-documented
ontogenic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
(growth) series of other dinosaurs evidence that ornamentation increases throughout growth. The latter may suggest that adult ''Alioramus'' possessed a rather elaborate cranial ornamentation. Examinations 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 its comparisons with both species of ''Alioramus'' published in 2022 suggests that both Alioramus species are known from juvenile specimens in different growth stages, and that ''Qianzhousaurus'' represents an adult example of the alioramini. The examinations also suggest that the variation seen between the various species is consistent with the growth trends seen in other tyrannosaurid genera, though specimens that could constitute a full growth series from infant to adult for each species have not been recovered for any of these tyrannosaurs. One part of the growth series across all specimens in this study was discovered to remain unique to alioramin tyrannosaurs; the rugose process of the jugal starts small and conical in early life, but becomes massive and indistinct as the animals grow. This same study also suggests Alioramins did not undergo a secondary metamorphosis from slender juveniles to robust adults like other tyrannosaurs such as ''Tarbosaurus'' and ''Tyrannosaurus'' did, but maintained a unique physiology better suited to pursuit of fast, small prey.


Paleoenvironment

The Beds of Nogon-Tsav are considered to be the same age as 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 ...
. 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 dated radiometrically, but the
fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
present in the fossil record indicate it was probably deposited during the
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 Stage, stages, or staging may refer to: Arts and media Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly Brit ...
, at the 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 ...
. The Maastrichtian stage in Mongolia, as preserved in the Nemegt Formation and at Nogon-Tsav, was characterized by a wetter and more humid climate compared with the semi-arid environment preserved in the earlier, underlying Barun Goyot and
Djadochta 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 ...
s. Nemegt sediments preserve
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 ...
s, large
river A river is a natural stream of fresh water that flows on land or inside Subterranean river, caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of ...
channels and soil deposits, but
caliche Caliche () is a soil accumulation of soluble calcium carbonate at depth, where it precipitates and binds other materials—such as gravel, sand, clay, and silt. It occurs worldwide, in aridisol and mollisol soil orders—generally in arid or se ...
deposits indicate periodic droughts. This environment supported a more diverse and generally larger dinosaur fauna than in earlier times. Kurzanov reported that other theropods, including ''Tarbosaurus'', ornithomimosaurs and
therizinosaur Therizinosaurs (; once called segnosaurs) are an extinct group of large herbivorous theropod dinosaurs whose fossils have been mainly discovered from Cretaceous deposits in Asia and North America. Potential fragmentary remains have also been foun ...
s were discovered at the same locality, but these remains have never been reported in detail. If the Nogon Tsav
fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
was similar to that of the Nemegt Formation,
troodontid Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
theropods, as well as
pachycephalosaur Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycepha ...
s,
ankylosaurid 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 ...
s and
hadrosaur 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 ...
s would also have been present.
Titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...
n
sauropod 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 ...
s were also potential prey for predators in the Nemegt.


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

* *
3D holotype braincase of ''A. altai''
at
DigiMorph Digital Morphology (DigiMorph), part of the National Science Foundation Digital Libraries Initiative, creates and shares 2D and 3D visualizations of the internal and external structure of living and extinct vertebrates, and a growing number of 'inve ...
{{Taxonbar, from=Q131174 Tyrannosauridae Dinosaur genera Maastrichtian dinosaurs Nemegt Formation Dinosaurs of Mongolia Dinosaurs of China Fossil taxa described in 1976 Taxa named by Sergei Kurzanov